Marine Decapod Crustacea: A Guide to Families and Genera of the World 1032138025, 9781032138022

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Marine Decapod Crustacea: A Guide to Families and Genera of the World
 1032138025, 9781032138022

Table of contents :
Cover
Title Page
Copyright
Contents
Acknowledgements
1. Introduction
What is a crustacean?
Scope of this work
History and resources
Structure of this book, and some advice!
2. Systematics
Evolution of the Crustacea
Classification of the Crustacea
Classification of the Decapoda
Key to suborders and infraorders of Decapoda and shrimp-like marine crustaceans
3. Dendrobranchiata – prawns and shrimps
Penaeoidea
Aristeidae
Benthesicymidae
Penaeidae
Sicyoniidae
Solenoceridae
Sergestoidea
Luciferidae
Sergestidae
4. Stenopodidea – coral shrimps and Venus shrimps
Macromaxillocarididae
Spongicolidae
Stenopodidae
5. Procarididea – shrimps
Procarididae
6. Caridea – shrimps
Alpheoidea
Alpheidae
Barbouriidae
Bythocarididae
Hippolytidae
Lysmatidae
Merguiidae
Ogyrididae
Thoridae
Atyoidea
Atyidae
Bresilioidea
Agostocarididae
Alvinocarididae
Bresiliidae
Disciadidae
Pseudochelidae
Campylonotoidea
Bathypalaemonellidae
Campylonotidae
Crangonoidea
Crangonidae
Glyphocrangonidae
Nematocarcinoidea
Eugonatonotidae
Lipkiidae
Nematocarcinidae
Rhynchocinetidae
Oplophoroidea
Acanthephyridae
Oplophoridae
Palaemonoidea
Palaemonidae
Pandaloidea
Chlorotocellidae
Pandalidae
Pasiphaeoidea
Pasiphaeidae
Physetocaridoidea
Physetocarididae
Processoidea
Processidae
Psalidopodoidea
Psalidopodidae
Stylodactyloidea
Stylodactylidae
Family incerta sedis Anchialocarididae
7. Astacidea – scampi and crayfish
Enoplometopidae
Nephropidae
8. Glypheidea – lobsters
Glypheidae
9. Axiidea – sponge shrimps, ghost shrimps and mud lobsters
Anacalliacidae
Axiidae
Callianassidae
Callianideidae
Callianopsidae
Callichiridae
Ctenochelidae
Eucalliacidae
Micheleidae
Paracalliacidae
Strahlaxiidae
10. Gebiidea – mud lobsters and sponge shrimps
Laomediidae
Thalassinidae
Upogebiidae
11. Achelata – spiny lobsters, slipper lobsters and bugs
Palinuridae
Scyllaridae
12. Polychelida – deep-sea lobsters
Polychelidae
13. Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters
Chirostyloidea
Chirostylidae
Eumunididae
Kiwaidae
Sternostylidae
Galatheoidea
Galatheidae
Munididae
Munidopsidae
Porcellanidae
Hippoidea
Albuneidae
Blepharipodidae
Hippidae
Lomisoidea
Lomisidae
Paguroidea
Coenobitidae
Diogenidae
Lithodidae
Paguridae
Parapaguridae
Pylochelidae
Pylojacquesidae
Xylopaguridae
Colour plates
14. Brachyura – crabs
Section Cyclodorippoida
Cyclodorippoidea
Cyclodorippidae
Cymonomidae
Phyllotymolinidae
Section Dromiacea
Dromioidea
Dromiidae
Dynomenidae
Homolodromioidea
Homolodromiidae
Section Homoloida
Homoloidea
Homolidae
Latreilliidae
Poupiniidae
Section Raninoida
Raninoidea
Lyreididae
Raninidae
Section Eubrachyura
Subsection Heterotremata
Aethroidea
Aethridae
Bellioidea
Belliidae
Heteroziidae
Bythograeoidea
Bythograeidae
Calappoidea
Calappidae
Matutidae
Cancroidea
Atelecyclidae
Cancridae
Carpilioidea
Carpiliidae
Cheiragonoidea
Cheiragonidae
Corystoidea
Corystidae
Dairoidea
Dairidae
Dorippoidea
Dorippidae
Ethusidae
Eriphioidea
Dacryopilumnidae
Eriphiidae
Hypothalassiidae
Menippidae
Oziidae
Platyxanthidae
Goneplacoidea
Acidopsidae
Chasmocarcinidae
Conleyidae
Euryplacidae
Goneplacidae
Litocheiridae
Mathildellidae
Progeryonidae
Scalopidiidae
Sotoplacidae
Vultocinidae
Hexapodoidea
Hexapodidae
Hymenosomatoidea
Hymenosomatidae
Leucosioidea
Iphiculidae
Leucosiidae
Majoidea
Epialtidae
Inachidae
Inachoididae
Majidae
Mithracidae
Oregoniidae
Orithyioidea
Orithyiidae
Palicoidea
Crossotonotidae
Palicidae
Parthenopoidea
Dairoididae
Parthenopidae
Pilumnoidea
Galenidae
Pilumnidae
Tanaochelidae
Portunoidea
Brusiniidae
Carcinidae
Geryonidae
Nautilocorystidae
Ovalipidae
Pirimelidae
Polybiidae
Portunidae
Thiidae
Pseudocarcinoidea
Pseudocarcinidae
Pseudozioidea
Christmaplacidae
Pilumnoididae
Planopilumnidae
Pseudoziidae
Retroplumoidea
Retroplumidae
Trapezioidea
Domeciidae
Tetraliidae
Trapeziidae
Trichopeltarioidea
Trichopeltariidae
Xanthoidea
Linnaeoxanthidae
Panopeidae
Pseudorhombilidae
Xanthidae
Subsection Thoracotremata
Aphanodactyloidea
Aphanodactylidae
Cryptochiroidea
Cryptochiridae
Grapsoidea
Grapsidae
Leptograpsodidae
Percnidae
Plagusiidae
Sesarmidae
Varunidae
Xenograpsidae
Ocypodoidea
Camptandriidae
Dotillidae
Heloeciidae
Macrophthalmidae
Mictyridae
Ocypodidae
Xenophthalmidae
Pinnotheroidea
Pinnotheridae
Glossary
Credits
Index to genus-group, family-group and higher taxonomic names

Citation preview

Marine Decapod Crustacea A Guide to Families and Genera of the World

Gary C. B. Poore and Shane T. Ahyong

Marine Decapod Crustacea A Guide to Families and Genera of the World

Gary C. B. Poore and Shane T. Ahyong

© Gary Poore and Shane Ahyong 2023 All rights reserved. Except as permitted by applicable copyright laws, no part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, duplicating or otherwise, without the prior permission of the copyright owner. Contact CSIRO Publishing for all permission requests. Gary Poore and Shane Ahyong assert their right to be known as the authors of this work. A catalogue record for this book is available from the National Library of Australia. ISBN: 9781486311781 (hbk) ISBN: 9781486311798 (epdf) ISBN: 9781486311804 (epub) Published in print in Australia and New Zealand, and in all other formats throughout the world, by CSIRO Publishing. CSIRO Publishing Private Bag 10 Clayton South VIC 3169 Australia Telephone: +61 3 9545 8400 Email: [email protected] Website: www.publish.csiro.au Sign up to our email alerts: publish.csiro.au/earlyalert Published in print only, throughout the world (except in Australia and New Zealand), by CRC Press, with ISBN 9781032138022. CRC Press 6000 Broken Sound Parkway NW, Suite 300, Boca Raton, FL 33487-2742 and 2 Park Square, Milton Park, Abingdon, Oxon, OX14 4RN Website: www.routledge.com CRC Press is an imprint of Taylor & Francis Group, LLC Front cover: (top, left to right) Hoplophrys oatesi (photo by Tin-Yam Chan), Goniopsis cruentata (photo by Arthur Anker), Ranilia muricata (photo by Darryl L. Felder), Puerulus gibbosus (photo by Tin-Yam Chan), Ciliopagurus strigatus (photo by Andrey Ryanskyi); (bottom) Enoplometopus crosnieri (photo by Tin-Yam Chan) Spine: Ovalipes catharus (photo by Shane T. Ahyong) Back cover: (right, top to bottom) Alienaxiopsis clypeata (photo by Zdeněk Ďuriš), Cervimunida johni (photo by Arthur Anker), Aristaeopsis edwardsiana (photo by Tin-Yam Chan); (left) Trichia horii (photo by Tin-Yam Chan) Cover design by Cath Pirret Typeset by Envisage Information Technology Printed in Singapore by Markono Print Media Pte Ltd CSIRO Publishing publishes and distributes scientific, technical and health science books, magazines and journals from Australia to a worldwide audience and conducts these activities autonomously from the research activities of the Commonwealth Scientific and Industrial Research Organisation (CSIRO). The views expressed in this publication are those of the author(s) and do not necessarily represent those of, and should not be attributed to, the publisher or CSIRO. The copyright owner shall not be liable for technical or other errors or omissions contained herein. The reader/ user accepts all risks and responsibility for losses, damages, costs and other consequences resulting directly or indirectly from using this information. CSIRO acknowledges the Traditional Owners of the lands that we live and work on across Australia and pays its respect to Elders past and present. CSIRO recognises that Aboriginal and Torres Strait Islander peoples have made and will continue to make extraordinary contributions to all aspects of Australian life including culture, economy and science. CSIRO is committed to reconciliation and demonstrating respect for Indigenous knowledge and science. The use of Western science in this publication should not be interpreted as diminishing the knowledge of plants, animals and environment from Indigenous ecological knowledge systems. The paper this book is printed on is in accordance with the standards of the Forest Stewardship Council ® and other controlled material. The FSC ® promotes environmentally responsible, socially beneficial and economically viable management of the world’s forests.

Oct22_01

Contents

Acknowledgements

xi

1. Introduction

1

What is a crustacean? Scope of this work History and resources Structure of this book, and some advice! 2. Systematics

3.

1 2 2 4 9

Evolution of the Crustacea Classification of the Crustacea Classification of the Decapoda Key to suborders and infrorders of Decapoda and shrimp-like marine crustaceans

9 10 14 14

Dendrobranchiata – prawns and shrimps

19

Penaeoidea  21 Aristeidae21 Benthesicymidae23 Penaeidae27 Sicyoniidae38 Solenoceridae38 Sergestoidea41 Luciferidae41 Sergestidae42 4.

Stenopodidea – coral shrimps and Venus shrimps

51

Macromaxillocarididae52 Spongicolidae52 Stenopodidae55 5.

Procarididea – shrimps

59

Procarididae59 6.

Caridea – shrimps

61

Alpheoidea65 Alpheidae65 Barbouriidae80

iv

Marine Decapod Crustacea

Bythocarididae82 Hippolytidae  83 Lysmatidae88 Merguiidae90 Ogyrididae91 Thoridae91 Atyoidea95 Atyidae95 Bresilioidea95 Agostocarididae  96 Alvinocarididae96 Bresiliidae99 Disciadidae100 Pseudochelidae102 Campylonotoidea102 Bathypalaemonellidae102 Campylonotidae103 Crangonoidea103 Crangonidae103 Glyphocrangonidae110 Nematocarcinoidea110 Eugonatonotidae111 Lipkiidae111 Nematocarcinidae112 Rhynchocinetidae114 Oplophoroidea114 Acanthephyridae115 Oplophoridae117 Palaemonoidea119 Palaemonidae119 Pandaloidea158 Chlorotocellidae158 Pandalidae160 Pasiphaeoidea165 Pasiphaeidae165 Physetocaridoidea168 Physetocarididae168 Processoidea168 Processidae168 Psalidopodoidea170 Psalidopodidae170

Contents

v

Stylodactyloidea170 Stylodactylidae171

7.

Family incerta sedis Anchialocarididae

172

Astacidea – scampi and crayfish

175

Enoplometopidae176 Nephropidae177 8.

Glypheidea – lobsters

183

Glypheidae183 9.

Axiidea – sponge shrimps, ghost shrimps and mud lobsters

185

Anacalliacidae187 Axiidae187 Callianassidae204 Callianideidae215 Callianopsidae217 Callichiridae219 Ctenochelidae225 Eucalliacidae228 Micheleidae231 Paracalliacidae233 Strahlaxiidae233 10.

Gebiidea – mud lobsters and sponge shrimps

237

Laomediidae237 Thalassinidae240 Upogebiidae241 11.

Achelata – spiny lobsters, slipper lobsters and bugs

247

Palinuridae248 Scyllaridae252 12.

Polychelida – deep-sea lobsters

259

Polychelidae259 13.

Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

263

Chirostyloidea265 Chirostylidae266 Eumunididae269 Kiwaidae270 Sternostylidae271

vi

Marine Decapod Crustacea

Galatheoidea  272 Galatheidae273 Munididae277 Munidopsidae283 Porcellanidae286 Hippoidea295 Albuneidae296 Blepharipodidae298 Hippidae299 Lomisoidea300 Lomisidae300 Paguroidea301 Coenobitidae303 Diogenidae304 Lithodidae311 Paguridae317 Parapaguridae339 Pylochelidae342 Pylojacquesidae345 Xylopaguridae346

Colour plates

349

14.

Brachyura – crabs

421

Section Cyclodorippoida  434 Cyclodorippoidea434 Cyclodorippidae435 Cymonomidae439 Phyllotymolinidae441 Section Dromiacea 442 Dromioidea443 Dromiidae443 Dynomenidae455 Homolodromioidea457 Homolodromiidae457 Section Homoloida 459 Homoloidea459 Homolidae460 Latreilliidae465 Poupiniidae466 Section Raninoida

466

Contents

vii

Raninoidea467 Lyreididae468 Raninidae468 Section Eubrachyura 473 Subsection Heterotremata 473 Aethroidea474 Aethridae474 Bellioidea477 Belliidae477 Heteroziidae478 Bythograeoidea478 Bythograeidae479 Calappoidea481 Calappidae482 Matutidae486 Cancroidea487 Atelecyclidae488 Cancridae488 Carpilioidea491 Carpiliidae491 Cheiragonoidea492 Cheiragonidae492 Corystoidea492 Corystidae493 Dairoidea494 Dairidae495 Dorippoidea495 Dorippidae496 Ethusidae498 Eriphioidea500 Dacryopilumnidae501 Eriphiidae502 Hypothalassiidae503 Menippidae504 Oziidae505 Platyxanthidae508 Goneplacoidea510 Acidopsidae511 Chasmocarcinidae514 Conleyidae519 Euryplacidae519

viii

Marine Decapod Crustacea

Goneplacidae523 Litocheiridae530 Mathildellidae531 Progeryonidae533 Scalopidiidae534 Sotoplacidae534 Vultocinidae536 Hexapodoidea536 Hexapodidae536 Hymenosomatoidea540 Hymenosomatidae541 Leucosioidea550 Iphiculidae551 Leucosiidae551 Majoidea577 Epialtidae580 Inachidae608 Inachoididae616 Majidae621 Mithracidae634 Oregoniidae639 Orithyioidea646 Orithyiidae646 Palicoidea647 Crossotonotidae647 Palicidae648 Parthenopoidea650 Dairoididae651 Parthenopidae652 Pilumnoidea664 Galenidae665 Pilumnidae666 Tanaochelidae687 Portunoidea687 Brusiniidae689 Carcinidae690 Geryonidae691 Nautilocorystidae694 Ovalipidae695 Pirimelidae696

Contents

ix

Polybiidae697 Portunidae700 Thiidae717 Pseudocarcinoidea718 Pseudocarcinidae718 Pseudozioidea719 Christmaplacidae720 Pilumnoididae720 Planopilumnidae722 Pseudoziidae723 Retroplumoidea724 Retroplumidae724 Trapezioidea725 Domeciidae726 Tetraliidae727 Trapeziidae728 Trichopeltarioidea731 Trichopeltariidae731 Xanthoidea733 Linnaeoxanthidae735 Panopeidae735 Pseudorhombilidae741 Xanthidae749 Subsection Thoracotremata 792 Aphanodactyloidea792 Aphanodactylidae793 Cryptochiroidea795 Cryptochiridae795 Grapsoidea801 Grapsidae803 Leptograpsodidae806 Percnidae806 Plagusiidae807 Sesarmidae809 Varunidae824 Xenograpsidae836 Ocypodoidea837 Camptandriidae838 Dotillidae845 Heloeciidae849

x

Marine Decapod Crustacea

Macrophthalmidae849 Mictyridae854 Ocypodidae855 Xenophthalmidae860 Pinnotheroidea861 Pinnotheridae862 Glossary

881

Credits

887



891

Index to genus-group, family-group and higher taxonomic names

Acknowledgements

Our interest in crustacean taxonomy, that of decapods in particular, is deep and longstanding. It has been encouraged by our opportunities to work with many specialists, in the laboratory and in the field, whose enthusiasm has rubbed off. We thank them all. Some have since passed: Sandy Bruce, Fenner Chace, Alain Crosnier, Jacques Forest, Lipke Holthuis, Brian Kensley, Ray Manning, Pat McLaughlin, Ngan Kee Ng, Michèle de Saint Laurent and Vasily Spiridonov. Others continue to be inspirational today: Arthur Anker, Akira Asakura, Keiji Baba, Chris Boyko, Peter Castro, Tin-Yam Chan, Paul Clark, Peter Davie, Sammy De Grave, Zdeněk Ďuriš, Peter Dworschak, Darryl Felder, Bella Galil, Danièle Guinot, Tomoyuki Komai, Rafael Lemaitre, Colin McLay, Enrique Macpherson, Jose Christopher Escano Mendoza, Tohru Naruse, Peter K. L. Ng, Masayuki Osawa, Bertrand Richer de Forges, Kareen Schnabel, Masatsune Takeda, Joanne Taylor and Marcos Tavares. All of these, either directly or indirectly, helped with this work. This review has greatly benefitted from ready access to lists of taxa and published works. We appreciate very much the catalogue at WoRMS (World Register of Marine Species) and thank the Database Management Team led by Leen Vandepitte and the decapod editors who keep the catalogue up to date. We are also thankful for the literature available on the Biodiversity Heritage Library and commend all those responsible for making this available. Although this is largely a synthetic work relying primarily on what has been published, it has only been made possible with access to diverse and well catalogued collections at Museums Victoria, Melbourne, the Australian Museum, Sydney, the Lee Kong Chian Natural History Museum,

Singapore, the Muséum nationale d’Histoire naturelle, Paris, and the National Museum of Natural History, Smithsonian Institution, Washington, D.C. Collection and expedition staff at these and other museums have enabled us to explore decapod diversity. Most illustrations are original but we thank the publishers of several journals for allowing us to reproduce figures directly: Australian Museum, Bishop Museum Press, E.J. Brill, CSIRO Publishing, Lee Kong Chian Natural History Museum, Oxford University Press, Pensoft Publishing, Queensland Museum, Magnolia Press, Muséum National d’Histoire Naturelle, Paris, National Science Museum, Tokyo, Royal Society of New Zealand, and University of Hawaii Press. Several photographers allowed us to reproduce their colour or black and white images. We thank all those mentioned in the credits but special mention to those who generously provided dozens of photos: Arthur Anker, Laure Corbari, Tin-Yam Chan, Darryl Felder, Karen Gowlett-Holmes, the late Michael Marmach, Mos Natapatt, Gustav Paulay, Ondřej Radosta, Andrey Ryanskyi and Swee-Hee Tan. We appreciate the confidence of Briana Melideo, Eloise Moir-Ford, Tracey Kudis and Mark Hamilton, CSIRO Publishing, in this project and their patience in getting it published. More than anyone we thank our wives. Gary thanks Lynsey for her support and encouragement in this and other research projects over many decades. Shane is most grateful to Rachel for her patience, constant support and for allowing decapods to hold more than their fair share of attention.

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1. Introduction

Identification of species, the most fundamental unit in biology, is the essential first step of any biological study. Confidence in any report on a species’ ecology, physiology or behaviour, and trust in a species list in multispecies research depend on the study organism, or organisms, being reliably labelled with their current Linnean name. Knowing that one’s target organism is several species rather than one, or vice versa, has considerable impact on the conclusions that can be drawn (Schram and Ng 2012). Naming of species is managed by taxonomists and is governed by rules and conventions (ICZN 1999). Getting into the taxonomic literature to ensure that the subject of a researcher’s study is correctly identified can be difficult for a non-taxonomist. Names and the classificatory hierarchy may change with time in the light of research on taxonomic affinities and relationships, but changes are traceable. The problem is exacerbated in multispecies projects. This guidebook aims to enable biologists to identify any marine decapod to the level of family and genus. The dichotomous keys lead to diagnoses of 189 families and 2121 genera. A book this size cannot be a resource for identification to species level but references to key literature are included that should facilitate more precise identification. Charles Darwin’s friend and advocate, Thomas Huxley, wrote a book about the crayfish in 1880 that he hoped would stimulate an interest in the zoological questions that were important then. Many of ‘the great zoological questions’ to which he referred are still being addressed 140 years later. This guidebook is an overview of the players providing evidence for these questions. Crab, shrimp, prawn, lobster, crayfish, yabby, and bug are just some of the names used in English to describe members of the decapod Crustacea. To these can be added more culinary terms from other languages  – langoustine, scampi and udang. Most people would recognise a crustacean on the sea shore or on the dining table. Crustaceans are a group of related animals, meaning they had a common ancestor some time in the past. The features that they share are described in this introductory chapter. This volume deals with only some crustaceans, those called ‘decapods’ and only those found in marine environments. Much has been written about crustaceans before, and, in Chapter 1, this is reviewed to place presentday knowledge in context. This introductory chapter explains to the reader how the book has been planned and what to expect from each section.

Chapter 2 deals with the higher systematics of Crustacea and explains how the Decapoda are related to the many other crustaceans. General morphology is dealt with and specialist terms introduced. Chapters 3 to 14 cover the major decapod crustacean groups in turn. Chapter 3 deals with what we Australians call prawns but are shrimp in the USA, Chapter 4 covers coral shrimps, Chapters 5 and 6 deal with all other shrimps, Chapter 7 with crayfish and scampi, Chapter 8 with glypheid lobsters, Chapters 9 and 10 with sponge shrimps and ghost shrimps, Chapter 11 with rock lobsters and bugs, Chapter 12 with deep-sea blind lobsters, Chapter 13 with hermit crabs and squat lobsters, and Chapter 14 with ‘true’ crabs. There follows a glossary because we have no choice but to learn a new language to identify these animals. Diagrams further help to explain the special terms applicable to different groups of decapods.

What is a crustacean? Insects, spiders, scorpions, millipedes are just some of the Phylum Arthropoda, that large group of animals distinguished from all others by their hard skin, or exoskeleton, and a segmented body. Crustaceans belong to this phylum too and may be called the insects of the sea. In fact, it may be more appropriate to call insects crustaceans of the land, because insects are now believed to have evolved from within the crustaceans (Oakley et  al. 2012; von Reumont and Edgecombe 2020). While insects are diverse and common on land, crustaceans are diverse and common in the sea (Plate 1). Unlike insects, which are more or less excluded from marine environments, crustaceans do get on to land and into fresh water. Crustaceans, then, are arthropods that usually have many pairs of legs, a pair per body segment, and above all, have two pairs of antennae (special sensory limbs), one pair on each of the first two segments of the head. Insects, in contrast, have relatively few pairs of legs, and only one pair of antennae. Chapter 2 goes into more detail about the classification of crustaceans. These details are of little importance when it comes to recognising a decapod crustacean. We know from childhood what is a crab, shrimp or lobster and have never counted legs or antennae. In a catch of animals from a marine habitat it is simple to separate crustaceans from fish, snails, worms and seastars. Crustaceans are obvious to the marine biologist in the way that ants and flies are to

2

Marine Decapod Crustacea

land-lubbers for the simple reason that they are so abundant and so diverse. Densities may be in thousands of individuals per square metre of sea floor or cubic metre of ocean water and commonly many different kinds live side by side. The global number of marine species for which we have names is more than 215 000 (Costello et al. 2013) but this is just a small fraction of the true number, which could be a million for all the world’s oceans (Appeltans et al. 2012) – or perhaps, ten million. The reason for this underestimate is that only a small percentage has been described and are known by formal names. The number of named species of Crustacea is at least 66  914 of which 51 267 are marine and 13 501, about one-quarter, are decapods (Ahyong et  al. 2011; Appeltans et  al. 2012, updated from WoRMS). The significance of crustaceans in the ecology of the oceans cannot be ignored – they can be scavengers, predators, parasites and food for many of their coinhabitants. Crustaceans range in size from one-tenth of a millimetre maximum diameter (a tantulocarid parasite) to four metres across (a Japanese spider crab). The mass of Antarctic krill in the oceans may be greater than that of any other single species. One species of oceanic copepod may be the most common animal on earth; and, crustaceans have been found almost as far back in the fossil record as any group of complex animals.

Scope of this work This guidebook tries to bridge the gap between taxonomists and other marine biologists. These non-taxonomists, ecologists, biogeographers, physiologists etc., are the target audience for this volume. The gap exists in part because, while taxonomists are experts in identification, they often write for each other in an arcane language. On the basis of new evidence they change accepted Linnean names of what were thought familiar species. Our aim is to write in a consistent language that is more accessible to all. This volume serves as a document on the state of play in decapod taxonomy in mid-2022. Systematics and classification are dynamic disciplines and the accepted family and generic names published today will not apply in the near future. Nevertheless, these data are a point of access to the taxonomic literature. We have not described new genera or species nor, with few exceptions, have we made any taxonomic changes. However, we have discovered errors and inconsistencies in past work and have corrected these with new interpretations. We have relied on the most recent, or most generally accepted, classifications and lists of names among which are a few we treat as new synonyms. The work is confined to marine decapods. Exclusively freshwater or terrestrial taxa are omitted, such as land and

freshwater crabs (Parathelphusidae, Potamoidea, Pseudothelphusoidea, Trichodactyloidea, Gecarcinidae, Glyptograpsidae), freshwater shrimps (Xiphocarididae, Desmocarididae, Euryrhynchidae and Typhlocarididae) and freshwater crayfish (Cambaridae, Cambaroididae and Parastacidae). Families with marine, estuarine and freshwater representatives are diagnosed but exclusively freshwater genera are listed without appearing in keys.

History and resources Like any group of animals, crustaceans have long attracted the attention of biologists and artists. Beautiful images of lobsters and crabs can be found in mosaics of ancient Roman cities, eighteenth century European still-life painting and in the paintings of indigenous peoples of all continents. The interest of these artists was surely because crustaceans are edible. Linnaeus included crustaceans in his Systema Naturae (Linnaeus 1758), mostly in the genus Cancer. The age of European biological ship-based exploring expeditions began in the late eighteenth century and continued through to the mid-twentieth century. The series of scientific reports, many taxonomic, associated with ships such as H.M.S. Challenger, R.I.M.S.S. Investigator, the Siboga, or with the United States Exploring Expedition remain relevant today. Today this tradition is continued by research vessels of several nations. Documentation of biodiversity in the Indo-West Pacific has benefitted from the MUSORSTOM, now Tropical Deep-Sea Benthos, series of volumes (Poore 2004a; Richer de Forges et al. 2021). General books on Crustacea are found only in specialist libraries. Old books have a fascination of their own because they established the classifications and terminology still used today. The Reverend T.R.R. Stebbing’s A history of Crustacea. Recent Malacostraca (1893) and William T.  ­Calman’s The life of Crustacea (1911) are classics. Patsy A. McLaughlin’s Comparative morphology of Recent Crustacea (1980) is a useful practical guide to the definition of living taxa but is not a taxonomic guide and has no ecological information. Alfred Kaestner’s (1980) Crustacea is a useful translation of a systematic German text but is an academic treatment, now out of print. Frederick R. Schram’s major volumes, Schram (1986) and his significantly expanded coauthored work (Schram and Koenemann 2021), are packed with information and include all the fossil taxa, but also have somewhat controversial and idiosyncratic classifications. They largely address other crustacean biologists, however, and are less accessible to non-specialists. Martin and Davis (2001) provided a valuable modern synthesis of all the families of Crustacea. This remains useful but quickly became out of date. For decapods, De Grave et al.

1 – Introduction

(2009) replaced their list. The recent and continuing series The Natural History of the Crustacea (Anger et  al. 2020; Derby and Thiel 2014; Poore and Thiel 2020; Thiel and Watling 2015; Thiel and Wellborn 2018; Watling and Thiel 2013) deals with all aspects of the biology of Crustacea. None of these texts is useful for species identification. Many regional studies are dedicated to that task. While these are valuable references some of the species names

given are no longer accepted due to later synonymy or rearrangement of higher classifications. Nevertheless, former species names can usually be traced and updated using the World Register of Marine Species (WoRMS). Here, we list the major regional studies geographically annotating each by the major taxon covered. Only works with keys or species diagnoses are listed. Faunal lists and picture atlases are not included.

Japan

Brachyura

Sakai (1976)

PR China

Brachyura

Dai and Yang (1991)

Taiwan

Primitive Brachyura

Ahyong et al. (2009)

Taiwan

Paguroidea

McLaughlin et al. (2007)

Taiwan

Crab-like Anomura

Chan (2010b)

Southern Australia

Decapoda

Poore (2004b)

New Zealand

Brachyura and crab-like Anomura

McLay (1988)

New Zealand

Dendrobranchiata, Caridea

Webber et al. (1990)

India

Brachyura

Dev Roy and Bhadra (2001, 2005, 2008, 2011); Dev Roy and Nandi (2012)

Persian Gulf

Brachyura

Naderloo (2017)

Southern Africa

Decapoda

Barnard (1950); Emmerson (2016)

Southern Africa

Dendrobranchiata, Caridea

Kensley (1972)

West Africa

Brachyura

Manning and Holthuis (1981)

France

Decapoda

Noël (1992)

United Kingdom

Shrimps and prawns

Smaldon et al. (1993)

United Kingdom

Brachyura

Ingle (1980)

United Kingdom

Lobsters, mud shrimps, anomuran crabs

Ingle and Christiansen (2004)

North-eastern Atlantic

Hermit crabs

Ingle (1993)

Americas

Brachyura

Rathbun (1918, 1937)

Eastern USA

Decapoda

Williams (1984)

Brazil

Decapoda

de Melo (1996, 1999)

Mexico, Pacific coast

Dendrobranchiata

Hendrickx and Estrada Navarrete (1996)

Mexico, Pacific coast

Decapoda

Hendrickx (1993)

Southern California, USA

Crustacea

SCAMIT (2009 onwards)

Chile

Brachyura

Garth (1957)

Chile

Anomura

Haig (1955)

Chile

Macrura

Holthuis (1952)

We have referred to all these books in the course of research for this volume. However, our task would not have been possible without three web-based resources. (1) The World Register of Marine Species (WoRMS, http:// www.marinespecies.org/) is ‘an authoritative classification and catalogue of marine names’ (WoRMS Editorial Board 2022). The WoRMS database is managed at the Flanders Marine Institute (VLIZ) and is updated by a

3

team of volunteer taxonomic editors. All family and generic names were downloaded from WoRMS in late 2018, and subsequently updated and revised, to become the foundation of our review. WoRMS also provides access to much of the relevant taxonomic literature, as citations and often as downloadable publications. (2) The Biodiversity Heritage Library (https://www.­ biodiversitylibrary.org/) houses scanned pages of most

4

Marine Decapod Crustacea

of the non-copyrighted books and journals held in museums and other institutions, a rich source of old taxonomic literature. (3) The Ocean Biodiverity Information System (OBIS, https://obis.org/) is ‘a global open-access data and information clearing-house on marine biodiversity for science, conservation and sustainable development’. OBIS provides distributional and depth-range data by species and higher taxa derived from museum, fisheries and other sources, and was one of our initial sources for distributional information.

Structure of this book, and some advice! The volume follows what is now accepted as the new classification of the decapods, each chapter devoted to a major group. This classification differs in some details from the most recent wide-ranging lists (Chan 2010a; De Grave and Fransen 2011; De Grave et al. 2009; Ng et al. 2008). These works provided catalogues of families, genera and in some cases species. All continue to be updated on WoRMS (WoRMS Editorial Board 2022). All chapters follow the same format: general introduction; diagnosis; a key to families and/or superfamilies; superfamilies and families treated in alphabetical order, each with an introduction, diagnosis and keys to genera; followed by all genera in alphabetical order (sometimes grouped in subfamilies). Each genus is diagnosed – maximum size, depth range, distribution and the number of described species are given. Diagnoses. Diagnoses of families and genera are original, usually simplified from more detailed diagnoses in the cited literature. All were generated initially from data stored in DELTA databases (Dallwitz et al. 1993 onwards). Diagnoses attempt to characterise each taxon with the same suite of characters but this was not always possible. DELTA facilitates use of ‘implicit attributes’, that is features that are common to most genera within a family. To save repetition these features are listed separately following the family diagnosis and only exceptions are included with relevant generic diagnoses. The size given is usually the carapace length (cl.) of the largest species in the genus. Rarely, other standard dimensions are used. Ecology. The ecological range of each genus is summarised as depth ranges following widely accepted bathymetric classes. For benthic groups these are: intertidal; subtidal (including inshore bays, 20 m or shallower); shelf (20–200 m); slope (200–2000 m); bathyal (2000–3500 m); abyssal (3500–6500 m); hadal (> 6500 m). Special ecological affinities are also noted for estuarine genera or those living in marine anchialine caves. For pelagic or planktonic groups depth ranges are as follows: neritic (< 200 m); mesopelagic

(200–1000 m); bathypelagic (> 1000 m). Actual recorded depth ranges are given when these data were readily available. Sometimes other ecological descriptors such as preferred sediment type or associated fauna are added but often this detail can be found in the family introduction. Distributions. Distributions of all benthic genera occurring down to slope depths are given in terms of the 12 MEOW (Marine Ecoregions of the World; Spalding et  al. 2006) realms as follows: Arctic; Temperate Northern Atlantic; Tropical Atlantic; Temperate Northern Pacific; Western Indo-Pacific; Central Indo-Pacific; Eastern Indo-Pacific; Tropical Eastern Pacific; Temperate South America; Temperate Southern Africa; Temperate Australasia; Southern Ocean (Fig. 1.1). The realms are annotated when they do not adequately describe the distribution. For example, the annotations E and W are added when the known range is only within the eastern and western part of a realm such as Temperate Northern Pacific and Tropical Atlantic. Further details are added or substituted where appropriate, e.g. ­Australia or New Zealand to Temperate Australasia. ‘IndoWest Pacific’ is substituted when a genus ranges across the Western Indo-Pacific, Central Indo-Pacific and Eastern Indo-Pacific realms. Distributions of all benthic genera occurring at bathyal or deeper depths are given in terms of the 14 GOODS bathyal provinces (Global Open Oceans and Deep Seabed; UNESCO 2009) as follows: Arctic; Northern Atlantic; Brazil Basin; Angola & Sierra Leone Basins; Argentine Basin; E Antarctic Basin; W Antarctic; Indian; E Pacific basins; S Pacific; Central Pacific; N Central Pacific; N  Pacific; W Pacific basins. For oceanic pelagic genera, many widespread, we follow the simple scheme of Vereshchaka et al. (2014): Atlantic Ocean; Indian Ocean; Central and W Pacific Oceans; E Pacific Ocean. References. The literature cited, over 2650 papers, books or chapters, concentrates on the most relevant taxonomic references, those with more extensive diagnoses and keys to species and/or information on distributions. Some key ecological works may be cited. Keys and illustrations. Identification is achieved through the use of dichotomous keys adapted from many originally published in the primary literature, or developed from scratch. The temptation when using keys is to assume when you reach the last couplet that your specimen belongs in that genus. It is worthwhile to check the diagnosis and the distribution given. The keys are supported by illustrations. Many of the illustrations are reworking of published figures, others are new or reproductions from the literature. References to elements in black and white figures are given in the key and some diagnoses; references to colour plates appear with each genus. Terminology. Keys written by expert taxonomists for other taxonomists use arcane language. To make matters

1 – Introduction

5

Fig. 1.1.  Biogeographic regions. Twelve realms of the Marine Ecoregions of the World (MEOW) apply from intertidal to slope depths (0–2000 m depth) and are: Arctic; Temperate Northern Atlantic; Tropical Atlantic; Temperate Northern Pacific; Western Indo-Pacific; Central Indo-Pacific; Eastern Indo-Pacific; Tropical Eastern Pacific; Temperate South America; Temperate Southern Africa; Temperate Australasia; and Southern Ocean (Spalding et al. 2006 for detail). Numbers in circles refer to the 14 Global Open Oceans and Deep Seabed (GOODS) lower bathyal provinces (800–3000 m depth) as follows: 1, Arctic; 2, Northern N Atlantic; 3, Northern N Pacific; 4, N Atlantic; 5, SE Pacific ridges; 6, New Zealand/Kermadec; 7, Cocos Plate; 8, Nazca Plate; 9, Antarctic; 10, Subantarctic; 11, Indian; 12, W Pacific; 13, S Atlantic; 14, N Pacific (UNESCO 2009 for detail).

worse many specialise in a limited range of taxa using terms that only those specialists appreciate, or spell the same term differently. Those exploring the diversity of copepods rarely venture into the classification of larger decapods. Even within decapods, few taxonomists are equally competent across this diverse taxon. One consequence of this is that each group of taxonomists develops its own jargon, using different terms for essentially the same structure depending on which group is being discussed. Further, relative descriptors, such as ‘small’ and ‘large’ are treated as diagnostic terms but are not meaningful to outsiders. Inconsistent taxonomic names, variable terminology and indeterminate descriptors are confusing for the non-specialist. We have prepared this volume using a consistent terminology that is hopefully understandable by our target audience. The terms used by crustacean taxonomists were compiled by Martin (2005) for the Assembling the Tree of Life (AToL) Decapod project. He relied for decapods on several references among this list (Bliss 1982; Chace and Hobbs 1969; Holthuis 1991; Ng 1998; Pérez Farfante and Kensley 1997; Poore 2004b; Williams 1984). The glossary at the end of this volume should be helpful and is illustrated with special terms applicable to only some groups.

References Ahyong ST, Naruse T, Tan SH, Ng PKL (2009) Part II. Infraorder Brachyura: Sections Dromiacea, Raninoida, Cyclodorippoida. In Crustacean fauna of Taiwan: Brachyuran crabs, volume 1 – Carcinology in Taiwan and Dromiacea, Raninoida, Cyclodorippoida. (Eds Chan TY, Ng PKL, Ahyong ST, Tan SH) pp. 27–198. National Taiwan Ocean University, Keelung. Ahyong ST, Lowry JK, Alonso M, Bamber RN, Boxshall GA, et al. (2011) Subphylum Crustacea Brünnich, 1772. In: Zhang Z-Q (Ed.) Animal biodiversity: an outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148, 164–191. Anger K, Harzsch S, Thiel M (Eds) (2020) Developmental biology and larval ecology. The Natural History of the Crustacea Vol. 7. Oxford University Press, New York. Appeltans W, Ahyong ST, Anderson G, Angel MV, Artois T, et al. (2012) The magnitude of global marine species diversity. Current Biology 22, 2189–2202. doi:10.1016/j.cub.2012.09.036 Barnard KH (1950) Descriptive catalogue of South African decapod Crustacea (crabs and shrimps). Annals of the South African Museum 38, 1–837. Bliss DE (1982) Shrimps, lobsters and crabs: their fascinating life story. New Century Publishers, Piscatawey. Calman WT (1911) The life of Crustacea. Methuen & Co., London. Chace FA, Hobbs HH (1969) Bredin-Archbold-Smithsonian Biological survey of Dominica. The freshwater and terrestrial decapod crustaceans of the West Indies with special reference to Dominica. Bulletin of the United States National Museum 292, 1–258. doi:10.5479/si.03629236.292.1

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Chan T-Y (2010a) Annotated checklist of the world’s marine lobsters (Crustacea: Decapoda: Astacidea, Glypheidea, Achelata, Polychelida). Raffles Bulletin of Zoology Supplement 23, 153–181. Chan T-Y (Ed.) (2010b) Crustacean fauna of Taiwan: Crab-like anomurans (Hippoidea, Lithodoidea and Porcellanidae). National Taiwan Ocean University, Keelung. Costello MJ, Bouchet P, Boxshall GA, Fauchald K, Gordon D, et al. (2013) Global coordination and standardisation in marine biodiversity through the World Register of Marine Species (WoRMS) and related databases. PLoS One 8, e51629. doi:10.1371/journal.pone.0051629 Dai A-Y, Yang S-L (1991) Crabs of the China Seas. English edn. China Ocean Press and Springer-Verlag, Beijing and Berlin. Dallwitz MJ, Paine TA, Zurcher EJ (1993 onwards) User’s guide to the DELTA system. A general system for processing taxonomic descriptions. 4.09 edn. CSIRO Division of Entomology, Canberra. . De Grave S, Fransen CHJM (2011) Carideorum Catalogus: the Recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoölogische Mededeelingen 85, 195–589. De Grave S, Pentcheff ND, Ahyong ST, Chan T-Y, Crandall KA, et  al. (2009) A classification of living and fossil genera of decapod crustaceans. Raffles Bulletin of Zoology Supplement 21, 1–109. Derby C, Thiel M (Eds) (2014) Crustacean nervous systems and their control of behavior. The Natural History of the Crustacea Vol. 3. Oxford University Press, New York. Dev Roy MK, Bhadra S (2001) Brachyuran crabs (Crustacea: Decapoda: Brachyura). In Estuarine Ecosystem Series. Vol. 4. pp. 35–54. Zoological Survey of India, Kolkata. Dev Roy MK, Bhadra S (2005) Marine and estuarine crabs (Crustacea: Decapoda: Brachyura). In State Fauna Series 5: Fauna of Andhra Pradesh. Part 5. pp. 357–535. Zoological Survey of India, Kolkata. Dev Roy MK, Bhadra S (2008) Marine and estuarine crabs (Crustacea: Decapoda: Brachyura). In State Fauna Series 16: Fauna of Goa. Part 7. pp. 109–154. Zoological Survey of India, Kolkata. Dev Roy MK, Bhadra S (2011) Brachyuran crabs (Crustacea: Decapoda: Brachyura). In State Fauna Series 17. Fauna of Tamil Nadu. Part 2. pp. 109–269. Zoological Survey of India, Kolkata. Dev Roy MK, Nandi NC (2012) Brachyuran crabs (Crustacea: Decapoda: Brachyura). In State Fauna Series. Vol. 19. Fauna of Andaman and Nicobar islands. Part 1. pp. 185–236. Zoological Survey of India, Kolkata. Emmerson WD (2016) A guide to, and checklist for, the Decapoda of Namibia, South Africa and Mozambique. Volumes 1–3. Cambridge Scholars Publishing, Cambridge. Garth JS (1957) Reports of the Lund University Chile Expedition 1948–49. 29. The Crustacea Decapoda Brachyura of Chile. Lunds Universitets Årsskrift N.F. Avd. 2 53, 1–129, pls 1–4. Haig J (1955) Reports of the Lund University Chile Expedition 1948–49. 20. The Crustacea Anomura of Chile. Lunds Universitets Årsskrift N.F. Avd. 2 51, 1–68, figs 1–12. Hendrickx ME (1993) Crustáceos decápodos del Pacífico mexicano. In Biodiversidad Marina y Costera de Mexico. (Eds Salazar-Vallejo SI, González NE) pp. 271–318. CONABIO, Chetumal. Hendrickx ME, Estrada Navarrete FD (1996) Los camarones pelágicos (Crustacea: Dendrobranchiata y Caridea) del Pacífico Mexicano. Instituto de Ciencias del Mar y Limnología Universidad Nacional Autónoma de Mexico. Holthuis LB (1952) Reports of the Lund Univeristy Chile Expedition 1948–49, 5. The Crustacea Decapoda Macrura of Chile. Lunds Universitets Årsskrift N.F. Avd. 2(47), 1–109. Holthuis LB (1991) FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue of species of interest to

fisheries known to date. FAO Fisheries Synopsis. Vol. 125. Food and Agriculture Organization of the United Nations, Rome. Huxley TH (1880) The crayfish. An introduction to the study of zoology (International Scientific Series Vol. 28). Paul & Co., London. ICZN (1999) International Code of Zoological Nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London. Ingle RW (1980) British crabs. Oxford University Press, Inc., New York. Ingle R (1993) Hermit crabs of the northeastern Atlantic Ocean and the Mediterranea Sea: an illustrated key. Chapman & Hall, London. Ingle RW, Christiansen ME (2004) Synopses of the British Fauna (New series) 55. Lobsters, mud shrimps and anomuran crabs. Keys and notes for the identification of the species. In: Crothers J, Hayward PJ, Published for The Linnean Society of London and The Estuarine and Coastal Sciences Association by Field Studies Council, Shrewsbury. Kaestner A (1980) Invertebrate Zoology. Vol. 3 Crustacea. Reprint edn. Kreiger Publishing Co., New York. Kensley B (1972) Shrimps and prawns of southern Africa. South African Museum, Cape Town. Linnaeus C (1758) Systema Naturae per Regna tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio Decima, reformata. Vol. 1 Animalia. 10th edn. Laurentii Salvii, Holmiae. Manning RB, Holthuis LB (1981) West African brachyuran crabs (Crustacea: Decapoda). Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Martin J (2005) Assembling the Tree of Life (AToL) Decapoda. Crustacea glossary. . Martin JW, Davis GE (2001) An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County Science Series 39, 1–124. McLaughlin PA (1980) Comparative morphology of Recent Crustacea. Freeman and Co., San Francisco. McLaughlin PA, Rahayu DL, Komai T, Chan T-Y (2007) A catalog of the hermit crabs (Paguroidea) of Taiwan. National Taiwan Ocean University, Keelung. McLay CL (1988) Brachyura and crab-like Anomura of New Zealand. Leigh Laboratory Bulletin 22, 1–463. Melo GASd (1996) Manual de identifição dos Brachyura (Caranguejos e siris) do litoral Brasileiro. Editora Plêiade Ltda, Sao Paulo. Melo GASd (1999) Manual de identifição dos Crustacea Decapoda do litoral Brasileiro: Anomura, Thalassinidea, Palinuridea, Astacidea. Editora Plêiade Ltda, São Paulo. Naderloo R (2017) Atlas of crabs of the Persian Gulf. Springer, Switzerland. Ng PKL (1998) Crabs. In FAO Species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 2. Cephalopods, crustaceans, holothurians and sharks. (Eds Carpenter KE, Niem VH) pp. 1045–1155. Food and Agriculture Organization, Rome. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Noël P (1992) Clé préliminaire d’identification des Crustacea Decapoda de France et des principales autres espèces d’Europe. Collections Patrimoines Naturels, série Patrimoine Scientifique (Muséum National d’Histoire Naturelle, Paris) 9, 1–145. Oakley TH, Wolfe JM, Lindgren AR, Zaharoff AK (2012) Phylotranscriptomics to bring the understudied into the fold: monophyletic Ostracoda, fossil placement, and pancrustacean phylogeny. Molecular Biology and Evolution 30, 215–233. Pérez Farfante I, Kensley B (1997) Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175, 1–233.

1 – Introduction

Poore GCB (2004a) Book Review. Tropical Deep Sea Benthos volume 23 (2004) (edited by Bruce A. Marshall and Bertrand Richer de Forges). Muséum National d’Histoire Naturelle - Publications Scientifiques ... Journal of Crustacean Biology 24, 680–681. Poore GCB (2004b) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Poore GCB, Thiel M (Eds) (2020) Evolution and biogeography. The Natural History of the Crustacea Vol. 8. Oxford University Press, New York. Rathbun MJ (1918) The grapsoid crabs of America. Bulletin of the United States National Museum 97, 1–461, pls 1–161. Rathbun MJ (1937) The oxystomatous and allied crabs of America. United States National Museum Bulletin 166, 1–278, pls 1–86. Richer de Forges B, Ahyong S, Castro P, Chan T-Y, Clark PF, et al. (2021) Alain Crosnier’s role in modern carcinology: exploration, international collaboration, and taxonomy. Journal of Crustacean Biology 41, ruab031. doi:10.1093/jcbiol/ruab031 Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo. SCAMIT (2009 onwards) Southern California Association of Marine Invertebrate Taxonomists. Taxonomic Toolbox. . Schram FR (1986) Crustacea. Oxford University Press, New York. Schram FR, Koenemann S (2021) Evolution and phylogeny of Pancrustacea: a story of scientific method. Oxford University Press, New York. Schram FR, Ng PKL (2012) What is Cancer? Journal of Crustacean Biology 32, 665–672. doi:10.1163/193724012X640650 Smaldon G, Holthuis LB, Fransen CHJM (1993) Synopses of the British Fauna (New Series) No. 15. Coastal shrimps and prawns. Keys and notes for identification of the species (Second Edition). In: Barnes RSK, Crothers J, Field Studies Council, Shrewsbury. Spalding M, Fox H, Davidson N, Ferdana Z, Finlayson M, et  al. (2006) Global coastal and marine biogeographic regionalization as a support

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tool for implementation of CBD programmes of work. COP Information Document. Vol. 34. Convention on Biological Diversity, Montreal. Stebbing TRR (1893) A History of Crustacea. Recent Malacostraca. Kegan Paul, Trench, Trübner and C., London. Thiel M, Watling L (Eds) (2015) Lifestyles and feeding biology. The Natural History of the Crustacea Vol. 2. Oxford University Press, New York. Thiel M, Wellborn GA (Eds) (2018) Life histories. The Natural History of the Crustacea Vol. 5. Oxford University Press, New York. UNESCO (2009) Global Open Oceans and Deep Seabed (GOODS)  – ­biogeographic classification. IOC Technical Series 84. UNESCO-IOC, Paris. Vereshchaka AL, Olesen J, Lunina AA (2014) Global diversity and phylogeny of pelagic shrimps of the former genera Sergestes and Sergia (Crustacea, Dendrobranchiata, Sergestidae), with definition of eight new genera. PLoS One 9(11), e112057. doi:10.1371/journal.pone.0112057 von Reumont BM, Edgecombe GD (2020) Crustaceans and insect origins. In The Natural History of the Crustacea. (Eds Poore GCB, Thiel M) pp. 105–120. Oxford University Press, New York. Watling L, Thiel M (Eds) (2013) Functional morphology and diversity. The Natural History of the Crustacea Vol. 1. Oxford University Press, New York. Webber WR, Fenaughty CM, Clark MR (1990) A guide to some common offshore shrimp and prawn species of New Zealand. New Zealand Fisheries Occasional Publication 6, 1–42. Williams AB (1984) Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C. WoRMS Editorial Board (2022) World Register of Marine Species. Available from at VLIZ. Accessed 1 May 2022.

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2. Systematics

Besides being extraordinarily abundant in the sea, crustaceans are amongst the most morphologically diverse of any taxon. More than 67  000 living species are known (Ahyong et  al. 2011), of which more than 51  000 are marine. They are far more varied than insects or spiders. Most are not crabs, shrimps or lobsters but are small, unfamiliar and rarely have common names. Biologists know them as ostracods, copepods, amphipods, cumaceans, isopods, tanaidaceans and a host of other names. Some are illustrated in Fig. 2.1 but unfortunately most will not be dealt with here.

Evolution of the Crustacea For the practical purposes of identification to species, debate over how crustaceans evolved does not matter. Ancestry does matter, however, at lower taxonomic levels. In theory at least, a species’ genus and family placement should tell you something about its ecology or evolutionary history. It is the attempts of taxonomists to reflect this history that induces them to move species from one genus or family to another. The crustaceans have origins in deep time. Definitive crustacean fossils date from ~500 million years ago in the upper Cambrian Orsten fauna of southern Sweden (Walossek and Müller 1998). What the first crustacean looked like has been much debated (for review, see Ahyong 2020; Richter and Wirkner 2020), but subsequent morphological diversification including increasing body size, most notably from the Silurian onward, has resulted in the major branches of the crustacean tree we recognise today  – Oligostraca (including Cephalocarida, Ichthyostraca, Mystacocarida, Ostracoda), Branchiopoda, Remipedia, Hexanauplia and Malacostraca. These major groupings follow differences in the way different parts of the body are specialised and organised, known as tagmatisation. Oligostracans have a short body with a short abdomen (except in the vermiform mystacocarids) and gonopores usually on the fourth thoracic somite. Although appearing very different, the hexanauplians, malacostracans, remipedes and cephalocarids have gonopores positioned on thoracic somites 6–8. The body plan of hexanauplians includes four abdominal somites, whereas malacostracans, apparently without a true abdomen, have the thorax differentiated into a pereon and pleon (according to Hox gene expression; Schram and Koenemann 2004). Remipedes lack trunk regionalisation and have gonopores on the seventh trunk somite in females

and on the fourteenth in males. Cephalocarids and branchiopods both have an elongated thorax and abdomen, but the gonopores are placed on the sixth and twelfth somites, respectively (Ahyong 2020). Any discussion of crustacean evolution must mention the unexpected and controversial finding, first touted by molecular phylogenetics, that insects, far from being distant relatives of crustaceans, were derived from within the ­Crustacea. That is, insects are, technically speaking, crustaceans. Crustaceans were long thought to dominate aquatic habitats but had little success out of the water. Evidently, however, crustaceans successfully colonised the land too, as insects (Glenner et al. 2006). This crustacean-insect clade is variously known as Pancrustacea, Tetraconata, or simply Crustacea (Ahyong 2020; Bracken-Grissom and Wolfe 2020; Richter and Wirkner 2020). The early crustacean fossil record is dominated by ostracods (especially in the Ordovician) whereas malacostracans, despite their Cambrian origins, did not significantly radiate until the Mesozoic. Eumalacostraca, of which Decapoda is a part, continued to actively radiate in the Cenozoic and are now the most ubiquitous and morphologically disparate crustaceans. The decapod fossil record is substantial. Molecular estimates suggest Decapoda arose in the Ordovician (Wolfe et  al. 2019) although the Dendrobranchiata and Stenopodidea first appear in the Late Devonian fossil record and Caridea the Late Triassic (Garassino and Teruzzi 1993; Jones et  al. 2014). Glypheidea, Polychelida and Achelata, first appeared in the Triassic (Audo et al. 2021; Charbonnier et  al. 2013) and clawed lobsters, ‘thalassinidean’ mud shrimps, anomurans and brachyurans followed closely behind. As a result, most major decapod infraorders diverged in the Paleozoic, but major radiations began in the Mesozoic in more or less successive waves with distinct faunal turnover (Schweitzer and Feldmann 2015). According to the fossil record, radiations of the dendrobranchiate shrimps and glypheid lobsters peaked in the Late Triassic with rapid subsequent decline. Polychelidan lobsters were most diverse in the Jurassic. Clawed lobsters peaked in the Mid- Jurassic to early Cretaceous and declined into the Cenozoic. Anomurans and ghost shrimps have remained relatively stable since the Palaeozoic. On the other hand, Brachyura appeared in three successive waves, with the ‘primitive’ podotreme crabs in the Jurassic and Cretaceous, followed by more ‘advanced’ heterotreme and

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Marine Decapod Crustacea

thoracotreme crabs, significantly radiating from the Eocene onwards. Thus, after the earliest decapod origins, the fossil record depicts major Triassic radiations among shrimps and lobsters, moderate ‘activity’ during the Jurassic in the rise of anomurans and podotreme crabs (peaking in the Cretaceous), followed by extensive radiations of heterotreme and thoracotreme crabs starting in the Eocene and which have dominated the decapod record ever since (Schweitzer and Feldmann 2005). Modern distributions of major decapod groups reflect the Mesozoic timing of major radiations. Because of ancient continental movements that ultimately led to the formation of the Atlantic and Pacific ocean basins during the Eocene, most warm-water marine decapod groups now have substantially discrete Indo-West Pacific and Atlanto-East Pacific distributions, with few shared species, some shared genera, and mostly cosmopolitan families. Why is this of interest for an identification guide? The deep history of decapod lineages means that today, most living genera (and their species) occur either in the Indo-West Pacific region or Atlantic Ocean, but not usually both. Some genera are cosmopolitan, but not most. As a result, knowing the natural geographic range of a genus (indicated in this work) can assist in corroborating or correcting identifications made using the keys. So, if you find a specimen from Brazil that belongs to a genus that is otherwise known only from the Indo-Pacific, be sure to double-check the identification – it could be incorrect, an introduced species or perhaps a new scientific discovery.

Classification of the Crustacea Table 2.1 reflects one hierarchy of all the major groups of Crustacea where it can be seen that the Decapoda is just one order among many. This hierarchy of groups within groups is just the latest of several and is subject to change as more data from morphology and DNA become available. The Subphylum Crustacea is divided into nine classes, of which four belong to the superclass Allotriocarida, three to Oligostraca and two to Multicrustacea (Bracken-Grissom and Wolfe 2020; Oakley et al. 2012; Regier et al. 2010): Branchiopoda  – despite a diversity of body forms branchiopods typically share a series of uniform, phyllopodous trunk limbs and a limbless abdomen (if present). Many branchiopods live in fresh water and only the Cladocera, popularly known as water fleas, have marine representatives (Olesen 2009). Cephalocarida  – microscopic thin animals with a broad, shield-like cephalon, followed by a thorax of eight segments

and an abdomen of twelve, of which the last abdominal segment bears paired caudal rami. Cephalocarids live between the sand grains on beaches and marine sediments (Hessler and Elofsson 2013). Remipedia – an unusual group of many-legged centipedelike predators, that, uniquely among crustaceans, produce venom delivered through the maxilla. They are confined to fresh water in caves and anchialine aquifers (Koenemann et al. 2003). Hexapoda – insects. Ichthyostraca  – includes two groups of parasitic crustaceans. Branchiura are external parasites of fishes and amphibians (Moller 2009) while Pentastomida, tongue worms, are lung parasites of vertebrates, especially reptiles (Christoffersen and Assis 2015). Mystacocarida – a group of vermiform interstitial crustaceans from beaches (Boxshall and Defaye 1996) Ostracoda – sometimes called seed shrimps, a group rich in species in marine, freshwater and terrestrial environments. They have a bivalved carapace and are well represented as fossils. Relationships among the two major groups are unresolved (Wolfe and Hegna 2014). The class Hexanauplia includes the copepods and barnacles plus another enigmatic group. Copepods are diverse and abundant in marine environments with diverse pelagic and benthic forms (Boxshall and Halsey 2004; Huys and Boxshall 1991). Many copepods are parasitic or symbiotic. Likewise, the subclass Thecostraca includes a wide range of attached, boring and parasitic barnacles (Chan et al. 2020; Høeg 1995; Newman 1996) The last class Malacostraca includes the most familiar crustaceans. They are variously arranged in two or three subclasses and 19 extant orders. One subclass includes only the Leptostraca (WalkerSmith and Poore 2001). The second subclass, Hoplocarida, includes Stomatopoda, mantis shrimps (Ahyong and Jarman 2009) and has variously been treated as a separate subclass or included within the largest subclass, Eumalacostraca (Ahyong 2020; Richter and Scholtz 2001). All eumalacostracans have 20 body segments (all except the first with paired limbs) plus a terminal telson. The body segments, or somites, are arranged into three sections, or tagmata, head or cephalon, thorax (or pereon) and pleon (or abdomen). The head has eight pairs of limbs involved in sensation and feeding, the thorax eight pairs usually locomotory and variously involved with feeding, and the

2 – Systematics

Fig 2.1.  Representatives of some basic crustacean types. a, Copepoda. b, Ostracoda. c, Anostraca. d, Notostraca. e, Onchycaudata. f, Leptostraca. g, Anaspidacea. h, Cirripedia. i, Stomatopoda. j, Mysida. k, Tanaidacea. l, Isopoda. m, Amphipoda. n, Euphausiacea. o, Dendrobranchiata. p, Caridea. q, Astacidea. r, Anomura. s, Achelata. t, Gebiidea. u, Brachyura. v, Polychelida.

11

Branchiopoda

Cephalocarida Remipedia Hexapoda Ichthyostraca

Mystacocarida Ostracoda Hexanauplia

Allotriocarida

Oligostraca

Multicrustacea

Malacostraca

Class

Superclass

Thecostaca/ Facetotecta Eumalacostraca

4 subclasses Copepoda Tantulocarida Thecostaca / Ascothoracica Thecostaca / Cirripedia

Branchiura Pentastomida

Sarcostraca

Phyllopoda

Subclass/ Infraclass

Eucarida

Thoracica

Acrothoracica

Superorder

Euphausiacea Decapoda

5 orders

2 orders

10 orders

Notostraca Anostraca Brachypoda Nectipoda

Diplostraca

Order

Dendrobranchiata Pleocyemata

Laevicaudata Onchycaudata

Suborder

Table 2.1.  Classification of living members of the subphylum Crustacea (Decapoda covered in this volume shaded)

Axiidea Brachyura Caridea Gebiidea Glypheidea Polychelida Procarididea Stenopodidea

Astacidea

Achelata Anomura

Infraorder

krill prawns spiny lobsters, bugs hermit crabs, king crabs, mole crabs, porcelain crabs, squat lobsters clawed lobsters, crayfish, yabbies ghost and sponge shrimps true crabs shrimps mud and sponge lobsters glypheid lobsters deep-sea blind lobsters shrimps coral shrimps

parasitic barnacles goose barnacles, wart barnacles, acorn barnacles y-larvae

boring barnacles

mystacocarids seed shrimps, ostracods copepods tantulocarids ascothoracican barnacles

clam shrimps water fleas shield shrimps fairy shrimps cephalocarids remipedes insects fish lice tongue worms

Common name

12 Marine Decapod Crustacea

Superclass

Class

Phyllocarida

Subclass/ Infraclass

Hoplocarida

spelaeogriphaceans cave mysids thermosbaenaceans tanaidaceans

Stygiomysida Thermosbaenacea Tanaidacea Leptostraca

leptostracans, nebaliaceans

mantis shrimps

opposum shrimps

Spelaeogriphacea

Stomatopoda

mictaceans

Mysida

2 suborders

lophogastrids

Isopods, slaters, woodlice

Isopoda Mictacea

ingolfiellids 11 suborders

comma cumaceans

Ingolfiellida Lophogastrida

hirsutiids

Amphipods, skeleton shrimps, whale lice

Cumacea

6 suborders

Bochusacea

Amphipoda

mountain shrimps

Peracarida

bathynellaceans

Anaspidacea

Common name

Bathynellacea

Infraorder

Syncarida

Suborder

Order

Superorder

2 – Systematics 13

14

Marine Decapod Crustacea

pleon has six pairs of limbs involved with respiration, reproduction and swimming. The mantis shrimps, superorder Hoplocarida, are marine, mostly benthic predators that are most common on tropical shores. They are recognised by their short, shield-like carapace, triflagellate antennules and their trademark adaptations  – praying mantis-like raptorial claws and highly complex eyes (Ahyong and Jarman 2009; Van Der Wal et al. 2017). The superorder Syncarida is mostly from fresh water and includes Anaspidacea or mountain shrimp (Ahyong 2016) and more cryptic cave and interstitial forms in the Bathynellacea (Camacho et al. 2018a, b). The superorder Peracarida includes 12 orders of small benthic or epibenthic crustaceans. Their relationships have not been agreed on (Poore 2005; Wilson 2009). Some are exceptionally abundant in marine environments. The diverse and common Amphipoda were separated from the rarer Ingolfiellida by Lowry and Myers (2017). Cumacea, unlike many other crustacean groups, are taxonomically stable although relationships between the families are debated (Haye et al. 2004). Isopoda include woodlice, one of the few large groups to have colonised terrestrial environments (Sfenthourakis et al. 2020), and take many forms in marine and fresh water (Brandt and Poore 2003). Lophogastrida, Mysida and Stygiomysida, epipelagic or mesopelagic shrimps, were once united but now treated as three orders (Meland et  al. 2015; Meland and Willassen 2007). Tanaidacea too are abundant but poorly known (Błażewicz-Paszkowycz et  al. 2012). Their relationships have begun to be addressed (Bird and Larsen 2009; Drumm 2010; Larsen and Wilson 2002). The remaining peracaridan orders contain few species and with the exception of the last are rare: Bochusacea, Mictacea, Spelaeogriphacea (Poore 2015) and Thermosbaenacea (Wagner 1994). The superorder Eucarida is characterised by a carapace – the shield or shell encasing the segments of the head and thorax in such a way that the segmentation is concealed. In most other crustaceans many of the segments are quite visible. All eucarids have eyes on movable stalks. Eucarids comprise two orders, Euphausiacea or krill (Baker et  al. 1990; Jarman 2001; Mauchline 1980), pelagic crustaceans famous for their role in the diet of whales, and Decapoda. It is only the last that features in this guide.

Classification of the Decapoda Decapods differ from other eucarids in having the first three pairs of thoracic limbs modified. The modified limbs, called maxillipeds, are feeding appendages. Consequently, only the last five pairs of thoracic limbs are involved in locomotion. These five pairs give the group its name, Decapoda, meaning ten-footed. The ten legs refer only to the claws and walking legs. The monophyly of the Decapoda is without doubt but the arrangement of taxonomic groups within the order has been debated over more than a century (Ahyong and O’Meally 2004; Bracken et al. 2009; Chu et al. 2009; Dixon et al. 2003; Martin et al. 2009; Porter et al. 2005; Toon et al. 2009; Wolfe et  al. 2019). The names used for these subdivisions have changed considerably (Poore 2016). This guidebook adopts the widely held classification into two suborders, Dendrobranchiata and Pleocyemata, differentiated on the structure of the gills and male pleopods 1 (Burkenroad 1981). Dendrobranchiata (diagnosed in detail below) have trichobranchiate gills and the male pleopods 1 joined to form a ‘petasma’. Pleocyemata are diagnosed as having phyllobranchiate or sometimes trichobranchiate gills and the endopods of the male pleopods 1 (if present) independent. Pleocyemata are divided into infraorders, the number now stabilised at eleven. Four recent changes are notable. The recognition of the differences between the two ‘thalassinidean’ infraorders Axiidea and Gebiidea was first proposed on morphological criteria (de Saint Laurent 1979; Sakai 2005) and later corroborated by molecular studies (Robles et  al. 2009). Similarly, exclusion of procarididean shrimps from Caridea (Felgenhauer and Abele 1983) was later confirmed by genetics (Bracken et  al. 2009; Felgenhauer and Abele 1983; Wolfe et al. 2019). Doubts about the monophyly of the ‘Palinura’ were expressed by Scholtz and Richter (1995) and Schram and Ahyong (2002). In Scholtz and Richter’s (1995) phylogeny, Palinura is not recognised as such but comprise at least two clades not closely related, Polychelida and Achelata. Likewise, Dixon et al. (2003) and Ahyong and O’Meally (2004) showed that the mostly extinct glypheoid lobsters also belonged in a different infraorder, Glypheidea, quite separate from the palinurans. This break-up of the old Palinura was subsequently corroborated by extensive molecular analysis (Bracken-Grissom et al. 2014).

Key to suborders and infraorders of Decapoda and shrimp-like marine crustaceans A key to differentiate the traditional suborders and infraorders of Decapoda may seem unnecessary but is included as a quick critical guide. Euphausiacean, lophogastrids and mysids are not decapods but are sufficiently shrimp-like to sometimes be confused. Exceptions could be found for most couplets, for example, secondarily chelate or non-chelate limbs appear in most orders.

2 – Systematics

1.

15

Carapace short, exposing feathery thoracic gills. Thoracic appendages similar, or last 1 or 2 lost, or first few modified. Pelagic, shrimp-like������������������������������������������������������������������������������������������������������ Euphausiacea – Carapace usually covering all gills (if short, gills absent). Thoracic appendages 1 or 1–3 modified as maxillipeds. Pelagic or benthic�������������������������������������������������������������������������������������������������������������������������������������������������2 2. Shrimp-like. With 1 pair of maxillipeds, 6 or 7 pairs of biramous thoracic swimming appendages. Females with thoracic oostegites (plates between legs and holding eggs)�����������������������������������������������������������������������������������������3 – Shrimp-, lobster- or crab-like. Thoracic appendages 1–3 modified as maxillipeds, 4–8 modified as pereopods 1–5. Females without oostegites������������������������������������������������������������������������������������������������������������� Decapoda … 4 3. Carapace with clear longitudinal keels and/or spines, covering all thoracic somites; with dominant triangular rostrum, extending well beyond eyes. Uropod without statocysts������������������������������������������ Lophogastrida – Carapace usually unornamented, often not covering posterior thoracic somite; with short rostrum, at most, extending little beyond eyes. Uropod with statocysts at base of endopods (swollen cavities containing granules)������������������������������������������������������������������������������������������������������������������������������������������Mysida 4. Prawn-like. Gills with branching fine filaments (trichobranchiate). Male pleopods 1 endopods joined to form petasma. Females never carrying egg masses on pleopods�������������������������� Dendrobranchiata … p. 19 – Shrimp-, lobster- or crab-like. Gills with stacked plate-like branches (phyllobranchiate or sometimes trichobranchiate). Male pleopods 1 endopods (if present) independent. Ovigerous females carrying egg masses on pleopods��������������������������������������������������������������������������������������������������Pleocyemata … 5 5. Crabs. Pereopod 1 chelate, usually greatly enlarged, pereopods 2–5 as walking legs or flat swimming legs, usually simple. Pleon short, folded under carapace. Uropod usually absent��������������������������� Brachyura… p. 421 – Shrimps, lobsters, or secondarily crab-like. Pereopods 1–5 variously chelate or simple. Pleon long, muscular or, if short, not usually engaging closely with underside of carapace. Uropod usually present�������������������������������6 6. Pereopod 3 longest��������������������������������������������������������������������������������������������������������������������������������������� Stenopodidea… p. 51 – Pereopods 1 or 2 longest������������������������������������������������������������������������������������������������������������������������������������������������������������������7 7. Deep-sea blind lobsters. Pereopods 1–4 or 1–5 chelate, first longest, thin. Carapace flat������������������Polychelida… p. 259 – Lobsters, shrimps or crabs. With 0–3 pairs of chelate limbs (if other limbs chelate of different form)�����������������������������8 8. Clawed lobsters, freshwater crayfish. Pereopods 1–3 chelate, first greatly enlarged (fifth sometimes also minutely chelate)���������������������������������������������������������������������������������������������������Astacidea … p. 175 – With fewer than 3 pairs of chelate pereopods������������������������������������������������������������������������������������������������������������������������������9 9. Without chelate pereopods�����������������������������������������������������������������������������������������������������������������������������������������������������������10 – With 1 or 2 pairs of chelate pereopods��������������������������������������������������������������������������������������������������������������������������������������� 12 10. Shrimps from anchialine or similar environments, or rock pools. Exoskeleton soft, smooth, chitinised. Pereopods with exopod��������������������������������������������������������������������������������������������������������������������������� Procarididea ... p. 59 – Lobsters from marine environments. Exoskeleton hard, spiny or rough, calcified. Pereopods without exopod�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������11 11. Rock lobsters, slipper lobsters. Basal antennular and antennal articles fused to epistome. Thoracic sternite 8 fused to sternal plastron������������������������������������������������������������������������������������������� Achelata … p. 247 – Basal antennular and antennal articles free, not fused to epistome. Thoracic sternite 8 fused articulating, not fused to sternal plastron�����������������������������������������������Glypheidea … p. 183 12. Shrimps. Thoracic sternites 1–8 fused. Pereopods 1, 2 or both chelate, second often strongest�������������� Caridea … p. 61 – Thoracic sternite 8 free from sternites 1–7. Pereopod 1 or 1+2 chelate, first pair much stronger than pereopods 2–5���������������������������������������������������������������������������������������������������������������������������������������������������13 13. Hermit-crab or crab-like. Pleon coiled or flattened. Pereopod 5 greatly reduced, held folded, not used for locomotion�����������������������������������������������������������������������������������������������������������������������������Anomura … p. 263 – Lobster-like. Pleon elongate, muscular. Pereopod 5 slightly reduced, used for locomotion����������������������������������������������14 14. Pereopods 1, 2 chelate, first a large cheliped��������������������������������������������������������������������������������������������������� Axiidea … p. 185 – Pereopod 1 a large chelate or subchelate cheliped; pereopod 2 subchelate or simple����������������������������� Gebiidea … p. 237

16

Marine Decapod Crustacea

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2 – Systematics

Islands. Contributions to Zoology 72, 227–252. doi:10.1163/1875986607204004 Larsen K, Wilson GDF (2002) Tanaidacean phylogeny, the first step: the superfamily Paratanaidoidea. Journal of Zoological Systematics and Evolutionary Research 40, 205–222. doi:10.1046/j.1439-0469.2002.​00193.x Lowry J, Myers A (2017) A phylogeny and classification of the Amphipoda with the establishment of the new order Ingolfiellida (Crustacea: Peracarida). Zootaxa 4265, 1–89. doi:10.11646/zootaxa.4265.1.1 Martin JW, Crandall KA, Felder DL (Eds) (2009) Decapod Crustacean Phylogenetics.Crustacean Issues Vol. 18. CRC Press, Taylor & Francis Group, Boca Raton, London, New York. Mauchline J (1980) The biology of mysids and euphausiids. Advances in Marine Biology 18, 1–680. Meland K, Willassen E (2007) The disunity of “Mysidacea” (Crustacea). Molecular Phylogenetics and Evolution 44, 1083–1104. doi:10.1016/j. ympev.2007.02.009 Meland K, Mees J, Porter M, Wittmann KJ (2015) Taxonomic review of the orders Mysida and Stygiomysida (Crustacea, Peracarida). PLoS One 10, e0124656. doi:10.1371/journal.pone.0124656 Moller OS (2009) Branchiura (Crustacea) – survey of historical literature and taxonomy. Arthropod Structure & Development 67, 41–55. Newman WA (1996) Sous-classe des Cirripèdes (Cirripedia Burmeister, 1834) Super-orders des Thoraciques et des Acrothoraciques (Thoracica Darwin, 1854  – Acrothoracica Gruvel, 1905). In Traité de Zoologie sous la direction de P.-P. Grassé. (Ed. Forest J) pp. 453–540. Masson éditeur, Paris. Oakley TH, Wolfe JM, Lindgren AR, Zaharoff AK (2012) Phylotranscriptomics to bring the understudied into the fold: monophyletic Ostracoda, fossil placement, and pancrustacean phylogeny. Molecular Biology and Evolution 30, 215–233. Olesen J (2009) Phylogeny of Branchiopoda (Crustacea)  – character evolution and contribution of uniquely preserved fossils. Arthropod Systematics & Phylogeny 67, 3–39. Poore GCB (2005) Peracarida: monophyly, relationships and evolutionary success. Nauplius 13, 1–27. Poore GCB (2015) Chapter 53. Orders Bochusacea, Mictacea and Spelaeogriphacea. In Treatise on Zoology  – Anatomy, Taxonomy, Biology. Revised and updated, as well as extended from the Traité de Zoologie. (Eds von Vaupel Klein JC, Charmantier-Duares M, Schram FR) pp. 77–92. Brill, Leiden. Poore GCB (2016) The names of the higher taxa of Crustacea Decapoda. Journal of Crustacean Biology 36, 248–255. Porter ML, Pérez-Losada M, Crandall KA (2005) Model-based multilocus estimation of decapod phylogeny and divergence times. Molecular Phylogenetics and Evolution 37, 355–369. doi:10.1016/j. ympev.2005.06.021 Regier JC, Shultz JW, Zwick A, Hussey A, Ball B, et al. (2010) Arthropod relationships revealed by phylogenomic analysis of nuclear proteincoding sequences. Nature 463, 1079–1083. doi:10.1038/nature08742 Richter S, Scholtz G (2001) Phylogenetic analysis of the Malacostraca (Crustacea). Journal of Zoological Systematics and Evolutionary Research 39, 113–136. doi:10.1046/j.1439-0469.2001.00164.x Richter S, Wirkner CS (2020) What the ur-crustacean looked like. In The Natural History of the Crustacea. (Eds Poore GCB, Thiel M) pp. 1–20. Oxford University Press, New York.

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Robles R, Tudge CC, Dworschak PC, Poore GCB, Felder DL (2009) Molecular phylogeny of the Thalassinidea based on nuclear and mitochondrial genes. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 309–326. CRC Press, Boca Raton. Sakai K (2005) The diphyletic nature of the infraorder Thalassinidea (Decapoda, Pleocyemata) as derived from the morphology of the gastric mill. Crustaceana 77, 1117–1129. doi:10.1163/1568540042900268 Scholtz G, Richter S (1995) Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca). Zoological Journal of the Linnean Society 113, 289–328. doi:10.1006/zjls.1995.0011 Schram FR, Ahyong S (2002) The higher affinities of Neoglyphea inopinata in particular and the Glypheoidea (Decapoda, Reptantia) in general. Crustaceana 75, 629–635. doi:10.1163/156854002760095651 Schram FR, Koenemann S (2004) Are the crustaceans monophyletic? In Assembling the tree of life. (Eds Cracraft J, Donaghue MJ) pp. 319–329. Oxford University Press, New York. Schweitzer CE, Feldmann RM (2005) Decapod crustaceans, the K/P event, and paleocene recovery. In: Koenemann S, and Vonk R (Eds), Crustacea and arthropod relationships. Crustacean Issues 16, 17–53. Schweitzer CE, Feldmann RM (2015) Faunal turnover and niche stability in marine Decapoda in the Phanerozoic. Journal of Crustacean Biology 35, 633–649. doi:10.1163/1937240X-00002359 Sfenthourakis S, Myers AA, Taiti S, Lowry JK (2020) Terrestrial environments. In The Natural History of the Crustacea. (Eds Poore GCB, Thiel M) pp. 359–388. Oxford University Press, New York. Toon A, Finley M, Staples J, Crandall KA (2009) Decapod phylogenetics and molecular evolution. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 15–29. CRC Press, Boca Raton. Van Der Wal C, Ahyong ST, Ho SYW, Lo N (2017) The evolutionary history of Stomatopoda (Crustacea: Malacostraca) inferred from molecular data. PeerJ 5, e3844. doi:10.7717/peerj.3844 Wagner HP (1994) A monographic review of the Thermosbaenacea (Crustacea: Peracarida). Zoölogische Verhandelingen 291, 1–338. Walker-Smith GK, Poore GCB (2001) A phylogeny of the Leptostraca (Crustacea) with keys to the families and genera. Memoirs of Museum Victoria 58, 383–410. doi:10.24199/j.mmv.2001.58.21 Walossek D, Müller KJ (1998) Cambrian ‘Orsten’-type arthropods and the phylogeny of the Crustacea. In Arthropod relationships. The Systematics Association Special Volume Series 55. (Eds Fortey RA, Thomas RH) pp. 139–154. Chapman and Hall, London. Wilson GDF (2009) The phylogenetic position of the Isopoda in the Peracarida (Crustacea: Malacostraca). Arthropod Systematics & Phylogeny 67, 159–198. Wolfe JM, Hegna TA (2014) Testing the phylogenetic position of Cambrian pancrustacean larval fossils by coding ontogenetic stages. Cladistics 30, 366–390. doi:10.1111/cla.12051 Wolfe JM, Breinholt JW, Crandall KA, Lemmon AR, Lemmon EM, et  al. (2019) A phylogenomic framework, evolutionary timeline and genomic resources for comparative studies of decapod crustaceans. Proceedings. Biological Sciences 286, 20190079. doi:10.1098/rspb.2019.0079

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3. Dendrobranchiata – prawns and shrimps

This chapter deals with Dendrobranchiata Bate, 1888, one of the two major divisions of the Decapoda – all other chapters treat sections of the other division, Pleocyemata. The Dendrobranchiata, therefore, are fundamentally different from all the others anatomically and ecologically. The essential anatomical character is found in the gills (branchiae) under the sides of the carapace. In dendrobranchiates, the gills, when present, are ‘dendrobranchiate’ or composed of many branching filaments attached to a major stem. In pleocyemate decapods, crabs, shrimps, lobsters etc., the gills look like a stack of plates in a row along a main axis. The usual dendrobranchiate reproductive strategy also differs from that of the Pleocyemata. Eggs are released from the female and develop in the plankton until their time to settle. One family, Luciferidae, is exceptional in that the eggs are brooded by the female on pereopod 3. Crabs and their relatives brood their eggs by attaching them to setae on the pleopods until they hatch as zoeal larvae. Dendrobranchiate shrimps, or prawns as edible penaeids are known in Australia, superficially look like caridean shrimps and in midwater and inshore benthic samples, examples of both often occur together. Carideans, besides having plate-like gills, differ from prawns in having the second pleonal pleuron (plate extending ventrally to shelter the pleopods) covering both the pleuron in front and behind. In dendrobranchiates, the pleura overlap like tiles, the first over the second, second over the third and so on. Also, dendrobranchiates have three pairs of claws (pereopods 1–3 chelate, although minutely so) while carideans usually have two pairs of chelate pereopods, the second often enlarged as claws. If you observe an egg mass carried under the pleon, your specimen is not a dendrobranchiate. Burkenroad’s (1983) discussion of a phylogeny of the Dendrobranchiata is responsible for the current classification into superfamilies. The ranks of some of his taxa have changed so his keys to families and genera have been superseded. His earlier work (Burkenroad 1963) was responsible for the separation of dendrobranchiates from the Pleocyemata, all other decapods and the abandoning of divisions between natant (swimming) and reptant (walking) taxa. This division has never been questioned and the most recent molecular analyses of the phylogeny of Decapoda assumes their monophyly (Tsang et  al. 2008). Two superfamilies are recognised.

– Sergestoidea lack pleurobranchs, never having more than two branchiae on each thoracic somite and eight per side. This describes one family, Sergestidae, but the other family, Luciferidae, lacks gills of any sort. – Penaeoidea have at least 11 branchiae on each side and usually pleurobranchs on some thoracic somites, some thoracic somites with at least three branchiae. Penaeoidea comprise five families. This division was tested by Tavares et al. (2009) using a parsimony analysis of morphological data. They concluded that the superfamilies and most families were monophyletic, although some lacked clear synapomorphies. They failed to demonstrate the monophyly of Penaeidae and Benthesicymidae but considered their results ‘preliminary’ because few species were used to represent each family. Penaeidae was also found to be paraphyletic and to have a close relationship to Solenoceridae by Hsiao and Lai (2006) and by Vazquez-Bader et  al. (2004) who also found Aristeidae, Benthesicymidae and Sicyoniidae to be closely associated. These findings were doubted by Chan et  al. (2008) who reviewed these and other molecular studies. Other studies supported the monophyly of Solenoceridae, Aristeidae, Benthesicymidae and Sicyoniidae but found these families nested within a paraphyletic Penaeidae (Chan 2012; Cheng et  al. 2018; Ma et  al. 2009). Robalino et al. (2016) went further and divided Penaeoidea into two clades: Phorcysida comprising Solenoceridae, Aristeidae and Benthesicymidae, with a marked preference for deep water; and Penaeidae (incorporating Sicyoniidae) with preference for shallow waters. Robalino et  al. (2016) collated considerable data on morphology, body size, depth range and behaviour. They were able to demonstrate that penaeids suitable for aquaculture are larger than others and that this large size is a synapomorphy of Penaeus s.l. De Grave and Fransen (2011) listed all species known then with synonyms and type localities. Their higher classification is followed here but not all the genera they recognised are accepted here. They catalogued 68 genera and 533 species but more have been added since. Identification of dendrobranchiates world-wide is facilitated by the well illustrated and explained keys to families and genera of Pérez Farfante and Kensley (1997). They also diagnosed all genera in detail, figured a representative of each, and listed all species with their distributions known at

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Marine Decapod Crustacea

that time. This chapter draws heavily on their treatment. While their family treatment is agreed on by most modern authors, there is less sympathy for their genera of penaeids (see under Penaeus below). Other keys are available, especially for species of commercial importance (Burkenroad 1983; Burukovskii 1983; Dore and Frimodt 1987; Holthuis 1980). The key to families below is a practical one and does not follow the division into superfamilies. Diagnosis. Carapace more or less cylindrical, longer than wide; epistome short, not fused to basal antennular and

antennal articles, lateral margins with narrow point of contact with carapace margins. Exoskeleton chitinised. Thoracic sternites 1–8 fused. Pleon elongate, muscular; pleura not over overlapping or overlapping next most posterior pleuron. Gills dendrobranchiate. Antennular penduncle articles aligned linearly, main flagellum long, lash-like, aesthetascs more-or-less evenly distributed. Pereopods 1–3 similar, usually chelate. Pereopod 5 of similar length to more anterior pereopods, longer than or reduced, or absent. Male pleopods 1 endopods joined to form petasma.

Key to families of Dendrobranchiata 1. – 2. – 3. – 4. – 5. – 6. –

Without branchiae. Body extremely thin, laterally compressed, anteriorly elongate (Fig. 3.8)�������������������������Luciferidae With branchiae. Body typically shrimp-like��������������������������������������������������������������������������������������������������������������������������������2 With no more than 2 branchiae per thoracic somite, at most 8 on each side (Figs 3.9–3.11)�����������������������������Sergestidae With at least 3 branchiae on some thoracic somites, at least 11 on each side�������������������������������������������������������������������������3 Rostrum short, triangular, with 1 or few postrostral spines (Fig. 3.2)�������������������������������������������������������� Benthesicymidae Rostrum elongate, blade-like with dorsal and sometimes ventral teeth (Figs 3.1, 3.4–3.7)��������������������������������������������������4 Pleopods 3–5 uniramous, lacking endopods. Integument rigid and stony (Fig. 3.6)����������������������������������������� Sicyoniidae Pleopods 3–5 biramous. Integument not rigid and stony����������������������������������������������������������������������������������������������������������5 Postorbital spine present (Fig. 3.7)��������������������������������������������������������������������������������������������������������������������������Solenoceridae Postorbital spine absent�������������������������������������������������������������������������������������������������������������������������������������������������������������������6 Prosartesma (mesial process on antennule article 1) well developed (Fig. 3.3b)����������������������������������������������������Penaeidae Prosartesma reduced to setose boss (Fig. 3.1e)������������������������������������������������������������������������������������������������������������ Aristeidae

References Burkenroad MD (1963) The evolution of the Eucarida (Crustacea, Eumalacostraca) in relation to the fossil record. Tulane Studies in Geology 2, 3–16. Burkenroad MD (1983) Natural classification of Dendrobranchiata, with a key to Recent genera. In: Schram FR (Ed.), Crustacean phylogeny. Crustacean Issues 6, 279–290. Burukovskii RN (1983) Key to shrimps and lobsters. (Translation of 1974 Russian edn) AA Balkema, Rotterdam. Chan T-Y (2012) A new genus of deep-sea solenocerid shrimp (Decapoda: Penaeoidea) from Papua New Guinea. Journal of Crustacean Biology 32, 489–495. doi:10.1163/193724012X626557 Chan T-Y, Tong J, Tam YK, Chu KH (2008) Phylogenetic relationships among the genera of the Penaeidae (Crustacea: Decapoda) revealed by mitochondrial 16S rRNA gene sequences. Zootaxa 1694, 38–50. doi:10.11646/zootaxa.1694.1.2 Cheng J, Chan T-Y, Zhang N, Sun S, Sha Z-l (2018) Mitochondrial phylogenomics reveals insights into taxonomy and evolution of Penaeoidea (Crustacea: Decapoda). Zoologica Scripta 47, 582–594. doi:10.1111/ zsc.12298 De Grave S, Fransen CHJM (2011) Carideorum Catalogus: the Recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoölogische Mededeelingen 85, 195–589. Dore I, Frimodt C (1987) An illustrated guide to shrimp of the world. Van Nostrand Reinhold Co., New York. Holthuis LB (1980) FAO species catalogue. Vol. 1. Shrimps and prawns of the world. An annotated catalogue of species of interest to fisheries. FAO Fisheries Synopsis 125, 1–261.

Hsiao S-T, Lai J-L (2006) Phylogenetic analyses of economically important shrimp species in Taiwan based on DNA sequences of mitochondrial 16S rRNA. Journal of Taiwan Fisheries Research 14, 67–73. Ma KY, Chan T-Y, Chu KH (2009) Phylogeny of penaeoid shrimps (Decapoda: Penaeoidea) inferred from nuclear protein-coding genes. Molecular Phylogenetics and Evolution 53, 45–55. doi:10.1016/j. ympev.2009.05.019 Pérez Farfante I, Kensley B (1997) Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175, 1–233. Robalino J, Wilkins B, Bracken-Grissom HD, Chan T-Y, O’Leary MA (2016) The origin of large-bodied shrimp that dominate modern global aquaculture. PLoS One 11, e0158840. doi:10.1371/journal. pone.0158840 Tavares C, Serejo CS, Martin JW (2009) A preliminary phylogenetic analysis of the Dendrobranchiata based on morphological characters. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 261–279. CRC Press, Boca Raton. Tsang LM, Ma KY, Ahyong ST, Chan T-Y, Chu KH (2008) Phylogeny of Decapoda using two nuclear protein-coding genes: Origin and evolution of the Repantia. Molecular Phylogenetics and Evolution 48, 359–368. doi:10.1016/j.ympev.2008.04.009 Vazquez-Bader AR, Carrero JC, Garcia-Varela M, Gracia A, Laclette JP (2004) Molecular phylogeny of Superfamily Penaeoidea RafinesqueSchmaltz, 1815, based on mitochondrial 16S partial sequence analysis. Journal of Shellfish Research 23, 911–917.

3 – Dendrobranchiata – prawns and shrimps

Superfamily Penaeoidea Rafinesque-Schmaltz, 1815 Five families are included: Aristeidae, Benthesicymidae, Penaeidae, Sicyoniidae and Solenoceridae. Diagnosis. At least 11 branchiae on each side and usually pleurobranchs on some thoracic somites, some thoracic somites with at least 3 branchiae.

Aristeidae Wood-Mason in Wood-Mason & Alcock, 1891 Figure 3.1, Plate 2 Aristeid shrimps are deep-sea demersal or mesopelagic species found off the edge of the continental shelf at depths below 300 m. None occurs in shallow water. Two widespread species in particular, Aristaeomorpha foliacea and Aristeus antennata are of commercial interest because of their large size, up to

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350 mm length, and high local densities (Cau et al. 2002; Pezzuto et al. 2006). Others are exploited locally (Holthuis 1980). Some species may be confused superficially with oplophorid shrimps but can be separated on the dendrobranchiate gills and pleon. A key character distinguishing the family from other prawns is the disparity between the lengths of flagella of the antennule. Crosnier (1978) provided keys and detailed descriptions and figures of several species. Pérez Farfante and Kensley (1997) diagnosed the family and genera in more detail than given here, listed all species known at the time with distributions. Diagnosis. Body typically shrimp-like; integument soft, stiff but not rigid. Rostrum elongate, blade-like with dorsal and sometimes ventral teeth. Postorbital spine absent; with 3 postrostral teeth, or with 4 or more postrostral teeth. With at least 3 branchiae on some thoracic somites, at least 11 on each side. Prosartesma (mesial process on antennule article 1) reduced to setose boss (Fig. 3.1e).

Key to genera of Aristeidae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. –

With 5 or more rostral-postrostral teeth (Fig. 3.1a)�����������������������������������������������������������������������������������������Aristaeomorpha With 3 rostral-postrostral teeth (Fig. 3.1b–j)�������������������������������������������������������������������������������������������������������������������������������2 Postantennal spine present (anterior to gastro-orbital carina) (Fig. 3.1h)��������������������������������������������������� Parahepomadus Postantennal spine absent���������������������������������������������������������������������������������������������������������������������������������������������������������������3 Pereopods 1–5 without exopods�����������������������������������������������������������������������������������������������������������������������������������������������������4 Pereopods 1–5 with exopods (Fig. 3.1k)����������������������������������������������������������������������������������������������������������������������������������������7 Hepatic spine present (Fig. 3.1g)���������������������������������������������������������������������������������������������������������������������������������Hepomadus Hepatic spine absent (Figs 3.1b–d, f, i, j)���������������������������������������������������������������������������������������������������������������������������������������5 Pereopods 1 and 2 merus without movable spine��������������������������������������������������������������������������������������������������� Aristaeopsis Pereopods 1 and 2 merus with small distomesial movable spine���������������������������������������������������������������������������������������������6 Postcervical sulcus present dorsally (Fig. 3.1j)�������������������������������������������������������������������������������������������������������Pseudaristeus Postcervical sulcus visible only laterally, absent dorsally (Fig. 3.1c)��������������������������������������������������������������������������� Aristeus Pereopods 1 and 2 merus without movable spine�������������������������������������������������������������������������������������������������Hemipenaeus Pereopods 1 and 2 merus with small distomesial movable spine���������������������������������������������������������������������������������������������8 Carapace ridges and carinae weak. Pereopod 3 podobranch rudimentary. Male antennular lower flagellum modified (Fig. 3.1d, e)�����������������������������������������������������������������������������������������������������������������������������������������Austropenaeus Carapace ridges and carinae strong (Fig. 3.1i). Pereopod 3 podobranch large. Male antennular flagella similar�������������������������������������������������������������������������������������������������������������������������������������������������������������������������� Cerataspis Aristaeomorpha Wood-Mason, 1891

giant red shrimp Diagnosis. Carapace with 4 or more dorsal rostral/postrostral teeth; hepatic spine present; carapace ridges and carinae weak; postcervical sulcus visible only laterally. Pleonites 3–6 dorsally carinate. Pereopods 4, 5 without exopods; pereopod 3 podobranch large; pereopod 4 epipod large. Maximum cl. 76 mm (Pl. 2a).

Shelf, slope (diurnal vertical migrator; 60–1300 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 2 species. One of the two species, A. foliacea, is common, distributed widely and often illustrated (e.g. Fransen 2014). Dall (2001) distinguished the two species.

22

Marine Decapod Crustacea

Fig. 3.1.  Aristeidae. Habitus or carapace: a, Aristaeomorpha foliacea (Risso, 1827); b, Aristaeopsis edwardsiana (Johnson, 1868); c, Aristeus mabahissae Ramadan, 1938; d, e, Austropenaeus nitidus Barnard, 1947 (with male antennular flagella); f, Hemipenaeus carpenteri WoodMason & Alcock, 1891; g, Hepomadus tener Smith, 1884; h, Parahepomadus vaubani Crosnier, 1978; i, Cerataspis montrosus Gray, 1828; j, Pseudaristeus kathleenae Pérez Farfante, 1987; k, basis of pereopod with rudimentary exopod.

Aristaeopsis Wood-Mason, 1891 scarlet shrimp Diagnosis. Carapace ridges and carinae strong; postcervical sulcus visible only laterally. Pleonites 3–6 dorsally carinate. Scaphocerite of male distally elongate, more than in female. Pereopods 4, 5 without exopods; pereopod 3 podobranch large; pereopod 4 epipod large. Maximum cl. 95 mm (Pl. 2b). Slope (200–1850 m). Temperate Northern and Tropical Atlantic, Indo-West Pacific, Temperate Australasia. 1 species. The only species, A.  edwardsiana (Johnson, 1868), has been diagnosed and illustrated by many authors, Fransen (2014) for example, who also summarised its biology.

Aristeus Duvernoy, 1840

Slope (250–1200 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Temperate Australasia. 8 species. Aristeus antennatus is a commercial species and much studied (e.g. Kapiris and Thessalou-Legaki 2011). The weak cervical groove characterises the eight species, separated on subtle differences in proportions of articles of the pereopods (Crosnier 1978; Dall 2001).

Austropenaeus Pérez Farfante & Kensley, 1997 Diagnosis. Carapace ridges and carinae weak; postcervical sulcus visible only laterally. Pleonites 4–6 dorsally carinate. Pereopods 4, 5 with exopods; pereopod 3 podobranch rudimentary; pereopod 4 epipod large. Maximum cl. 38 mm (Pl. 2d). Slope (457–1530 m). Temperate Southern Africa, Temperate Australasia. 1 species (Pérez Farfante and Kensley 1997).

blue and red shrimp, red shrimp Diagnosis. Carapace ridges and carinae weak; postcervical sulcus visible only laterally. Pleonites 4–6 or 3–6 dorsally carinate. Pereopods 4, 5 without exopods; pereopod 3 podobranch absent; pereopod 4 epipod absent. Maximum cl. 77 mm (Pl. 2c).

Cerataspis Gray, 1828 scarlet shrimp Diagnosis. Carapace ridges and carinae strong; postcervical sulcus present dorsally, or visible only laterally. Pleonites 4–6

3 – Dendrobranchiata – prawns and shrimps

dorsally carinate. Pereopods 4, 5 with exopods; pereopod 3 podobranch large; pereopod 4 epipod large. Maximum cl. 78 mm (Pl. 2b). Slope–abyssal (752–5413 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate South America, Temperate Australasia. GOODS bathyal provinces: West Pacific Ocean. GOODS abyssal provinces: Northern Atlantic, Argentine Basin, Central Pacific Ocean, N Central Pacific. 2 species (Dall 2001: as members of Plesiopenaeus). Bracken-Grissom et al. (2012) showed by molecular sequencing that the enigmatic ‘cerataspis’ planktonic larval form belonged to adults of Plesiopenaeus armatus (Bate, 1881). Because the name Cerataspis monstrosus Gray, 1828 is a valid genus and species published before Bate’s name, not only does the genus have taxonomic priority but so too does the species  – C. monstrosus, has priority over P. armatus.

Hemipenaeus Bate, 1881 Diagnosis. Carapace ridges and carinae strong; postcervical sulcus present dorsally, or visible only laterally. Pleonites 3–6 dorsally carinate. Pereopods 4, 5 with exopods; pereopod 3 podobranch large; pereopod 4 epipod rudimentary. Maximum cl. 66 mm (Pl. 2f). Slope–abyssal (900–4022 m). GOODS abyssal provinces: Northern Atlantic, E Antarctic Basin, Indian Ocean, E Pacific basins, Central Pacific Ocean, N Central Pacific. 2 species (Crosnier 1978; Dall 2001).

Hepomadus Bate, 1881 Diagnosis. Hepatic spine present; carapace ridges and carinae weak; postcervical sulcus present dorsally. Pleonites 3–6 dorsally carinate. Pereopods 4, 5 with reduced or without exopods; pereopod 3 podobranch absent; pereopod 4 epipod rudimentary. Maximum cl. 81 mm. Slope–abyssal (765–5400 m). GOODS bathyal provinces: Northern N Pacific Ocean, North Atlantic Ocean, Indian Ocean, North Pacific Ocean. 3 species (Dall 2001: key to species).

Parahepomadus Crosnier, 1978 Diagnosis. Postantennal spine present; carapace ridges and carinae strong; postcervical sulcus visible only laterally. Pleonites 3–6 dorsally carinate. Pereopods 4, 5 without exopods; pereopod 3 podobranch absent; pereopod 4 epipod rudimentary. Maximum cl. 58 mm (Pl. 2g). Slope (750–1525 m). Western and Central Indo-Pacific. 1 species (Crosnier 1978).

Pseudaristeus Crosnier, 1978 Diagnosis. Carapace ridges and carinae strong; postcervical sulcus present dorsally. Pleonites 3–6 dorsally carinate. Pereopods 4, 5 without exopods; pereopod 3 podobranch absent; pereopod 4 epipod absent. Maximum cl. 24 mm.

23

Slope–abyssal (297–4847 m). Western and Central IndoPacific, Temperate Australasia. GOODS abyssal province: Argentine Basin. 6 species (Crosnier 1994; Dall 2001: key to 4 Indo-West Pacific species; Pérez Farfante 1987: key to species). References Bracken-Grissom HD, Felder DL, Vollmer NL, Martin JW, Crandall KA (2012) Phylogenetics links monster larva to deep-sea shrimp. Ecology and Evolution 2, 2367–2373. doi:10.1002/ece3.347 Cau M, Carbonelli A, Follesa MC, Mannini A, Norrito G, et  al. (2002) MEDITS-based information on the deep-water red shrimps Aristaeomorpha foliacea and Aristeus antennatus (Crustacea: Decapoda: Aristeidae). Scientia Marina 66(Suppl. 2), 103–124. doi:10.3989/ scimar.2002.66s2103 Crosnier A (1978) Crustacés Décapodes pénéides Aristeidae (Benthesicyminae, Aristeinae, Solenocerinae). Faune de Madagascar 46, 1–197. Crosnier A (1994) Crustacea Decapoda: Penaeoidea récoltés lors de la campagne KARUBAR en Indonesie. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, Volume 12. Mémoires du Muséum National d’Histoire Naturelle, Paris 161, 351–365. Dall W (2001) Australian species of Aristeidae and Benthesicymidae (Penaeoidea: Decapoda). Memoirs of the Queensland Museum 46, 409–441. Fransen CHJM (2014) Shrimps and prawns. In FAO species identification guide for fishery purposes. The living marine resources of the Eastern Central Atlantic. Volume 1. Introduction, crustaceans, chitons, and cephalopods. (Eds Carpenter KE, De Angelis N) pp. 37–196. FAO, Rome. Holthuis LB (1980) FAO species catalogue. Vol. 1. Shrimps and prawns of the world. An annotated catalogue of species of interest to fisheries. FAO Fisheries Synopsis 125, 1–261. Kapiris K, Thessalou-Legaki M (2011) Feeding ecology of the deep-water blue–red shrimp Aristeus antennatus (Decapoda: Aristeidae) in the Greek Ionian Sea (E. Mediterranean). Journal of Sea Research 65, 151–160. doi:10.1016/j.seares.2010.09.005 Pérez Farfante I (1987) Revision of the gamba prawn genus Pseudaristeus, with description of two new species (Crustacea: Decapoda Penaeoidea). Fishery Bulletin 85, 311–338. Pérez Farfante I, Kensley B (1997) Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175, 1–233. Pezzuto PR, Perez JAA, Wahrlich R (2006) Deep-sea shrimps (Decapoda: Aristeidae): new targets of the deep-water trawling fishery in Brazil. Brazilian Journal of Oceanography 54, 123–134. doi:10.1590/ S1679-87592006000200003

Benthesicymidae Wood-Mason in Wood-Mason & Alcock, 1891 Figure 3.2, Plate 2h Benthesicymid shrimps are, like aristeids, benthopelagic in deep offshore water. Many species have been collected over great depth ranges, from the surface down to thousands of metres and it is assumed that they migrate vertically. Benthesicymidae have long been recognised as morphologically similar to Aristeidae and this is borne out by molecular studies (Cheng et  al. 2018; Ma et  al. 2009; Robalino et  al. 2016; Vazquez-Bader et  al. 2004). Until recently ten genera were recognised, four of which were

24

Marine Decapod Crustacea

diagnosed with lists of species by Pérez Farfante and K ­ ensley (1997). De Grave and Fransen (2011) updated the list of taxa with synonymies but this has become outdated with recent revisions (Lunina et al. 2019; Vereshchaka et al. 2017, 2019, 2021). Two genera, Amalopenaeus Smith, 1882 and Notogennema Vereshchaka & Kulagin in Vereshchaka, Kulagin & Lunina, 2021, each with one species, were recognised after this work went to press (Vereshchaka et al. 2021). Both would key to Gennadas in the key below. These authors based their taxonomy on molecular and morphological data and provided an updated key to genera.

Diagnosis. Body typically shrimp-like; integument soft, stiff but not rigid. Rostrum short, triangular, with 1 or few postrostral spines. Postorbital spine absent; with 1 or 2 postrostral teeth. With at least 3 branchiae on some thoracic somites, at least 11 on each side. Prosartesma (mesial process on antennule article 1) reduced to setose boss. Implicit generic attributes. Rostrum with dorsal teeth only. Hepatic spine absent. Pereopods 4, 5 dactyli tapering, shorter than propodus.

Key to genera of Benthesicymidae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. – 9. –

Pleonites 4, 5 not carinate (Fig. 3.2b, e). Telson truncate (Fig. 3.2f)�����������������������������������������������������������������������������������������2 Pleonites 4, 5 carinate (Fig. 3.2a, c, d). Telson with apical triangular endpiece between terminal spiniform setae (Fig. 3.2g)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������4 Thelycum: thoracic sternite 7 entire. Petasma pars media entire, with straight nearly truncate lamina (Fig. 3.2j)�����������������������������������������������������������������������������������������������������������������������������������������������������������Bentheogennema Thelycum: thoracic sternite 7 divided. Petasma pars media divided, or convex if entire����������������������������������������������������3 Telson without dorsolateral spiniform setae. Thelycum: thoracic sternite 7 trapezoid, subtriangular, or narrow strip. Petasma: grasping structure and lobus accessoris present, pars externa divided or scooplike (Fig. 3.2o)��������������������������������������������������������������������������������������������������������������������������������������������������������������Gennadas Telson with pair of dorsolateral spiniform setae. Thelycum: thoracic sternite 7 pentagonal. Petasma: grasping structure and lobus accessoris absent, pars externa simple subtriangular lash (Fig. 3.2n)������������������������������������������������������������������������������������������������������������������������������������������������������Boreogennema Pereopods 4 and 5 dactyli segmented, much longer than propodus (Fig. 3.2d)������������������������������������������������Benthonectes Pereopods 4 and 5 dactyli unsegmented, tapering, shorter than propodus (Fig. 3.2b)���������������������������������������������������������5 Pleonite 3 dorsally carinate. Rostrum unarmed or bearing rudimentary dorsal tooth (Fig. 3.2a)�������������������� Altelatipes Pleonite 3 not dorsally carinate. Rostrum with 1 or more dorsal teeth (Fig. 3.2b–e)������������������������������������������������������������6 Hepatic spine present (Fig. 3.2c). Maxilliped 3 dactylus subtriangular, with 1 apical spine������������������������������������������������7 Hepatic spine absent. Maxilliped 3 dactylus subrectagular, with several apical spines��������������������������������������������������������8 Pleonite 4 posteriorly crenate. Pereopod 1 basis, ischium with distal tooth. Thelycum sternite 6 or 8 or both with strong median tooth (Fig. 3.2t). Petasma pars media modified (expanded) (Fig. 3.2k)��������Benthesicymus Pleonite 4 posteriorly entire. Pereopod 1 basis, ischium without distal tooth. Thelycum sternite 6 or 8 without median tooth (Fig. 3.2r). Petasma pars media not modified (Fig. 3.2i)�����������������������������������������������Bathicaris Pleonite 6 with posterodorsal margin curved upwards. Thelycum sternite 7 covered with solitary setae (Fig. 3.2o). Petasma pars media with lashes set in rows (grasping structure) (Fig. 3.2u)����������������������������������Dalicaris Pleonite 6 with posterodorsal margin straight. Thelycum sternite 7 without seta. Petasma pars media simple, lamina (Fig. 3.2l, q)�������������������������������������������������������������������������������������������������������������������������������������������������������9 Branchiostegal spine set on carapace anterior margin. Thelycum sternite 7 an anteriorly directed plate. Petasma pars externa not distally produced (Fig. 3.2q)��������������������������������������������������������������������������������� Maorrancaris Branchiostegal spine set behind carapace anterior margin. Thelycum sternite 7 trapezoid, covered with tufts of setae (Fig. 3.2s). Petasma pars externa distally produced (Fig. 3.2l)������������������������������������������������� Benthoecetes Altelatipes Crosnier & Vereshchaka, 2008

Diagnosis. Rostrum unarmed or bearing rudimentary dorsal tooth. Pleonites 3–6 dorsally carinate. Telson apex with apical triangular endpiece between terminal spiniform setae.

Thelycum: sternites 6, 8 without median tooth, sternite 7 covered with tufts of setae. Petasma pars media serrate, laminate. Maximum cl. 30 mm. Slope, bathyal (765–2308 m); benthopelagic. Atlantic, Pacific, Indian oceans. 3 species (Crosnier and Vereshchaka 2008).

3 – Dendrobranchiata – prawns and shrimps

25

Fig. 3.2.  Benthesicymidae. Habitus or carapace: a, Altelatipes falkenhaugae Crosnier & Vereshchaka, 2008; b, Bentheogennema intermedia (Bate, 1888); c, Benthesicymus urinator Burkenroad, 1936; d, Benthonectes filipes Smith, 1885; e, Gennadas elegans (Smith, 1882). Telson: f, Bentheogennema; g, Benthesicymus. Petasma; h, Altelatipes; i, Bathicaris; j, Bentheogennema; k, Benthesicymus; l, Benthoecetes; m, Benthonectes; n, Boreogennema; o, Dalicaris; p, Gennadas; q, Maorrancaris. Thylecum (thoracic sternites 6–8): r, Bathicaris; s, Benthoecetes; t, Benthesicymus; u, Dalicaris. pe, pars externa; pi, pars interna; pm, pars media.

Bathicaris Vereshchaka & Kulagin in Vereshchaka, Corbari, Kulagin, Lunina & Olesen, 2019 Diagnosis. Hepatic spine present. Pleonites 4–6 dorsally carinate. Telson apex with apical triangular endpiece between terminal spiniform setae. Maxilliped 3 dactylus subtriangular, with 1 apical spine. Thelycum: sternites 6, 8 without median tooth, sternite 7 with linguiform process. Petasma pars media serrate, laminate. Maximum cl. 50 mm.

Slope–abyssal (576–6500 m); benthopelagic. Cosmopolitan. 6 species (Vereshchaka et al. 2019).

Bentheogennema Burkenroad, 1936 Diagnosis. Pleonite 6 only dorsally carinate. Telson apex truncate. Thelycum: sternite 7 entire, trapezoid. Petasma pars media entire, with straight nearly truncate lamina. Maximum cl. 20 mm.

26

Marine Decapod Crustacea

Slope–abyssal (0–4360 m); bathypelagic. Atlantic, Indian and probably Pacific oceans. 5 species (Lunina et al. 2019: rediagnosis, key to species). All are probably vertical migrators in oceanic waters (e.g. Heffernan and Hopkins 1981).

Benthesicymus Bate, 1881 Diagnosis. Hepatic spine present. Pleonites 4–6 dorsally carinate. Telson apex with apical triangular endpiece between terminal spiniform setae. Maxilliped 3 dactylus subtriangular, with 1 apical spine. Thelycum: sternite 6 or 8 with median tooth, sternite 7 with sharp anterior process. Petasma pars media modified. Maximum cl. 55 mm. Slope–hadal (1325–9726 m); benthopelagic. Atlantic, Pacific, Indian oceans. 2 species (Vereshchaka et al. 2019: rediagnosis, key to species).

Benthoecetes Smith, 1884 Diagnosis. Branchiostegal spine set behind carapace anterior margin. Pleonites 4–6 dorsally carinate. Telson apex with apical triangular endpiece between terminal spiniform setae. Maxilliped 3 dactylus subrectagular, with several apical spines. Thelycum: trapezoid, covered with tufts of setae. Petasma pars media simple, lamina, distally produced. Maximum cl. 27 mm. Slope, bathyal (600–5777 m); benthopelagic. Atlantic, Pacific, Indian oceans. 3 species (Vereshchaka et al. 2019: key to species as Trichocaris, objective synonym).

Benthonectes Smith, 1885 Diagnosis. Hepatic spine present. Pleonites 4–6 dorsally carinate. Telson apex with apical triangular endpiece between terminal spiniform setae. Pereopods 4, 5 dactyli segmented, much longer than propodus. Thelycum: sternites 6, 8 without median tooth, sternite 7 covered with tufts of setae. Petasma pars media simple, lamina. Maximum cl. 21 mm. Slope (995–1908 m); benthopelagic. Atlantic, Pacific, Indian oceans. 1 species (Vereshchaka et al. 2019: rediagnosis).

Boreogennema Lunina, Kulagin & Vereshchaka, 2019 Diagnosis. Pleonites 6 only dorsally carinate. Telson with 1 pair of dorsolateral spines. Telson apex truncate. Thelycum: sternite 7 divided, pentagonal. Petasma grasping structure and lobus accessoris absent, pars externa simple subtriangular lash. Maximum cl. 20 mm. Mesopelagic, bathypelagic (200–3431 m). North Pacific 1 species (Lunina et al. 2019).

Dalicaris Vereshchaka & Lunina in Vereshchaka, Corbari, Kulagin, Lunina & Olesen, 2019 Diagnosis. Pleonites 4–6 dorsally carinate. Telson apex with apical triangular endpiece between terminal spiniform setae.

Maxilliped 3 dactylus subrectagular, with several apical spines. Thelycum: sternite 7 divided, with solitary setae. Petasma pars media with lashes set in rows (grasping structure). Maximum cl. 50 mm. Slope–abyssal (485–4130 m); benthopelagic. Atlantic, Pacific, Indian oceans. 1 species (Vereshchaka et al. 2019).

Gennadas Bate, 1881 Diagnosis. Pleonite 6 only dorsally carinate. Telson without dorsolateral spines. Telson apex truncate. Thelycum: sternite 7 divided, trapezoid, subtriangular, or narrow strip. Petasma grasping structure and lobus accessoris present, pars externa divided or scooplike. Maximum cl. 13 mm. Epipelagic, mesopelagic, bathypelagic (0–4167 m); vertical migrator. Cosmopolitan 17 species (Dall 2001: key to Indo-West Pacific species). Gennadas is the most diverse genus of Benthesicymidae and the most widely distributed. Four species are panoceanic, four species are confined to the Atlantic and eight inhabit the Indo-West Pacific (Dall 2001; De Grave and Fransen 2011; Pérez Farfante and Kensley 1997; Vereshchaka et al. 2017). Vereshchaka et al. (2017) emended the generic diagnosis, divided the genus into four clades each of 2–5 species, largely on the basis of a phylogenetic analysis of the structure of the petasma and thelycum, and published a key to species.

Maorrancaris Vereshchaka & Lunina in Vereshchaka, Corbari, Kulagin, Lunina & Olesen, 2019 Diagnosis. Pleonites 4–6 dorsally carinate. Telson apex with apical triangular endpiece between terminal spiniform setae. Maxilliped 3 dactylus subrectagular, with several apical spines. Thelycum: sternite 7 divided, without setae. Petasma pars media simple, lamina. Maximum cl. 28 mm (Pl. 2h). Slope (589–1690 m); benthopelagic. Indo-West Pacific. 1 species (Vereshchaka et al. 2019). References Cheng J, Chan T-Y, Zhang N, Sun S, Sha Z-l (2018) Mitochondrial phylogenomics reveals insights into taxonomy and evolution of Penaeoidea (Crustacea: Decapoda). Zoologica Scripta 47, 582–594. doi:10.1111/ zsc.12298 Crosnier A, Vereshchaka AL (2008) Altelatipes falkenhaugae n.gen., n.sp. (Crustacea, Decapoda, Benthesicymidae) de la ride médio-Atlantique nord. Zoosystema 30, 399–411. Dall W (2001) Australian species of Aristeidae and Benthesicymidae (Penaeoidea: Decapoda). Memoirs of the Queensland Museum 46, 409–441. De Grave S, Fransen CHJM (2011) Carideorum Catalogus: the Recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoölogische Mededeelingen 85, 195–589. Heffernan JJ, Hopkins TL (1981) Vertical distribution and feeding of the shrimp genera Gennadas and Bentheogennema (Decapoda: Penaeidea) in the eastern Gulf of Mexico. Journal of Crustacean Biology 1, 461–473. doi:10.2307/1548124

3 – Dendrobranchiata – prawns and shrimps

Lunina AA, Kulagin DN, Vereshchaka AL (2019) A hard-earned draw: phylogeny-based revision of the deep-sea shrimp Bentheogennema (Decapoda: Benthesicymidae) transfers two species to other genera and reveals two new species. Zoological Journal of the Linnean Society 187, 1155–1172. doi:10.1093/zoolinnean/zlz070 Ma KY, Chan T-Y, Chu KH (2009) Phylogeny of penaeoid shrimps (Decapoda: Penaeoidea) inferred from nuclear protein-coding genes. Molecular Phylogenetics and Evolution 53, 45–55. doi:10.1016/j. ympev.2009.05.019 Pérez Farfante I, Kensley B (1997) Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175, 1–233. Robalino J, Wilkins B, Bracken-Grissom HD, Chan T-Y, O’Leary MA (2016) The origin of large-bodied shrimp that dominate modern global aquaculture. PLoS One 11, e0158840. doi:10.1371/journal. pone.0158840 Vazquez-Bader AR, Carrero JC, Garcia-Varela M, Gracia A, Laclette JP (2004) Molecular phylogeny of Superfamily Penaeoidea RafinesqueSchmaltz, 1815, based on mitochondrial 16S partial sequence analysis. Journal of Shellfish Research 23, 911–917. Vereshchaka AL, Lunina AA, Olesen J (2017) The genus Gennadas (­Benthesicymidae: Decapoda): morphology of copulatory characters, phylogeny and coevolution of genital structures. Royal Society Open Science 4, doi:10.1098/rsos.171288. Vereshchaka AL, Corbari L, Kulagin DN, Lunina AA, Olesen J (2019) A phylogeny-based revision of the shrimp genera Altelatipes, Benthonectes and Benthesicymus (Crustacea: Decapoda: Benthesicymidae). Zoological Journal of the Linnean Society 2019, 1–21. Vereshchaka AL, Kulagin DN, Lunina AA (2021) Across the benthic and pelagic realms: a species-level phylogeny of Benthesicymidae (Crustacea: Decapoda). Invertebrate Systematics 35, 776–796.

Penaeidae Rafinesque-Schmaltz, 1815 shrimp, prawns Figures 3.3–3.5, Plate 3 Most edible shrimp, or prawns, as they are called in Australia, belong to Penaeidae. All have a strong serrated rostrum, a laterally flattened body and pleon, three pairs of minutely chelate legs, and pleonites overlapping tile-like from the front. The last features distinguish penaeids from caridean shrimps, also commercially fished, which have at most only two pairs of chelate legs, and the second pleonite overlapping segments both in front and behind. Shrimps or prawns sold in markets are caught with trawl nets skimming above the bottom in the muddy waters of shallow bays and estuaries, mostly in the tropics. The FAO recommends its own international common names (Holthuis 1980). Australians have their own colourful names for the edible prawns found in this country: bay prawn, greasy back prawn, school prawn, banana prawn, king prawn, tiger prawn, rough prawn and others. Many of these names refer to more than one species and some are prefixed by other adjectives, such as the blue-legged, western, eastern and red-spot king prawns, or brown, blue, green and jumbo tiger prawns. Because of their fishery

27

interest, shrimps are among the best understood of all Crustacea (Dall et al. 1990). While confusion over what common name to use can be expected because many names are only locally applicable, more certainty could be expected with taxonomic names. Unfortunately, this is not always the case. Most commercial shrimps have traditionally been placed in the genus Penaeus and only a few smaller genera recognised. Following a taxonomic revision based on morphological investigation by Pérez Farfante and Kensley (1997) Penaeus was divided into six smaller genera by reviving names already proposed but these have again fallen out of use (see discussion under Penaeus below). Pérez Farfante and Kensley’s names, and those of Sakai and Shinomiya (2011) who divided Parapenaeopsis into eight genera, appeared in a catalogue of species (De Grave and Fransen 2011). Chan et al. (2008) and Chan (2012), resolved three clades treated as tribes following Burkenroad (1983) (they could just as equally be considered subfamilies) but they persisted with the synonymised generic names: Penaeinae  – ‘Farfantepenaeus’, ‘Fenneropenaeus’, Funchalia, Heteropenaeus, ‘Litopenaeus’, ‘Marsupenaeus’, ‘Melicertus’, Pelagopenaeus, Penaeus s.s.; Trachypenaeinae – Atypopenaeus, Metapenaeus, Megokris, Parapenaeopsis, Rimapenaeus, Trachysalambria, Xiphopenaeus, to which could be added Sakai and Shinomiya’s genera; Parapenaeinae – Metapenaeopsis, Penaeopsis, Parapenaeus. Penaeidae has been found to be paraphyletic, Sicyoniidae being an offshoot (Ma et  al. 2009; Robalino et  al. 2016). Cheng et  al. (2018) went further, concluding the three ‘tribes’ are polyphyletic, one clade comprising Solenoceridae + Aristeidae + Benthesicymidae sister to Penaeinae, and Sicyoniidae derived within Trachypenaeinae. Cheng et  al. (2018) proposed elevating two tribes to family rank, Penaeidae s.s., Parapenaeidae, and Sicyoniidae as a senior synonym of Trachypenaeinae. Pérez Farfante and Kensley (1997) divided the large genus Penaeus by reviving names that had until then not be widely accepted: Farfantepenaeus, Fenneropenaeus, Litopenaeus, Marsupenaeus and Melicertus. Because their work became the definitive revision of dendrobranchiate classification some of these names began to appear in the literature. Davie (2002) argued that the commercial importance in fisheries and aquaculture of many species demanded that names be treated conservatively while uncertainty remains. Some of the genera proposed conflicted with molecular data (Dall 2007) and were not appreciated by those with more commercial interests (Flegel 2007, 2008). McLaughlin et al. (2008) advocated that the rules of nomenclature should

28

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prevail. Early molecular evidence was ambivalent (Baldwin et  al. 1998; Gusmão et  al. 2000; Maggioni et  al. 2001). A study involving 30 species of 20 nominal genera (Chan et al. 2008) raised further suspicion. This debate become moot in the light of more recent molecular research that has shown that Pérez Farfante and Kensley’s (1997) division may not be fully supported (Ma et al. 2011 and earlier studies). Ma et al. (2011) used mitochondrial and nuclear markers to confirm that Penaeus s.l. is monophyletic, Melicertus and Penaeus s.s. are paraphyletic and Farfantepenaeus is genetically distant from Melicertus. None of these groupings correlated with morphological traits. Pérez Farfante and Kensley’s generic names have fallen out of favour but sometimes continue to be used for established species and for new ones (e.g. Tavares and Gusmão 2016). Ramirez et al. (2021) found a high level of cryptic species diversity throughout the family that further complicated generic divisions. While the classification of Penaeidae, and penaeoids in general, remains so fluid we take a practical approach to identification of genera. Pérez Farfante and Kensley (1997) recognised 26 genera and provided a key. We have updated this taking into account the classification proposed by Ma et al. (2011) and adopted by WoRMS Editorial Board (2022). Genera are distinguished partly on the presence or absence of branchiae (gills) above the legs (Fig. 3.3d) and on the arrangement of carinae and sutures on the carapace. Carapace ornamentations are variously termed by different authors – those used here are shown in Fig. 3.3a but more detail is needed to separate species. Reproductive apparatus is also used to distinguish genera and species. The thelycum (Fig. 3.3g–j) is a peculiar modification of the last two or three thoracic sternites (sternites 6–8) of the female that serves for the storage and transfer of spermatophores from

the male. The thelycum may be closed (with seminal receptacles, paired or unpaired sacs opening through a slit or slits on sternite 8) or open (without seminal receptacles). The petasma (Fig.  3.3k–z) is a modification of the enlarged endopods of pleopods 1 of the male, hooked together by hooked setae. Each half is called a valve, comprising a median and a lateral lobe. The apex of each lobe is various convoluted with lobules, extensions and projections that serve to differentiate genera. The so-called ventral face of the petasma (actually the posterior face) is viewed when the petasma is folded anteriorly; the dorsal face when folded posteriorly. The thelycum and petasma of all genera were figured by Pérez Farfante and Kensley (1997). Diagnosis. Body typically shrimp-like; integument soft, stiff but not rigid. Rostrum elongate, blade-like with dorsal and sometimes ventral teeth. Postorbital spine absent; with 3 or more postrostral teeth postrostral teeth. With at least 3 branchiae on some thoracic somites, at least 11 on each side. Prosartesma (mesial process on antennule article 1) well developed (Fig. 3.3b) Implicit generic attributes. Rostrum with dorsal teeth only; adrostral sulcus and carina absent. Pterygostomian spine absent, anteroventral angle rounded; integument smooth; carapace without dorsolateral carina or suture. Telson unarmed. Antennular peduncle, article 1 without ventromesial spine. Pereopods 1–3 with usual chelae, fingers at least half length of propodus; pereopod 2 ischium without spine; pereopods 4 and 5 about as long as pereopods 1–3; 4, 5 dactyli entire, single article. Without pleurobranch on thoracic somite 8; pereopod 3 with epipod; exopods on maxillipeds 1–3 and pereopods 1–5. Petasma symmetrical; distomedian projections leaf-like; distolateral projections unilobed, without lobule.

Key to genera of Penaeidae The key contains multiple choices.

1. – 2. – 3. – 4.

Pleurobranchs on thoracic somite 8 (above pereopod 5). Rostrum with dorsal and ventral teeth (Figs 3.4d, 3.5d) (dorsal only in mesopelagic Funchalia)�������������������������������������������������������������������������������������������������������������������������2 Without pleurobranchs on thoracic somite 8 (above pereopod 5). Rostrum with dorsal teeth only����������������������������������5 Integument glabrous, polished. Pleonite 6 with 3 short well defined lateral raised ridges (Fig. 3.5d)������������������ Penaeus Integument fine, setose. Pleonite 6 with 1 long lateral ridge (Fig. 3.4d)����������������������������������������������������������������������������������3 Integument with numerous sulci overlapped by rows of dense stiff setae projecting from one margin. Telson with 4 pairs of movable lateral spines. Antennular flagella less than half length of carapace (Fig. 3.4d)��������������������������������������������������������������������������������������������������������������������������������������������������������������Heteropenaeus Integument with few sulci, not overlapped by rows of dense stiff setae projecting from one margin. Telson with 3 pairs of movable lateral spines. Antennular flagella longer than carapace�����������������������������������������������������������4 Rostrum with dorsal teeth only. Carapace without dorsolateral carina (between dorsal and branchiocardiac carinae) (Fig. 3.4c). Mandibular incisor elongate, scythe-like, crossing posterior to labrum (Fig. 3.3c). Petasma asymmetrical (Fig. 3.3o)��������������������������������������������������������������������������������������������������������� Funchalia

3 – Dendrobranchiata – prawns and shrimps



29

Rostrum with dorsal and ventral teeth. Carapace with dorsolateral carina (between dorsal and branchiocardiac carinae) (Fig. 3.5b). Mandibular incisor short, not crossing. Petasma symmetrical������������������Pelagopenaeus 5. Telson with pair of fixed subapical spines, preceded by lateral movable spines (Fig. 3.3f). Antennular article 1 usually with ventromesial spine��������������������������������������������������������������������������������������������������������������������������������������������6 – Telson without pair of fixed subapical spines, usually with lateral movable spines (Fig. 3.3e). Antennular article 1 without ventromesial spine����������������������������������������������������������������������������������������������������������������������������������������9 6. Carapace with longitudinal and transverse sutures (Fig. 3.5a). Telson with 1 pair of dorsolateral spines anterior to subapical spines���������������������������������������������������������������������������������������������������������������������������������� Parapenaeus – Carapace without longitudinal suture. Telson with 2 or more pair of dorsolateral spines anterior to subapical spines���������������������������������������������������������������������������������������������������������������������������������������������������������������������������7 7. Pterygostomian spine absent (anteroventral corner of carapace square) (Fig. 3.4a). Pereopod 1 basis without spine. Maxilliped 2 and all pereopods without exopod����������������������������������������������������������������������������������������Artemesia – Pterygostomian spine present (anteroventral corner of carapace acute) (Figs 3.4g, h, 3.5c). Pereopod 1 basis with spine. Maxillipeds and all pereopods with exopod������������������������������������������������������������������������������������������������������8 8. Maxilliped 3 and pereopod 2 bases with spine. Petasma asymmetrical (Fig. 3.3u)�������������������������������������Metapenaeopsis – Maxilliped 3 and pereopod 2 bases without spine. Petasma symmetrical�������������������������������������������������������������Penaeopsis 9. Pleurobranchs on thoracic somite 7 (above pereopod 4). Maxillipeds and pereopods 1–4 with exopods������������������������������������������������������������������������������������������������������������������������������������������������������������������� Metapenaeus – Pleurobranchs absent from thoracic somite 7 (above pereopod 4). Pereopod 1 only or pereopods 1–4 with exopods�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 10 10. Maxilliped 1 and pereopod 1 only with exopods. Petasma with lateral lobes distally produced, long, filamentous (Fig. 3.3s). Thylecum with lateral plates curved posteriorly, expanded as bulbous anterior swellings (Fig. 3.3g)���������������������������������������������������������������������������������������������������������������������������������������������Macropetasma – Maxilliped 3 and pereopods 1–5 only with exopods. Petasma with lateral lobes not distally produced. Thylecum with lateral plates not curved posteriorly, not expanded as bulbous anterior swelling������������������������������11 11. Carapace without longitudinal or transverse sutures (Fig. 3.5h). Telson with subapical pair of lateral movable spines on elongate shoulder. Petasma with ventrolateral lobe produced into 2 or 3 flaps (Fig. 3.3x)���������������������������������������������������������������������������������������������������������������������������������������������������������Trachypenaeopsis – Carapace with longitudinal or transverse sutures or both. Telson without lateral spines or with subapical pair of lateral movable spines not on shoulder. Petasma with ventrolateral lobe not produced as flaps������������������� 12 12. Maxilliped 2 without exopod. Pereopods 1–3 with elongate chela������������������������������������������������������������������� Protrachypene – Maxilliped 2 with exopod. Pereopods 1–3 with chela not elongate���������������������������������������������������������������������������������������13 13. Pereopods 4, 5 elongate, flagelliform, longer than pereopods 1–3 (Fig. 3.5g, j)�������������������������������������������������������������������14 – Pereopods 1–5 of similar lengths, none elongate����������������������������������������������������������������������������������������������������������������������15 14. Integument pubescent. Telson with 4 pairs of movable spines. Pereopods 4, 5 with dactylus entire (Fig. 3.5g)�����������������������������������������������������������������������������������������������������������������������������������������������������������������Tanypenaeus – Integument smooth. Telson unarmed. Pereopods 4, 5 with dactylus multiarticulate (Fig. 3.5j)�����������������Xiphopenaeus 15. Carapace without longitudinal suture (Fig. 3.4b). Pereopod 2 ischium with spine����������������������������������������Atypopenaeus – Carapace with longitudinal suture (Fig. 3.4j). Pereopod 2 ischium without spine��������������������������������������������������������������16 16. Body slender. Integument thin (Fig. 3.4j). Pereopod 3 without epipod���������������������������������������������������������������������������������17 – Body thickset. Integument thick (Fig. 3.5f). Pereopod 3 with epipod���������������������������������������������������������������������������������� 22 17. Petasma with distomedian projections leaf-like (Fig. 3.3v)����������������������������������������������������������������������������Mierspenaeopsis – Petasma with distomedian projections not leaf-like�����������������������������������������������������������������������������������������������������������������18 18. Petasma with distolateral projections divided into 2 lobes, a ventral hook and dorsal finger (Fig. 3.3l)��������������������������������������������������������������������������������������������������������������������������������������������������������Arafurapenaeopsis – Petasma with distolateral projections of 1 lobe�������������������������������������������������������������������������������������������������������������������������19 19. Petasma with distolateral projections each with a lobule������������������������������������������������������������������������������������������������������� 20 – Petasma with distolateral projections simple�����������������������������������������������������������������������������������������������������������������������������21

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Marine Decapod Crustacea

20. – 21. – – –

Petasma, lobules of distolateral projections dorsal (Fig. 3.3p)�������������������������������������������������������������������Ganjampenaeopsis Petasma, lobules of distolateral projections ventral (Fig. 3.3k)�������������������������������������������������������������������� Alcockpenaeopsis Petasma with distolateral projections diverging laterally at 90° (Fig. 3.3q)��������������������������������������������Holthuispenaeopsis Petasma with distolateral projections diverging posterolaterally (Fig. 3.3n)�������������������������������������������������Batepenaeopsis Petasma with distolateral projections diverging straight and anterolaterally at 45° (Fig. 3.3w)�����������������Parapenaeopsis Petasma with distolateral projections diverging anteriorly in proximal two-thirds and inwards in distal third (Fig. 3.3r)���������������������������������������������������������������������������������������������������������������������������������������� Kishinouyepenaeopsis 22. Carapace with longitudinal suture conspicuously overreaching hepatic spine (Fig. 3.5f). Maxilliped 3 basis with spine���������������������������������������������������������������������������������������������������������������������������������������������������������������Rimapenaeus

Fig. 3.3.  Penaeidae. a, carapace (features labelled: ac, adrostral carina; as, antennal spine; cc, cervical carina; et, epigastric tooth; hc, hepatic carina; ls, longitudinal suture; pt pterygostomian spine; rt, rostral teeth; ts, transverse suture); b, antennule, lateral; c, mandible, Funchalia; d, anterior pereopod; e, telson with lateral movable spines; f, telson with fixed subapical spines. Thylecum (anterior uppermost; lp, lateral plate): g, Macropetasma; h, Megokris; i, Trachypenaeus; j, Trachysalambria. Petasma (dorsal views except k, p, s, u, ventral views; dlp, dorsolateral projection; dmp, dorsomedian projection; l, lobule): k, Alcockpenaeopsis; l, Arafurapenaeopsis; m, Atypopenaeus; n, Batepenaeopsis; o, Funchalia; p, Ganjampenaeopsis; q, Holthuispenaeopsis; r, Kishinouyepenaeopsis; s, Macropetasma; t, Megokris; u, Metapenaeopsis; v, Mierspenaeopsis; w, Parapenaeopsis; x, Trachypenaeopsis; y, Trachypenaeus; z, Trachysalambria.

3 – Dendrobranchiata – prawns and shrimps

31

Fig. 3.4.  Penaeidae. Habitus or carapace: a, Artemesia longinaris Bate, 1888; b, Atypopenaeus bicornis Racek & Dall, 1965; c, Funchalia villosa (Bouvier, 1905); d, Heteropenaeus longimanus De Man, 1896; e, Macropetasma africana (Balss, 1913); f, Megokris granulosus (Haswell, 1879); g, Metapenaeopsis commensalis Borradaile, 1899 (without stridulating ridge); h, h’, Metapenaeopsis rosea Racek & Dall, 1965 (with stridulating ridge, detail); i, Metapenaeus affinis (H. Milne Edwards, 1837); j, Parapenaeopsis hardwicki (Miers, 1878).



Carapace with longitudinal suture ending well anterior to hepatic spine (Figs 3.4f, 3.5h, i). Maxilliped 3 basis without spine������������������������������������������������������������������������������������������������������������������������������������������������������������������� 23 23. Thelycum with plate on thoracic sternite 8 (between pereopods 5) short medially, deeply excavate, embracing extremely long caudal extension of medial protuberance (Fig. 3.3h). Petasma with distolateral projections extending to hook-like tip or narrowing rapidly from broad base to mesoanteriorly directed tip (Fig. 3.3t)�������������������������������������������������������������������������������������������������������������������������������������������������������������������Megokris

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Marine Decapod Crustacea

Fig. 3.5.  Penaeidae. Habitus or carapace: a, Parapenaeus fissurus (Bate, 1881); b, Pelagopenaeus balboae (Faxon, 1893); c, Penaeopsis eduardoi Pérez Farfante, 1977; d, Penaeus japonicus Bate, 1888; e, Protrachypene precipua Burkenroad, 1934; f, Rimapenaeus faoe (Obarrio, 1954); g, Tanypenaeus caribaeus Pérez Farfante, 1972; h, Trachypenaeopsis mobilispinis (Rathbun, 1915); i, Trachysalambria curvirostris (Stimpson, 1860); j, Xiphopenaeus kroyeri (Heller, 1862).

– –

Thelycum with plate on thoracic sternite 8 (between pereopods 5) long, not emarginate, without caudal extension (Fig. 3.3i). Petasma with distolateral projections broad, extending laterally to form apically long flat twisted process (Fig. 3.3y)����������������������������������������������������������������������������������������������������������������� Trachypenaeus Thelycum with plate on thoracic sternite 8 shallowly emarginate or slightly produced, without caudal extension (Fig. 3.3j). Petasma with distolateral projections tapering from narrow base, straight or curving backwards (Fig. 3.3z)����������������������������������������������������������������������������������������������������������������������� Trachysalambria

3 – Dendrobranchiata – prawns and shrimps

Alcockpenaeopsis Sakai & Shinomiya, 2011 dog shrimp Diagnosis. Carapace with transverse sutures; with longitudinal suture conspicuously overreaching hepatic spine. Pereopod 3 without epipod. Petasma distolateral projections unilobed, with ventral lobule. Maximum cl. 20 mm. Intertidal, subtidal (0–13 m). Central Indo-Pacific. 1 species. This and other penaeid genera named for early carcinologists were differentiated from Parapenaeopsis by Sakai and Shinomiya (2011) on the basis of variation in the shape of the petasma. Li et al. (2014) undertook a molecular study of six Chinese species of Parapenaeopsis, too few to confirm or refute the division.

Arafurapenaeopsis Sakai & Shinomiya, 2011 Arafura shrimp Diagnosis. Carapace with transverse sutures; with longitudinal suture conspicuously overreaching hepatic spine. Pereopod 3 without epipod. Petasma distolateral projections bilobed. Maximum cl. 25 mm. Subtidal, shelf (5–30 m); estuarine. Central Indo-Pacific (N Australia, Papua New Guinea). 1 species (Sakai and Shinomiya 2011).

Artemesia Bate, 1888 stiletto shrimp Diagnosis. Carapace without transverse sutures. Telson with pair of fixed subapical spines, with 2 or more pair of movable spines. Antennular peduncle, article 1 with ventromesial spine; flagella as long or longer than carapace. Exopods on maxillipeds 1 and 3 only. Maximum cl. 44 mm. Intertidal–shelf (0–125 m). Tropical W Atlantic. 1 species. Artemesia longinaris Bate, 1888 supports a fishery in eastern South America where it is well studied (da Costa et al. 2010).

Atypopenaeus Alcock, 1905 orange shrimp, periscope shrimp

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without epipod. Petasma distolateral projections diverging posterolaterally. Maximum cl. 20 mm. Subtidal, shelf (5.5–48 m). Temperate W Northern Pacific, Western and Central Indo-Pacific. 4 species. Some species of Batepenaeopsis are of minor commercial importance in SE Asia. Shinomiya and Sakai (2016) reviewed the species and provided a key.

Funchalia Johnson, 1868 Diagnosis. Pterygostomian spine present, anteroventral corner tooth-like; integument setose; carapace without transverse sutures. Telson with 3 pairs of fixed subapical spines preceded by smaller ones. Antennular flagella as long or longer than carapace. Pereopod 2 ischium with spine. With pleurobranch on thoracic somite 8. Petasma asymmetrical. Maximum cl. 36 mm (Pl. 3b). Mesopelagic (0–3000 m). Atlantic Ocean, Mediterranean, IndoWest Pacific. 5 species. Dall (1957) diagnosed the genus and provided a key to five species but one of these is now Pelagopenaeus balboae (Grippa 1987). Fujino’s (1975) species from the Pacific is doubtful.

Ganjampenaeopsis Sakai & Shinomiya, 2011 Diagnosis. Carapace with transverse sutures; with longitudinal suture conspicuously overreaching hepatic spine. Pereopod 3 without epipod. Petasma distolateral projections unilobed, with dorsal lobule. Maximum cl. 31 mm. Subtidal, shelf (5–82 m). Western Indo-Pacific (Bay of Bengal). 1 species (Alcock 1905).

Heteropenaeus de Man, 1896 Diagnosis. Rostrum with dorsal and ventral teeth. Integument with numerous sulci overlapped by rows of dense stiff setae projecting from one margin; carapace without transverse sutures. Telson with 4 pairs of movable spines. Antennular flagella less than length of carapace. With pleurobranch on thoracic somite 8. Maximum cl. 18 mm. Subtidal (4–17 m). Central Indo-Pacific. 1 species (Pérez Farfante and Kensley 1997).

Holthuispenaeopsis Sakai & Shinomiya, 2011

Diagnosis. Carapace without transverse sutures, or with transverse sutures. Antennular flagella as long or longer than carapace. Pereopod 2 ischium with spine; pereopods 4 as long as pereopod 3, pereopod 5 elongate, much longer than 3 and 4. Maximum cl. 38 mm (Pl. 3a). Subtidal, shelf (2–27 m); estuarine. Temperate Northern W Pacific, Western and Central Indo-Pacific. 5 species. Some species of Atypopenaeus are of minor commercial importance in India and Japan.

Diagnosis. Carapace with transverse sutures; with longitudinal suture conspicuously overreaching hepatic spine. Pereopod 3 without epipod. Petasma distolateral projections diverging laterally at 90°. Maximum cl. 22 mm. Estuarine, subtidal, shelf (1–60 m). Tropical E Atlantic. 1 species (Holthuis 1980).

Batepenaeopsis Sakai & Shinomiya, 2011

Kishinouyepenaeopsis Sakai & Shinomiya, 2011

Guinea shrimp

smoothshell shrimp

coral shrimp, torpedo shrimp

Diagnosis. Carapace with transverse sutures; with longitudinal suture conspicuously overreaching hepatic spine. Pereopod 3

Diagnosis. Carapace with transverse sutures; with longitudinal suture conspicuously overreaching hepatic spine. Pereopod 3

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Marine Decapod Crustacea

without epipod. Petasma distolateral projections diverging anteriorly in proximal two-thirds, then inwards. Maximum cl. 24 mm. Subtidal, shelf (1–37 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 4 species (Sakai and Shinomiya 2011).

Macropetasma Stebbing, 1914

Metapenaeus Wood-Mason in Wood-Mason & Alcock, 1891 jinga shrimp, fire shrimp, greentail shrimp, yellow shrimp, wood shrimp, school prawn, Endeavour prawn

Diagnosis. Pterygostomian spine absent, anteroventral angle rounded, or present, anteroventral corner tooth-like; integument setose; carapace with transverse sutures; carapace with short longitudinal suture above antennal spine. Telson with 3 pairs of movable spines. Antennular flagella less than length of carapace. Thelycum with plate on thoracic sternite 8 short medially, deeply excavate, embracing extremely long caudal extension of medial protuberance. Maximum cl. 26 mm. Subtidal, shelf (2–81 m). Western and Central Indo-Pacific. 9 species (Shinomiya and Sakai 2006: key to species).

Diagnosis. Rostrum without ventral teeth. Integument smooth, or setose; carapace without transverse sutures. Telson with numerous small movable spines. Antennular peduncle, article 1 with ventromesial spine; flagella less than length of carapace. Exopods on maxillipeds 1–3 and pereopods 1–4. Maximum cl. 50 mm. Estuarine, intertidal–shelf (some species entering fresh water) (0–92 m). Temperate Northern Atlantic (E Mediterranean), Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 28 species. Species of Metapenaeus are common in estuaries where they feature in amateur fisheries while larger representatives, such as Endeavour prawns, are more commercial. Racek and Dall (1965) provided a key to the Indo-West Pacific species then known. Miquel (1982) reviewed taxonomy, biology and fisheries world-wide for all 25 species and also provided a key to distinguish them. All species live in the Indo-West Pacific but two have entered the eastern Mediterranean through the Suez Canal or in ballast water (Aydin et al. 2009). Metapenaeus monoceros (Fabricius, 1798) is well established and the basis of an important fishery there and in its home range. Metapenaeus bennettae Racek & Dall, 1965, from eastern Australia, has become established in northern New Zealand (Ahyong and Wilkens 2011).

Metapenaeopsis Bouvier, 1905

Mierspenaeopsis Sakai & Shinomiya, 2011

swimming prawn Diagnosis. Integument setose (sparsely); carapace without transverse sutures. Telson with 4 pairs of movable spines. Antennular flagella as long or longer than carapace. Exopods on maxilliped 3 and pereopod 1. Petasma distolateral projections distally produced, long, filamentous. Maximum cl. 11 mm (Pl. 3c). Estuarine, subtidal, shelf (0–30 m). Temperate Southern Africa. 1 species (Cockcroft and Wooldridge 1987).

Megokris Pérez Farfante & Kensley, 1997

velvet shrimp, humpback shrimp, fiddler shrimp

spear shrimp, rainbow shrimp

Diagnosis. Pterygostomian spine present, anteroventral corner tooth-like; integument setose; carapace without transverse sutures. Telson with pair of fixed subapical spines, with 2 or more pair of movable spines. Antennular peduncle, article 1 with ventromesial spine; flagella less than length of carapace. Pereopod 2 ischium with spine. Petasma asymmetrical. Maximum cl. 40 mm (Pl. 3d). Subtidal–slope (few species) (0–894 m). Cosmopolitan except temperate South America and polar seas. 72 species. Metapenaeopsis is the largest genus of penaeids and well studied. Some species are called velvet shrimps because of the texture of the exoskeleton. The keys of Racek and Dall (1965) and Crosnier (1994) dealt with shallow-water species of the Indo-West Pacific with a stridulating organ (Fig.  3.4h) on the posterior margin of the carapace while those of Crosnier (1987, 1991) covered deep-water species in which this organ is absent. The division of most of the genus, those with a bivalved/symmetrical petasma, into two groups based on the presence or absence of the stridulating organ was supported by an analysis of molecular data (Cheng et al. 2015; Tong et al. 2000), but species with a univalved/asymmetrical petasma are excluded from these groups. While the species with a bivalved petasma inhabit sandy or muddy bottoms the univalved species are symbiotic and associated with coral reefs, at least in the Indo-West Pacific.

Diagnosis. Carapace with transverse sutures; with longitudinal suture conspicuously overreaching hepatic spine. Pereopod 3 without epipod. Petasma distomedian projections undeveloped. Maximum cl. 21 mm. Subtidal, shelf (1–90 mm). Western and Central Indo-Pacific. 4 species (Sakai and Shinomiya 2011).

Parapenaeopsis Alcock, 1901 hawknose shrimp, dwarf shrimp Diagnosis. Carapace with transverse sutures; with longitudinal suture conspicuously overreaching hepatic spine. Telson with pair of fixed subapical spines, with 2 or more pair of movable spines, or unarmed. Pereopod 3 without epipod. Petasma distolateral projections diverging straight, anterolaterally at 45°. Maximum cl. 20 mm (Pl. 3f). Subtidal, shelf (7–124 mm). Indo-West Pacific. 6 species. The 20 species of Parapenaeopsis listed by Pérez Farfante and Kensley (1997) were divided into eight genera by Sakai and Shinomiya (2011), all recognised here. Sakai and Shinomiya’s (2011) seven new genera have not been tested by molecular methods. Dall’s (1957) key to 13 species includes those assigned to the newer genera.

3 – Dendrobranchiata – prawns and shrimps

Parapenaeus Smith, 1885 rose shrimp Diagnosis. Pterygostomian spine present, anteroventral corner tooth-like; carapace with transverse sutures. Telson with pair of fixed subapical spines, with 1 pair of movable spines. Antennular peduncle, article 1 with ventromesial spine; flagella as long or longer than carapace. Maximum cl. 34 mm. Subtidal–slope (13–700 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 15 species. Crosnier’s (1986) key differentiated ten Indo-West Pacific species. Yang et al. (2015) recognised one group of species from the Atlantic and three from the Indo-West Pacific, more or less following Crosnier’s scheme.

Pelagopenaeus Pérez Farfante & Kensley, 1997 Diagnosis. Rostrum with dorsal and ventral teeth. Pterygostomian spine present, anteroventral corner tooth-like; integument setose; carapace without transverse sutures; carapace with elongate dorsolateral carina commencing well behind orbit, running between dorsal carina and hepatic-branchiocardiac carinae. Telson with 3 pairs of fixed subapical spines preceded by smaller ones. Antennular flagella as long or longer than carapace. Pereopod 2 ischium with spine. With pleurobranch on thoracic somite 8. Maximum cl. 28 mm. Bathypelagic (0–1609 m). 1 species (Grippa 1987). Tropical Indo-Pacific, Tropical Pacific, Tropical Atlantic.

Penaeopsis Bate, 1881 needle shrimp, megalops shrimp Diagnosis. Pterygostomian spine present, anteroventral corner tooth-like; carapace without transverse sutures. Telson with pair of fixed subapical spines, with 2 or more pairs of movable spines. Antennular peduncle, article 1 with ventromesial spine; flagella as long or longer than carapace. Maximum cl. 54 mm (Pl. 3e). Shelf, slope (120–750 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Western and Central IndoPacific, Temperate Southern Africa, Temperate Australasia. 7 species. Pérez Farfante (1980) rediagnosed the genus in which she recognised six species. Crosnier (2006) redescribed three species and added another. The shrimps are benthic on the upper part of continental slopes, especially in the tropics of the Indo-West Pacific. Some species are sufficiently abundant to be commercially exploited.

Penaeus Fabricius, 1798 Diagnosis. Rostrum with dorsal and ventral teeth; adrostral sulcus and carina present. Carapace without transverse sutures. Antennular flagella less than length of carapace. Pereopod 2 ischium without spine, or with spine. With pleurobranch on thoracic somite 8; exopods on maxillipeds 1–3 and pereopods 1–5, or on maxillipeds 1–3 and pereopods 1–4. Maximum cl. 90 mm (Pl. 3g). Estuarine (as juveniles), intertidal–slope (rarely) (0–365 m). Temperate Northern and Tropical Atlantic, Temperate Northern

35

Pacific, Indo-West Pacific, Tropical Eastern Pacific, Temperate Southern Africa, Temperate Australasia. 30 species. The taxonomy of Penaeus sensu lato remains problematic, as discussed under the family heading above, but we follow the single genus classification of Ma et al. (2011).

Penaeids are fished in commercial quantities in many parts of the world and have numerous local popular names. Aquaculture, however, is now the dominant source of shrimp for human consumption (Jory 2019). Penaeus vannamei Boone, 1931 (otherwise Litopenaeus vannamei), called western white shrimp over its native range, the Pacific coast of Central and South America, is the dominant cultivated species in Latin America. In south-east Asia, tropical Australia and countries bordering the Indian Ocean, P.  monodon Fabricius, 1798 is widely cultivated (FAO 2007–2019). This ‘species’ was recently found to harbour a pseudocryptic species in Indonesia (Chan et al. 2021). Most species of Penaeus live at shallow shelf depths having juveniles that enter estuaries (Primavera 1998) but P. canaliculatus (Olivier, 1811), P. latisulcatus Kishouye, 1896 and P. marginatus Randall, 1840 occur with other penaeids at slope depths near hydrothermal vents off T ­ aiwan (Wang et al. 2013). Protrachypene Burkenroad, 1934 titi shrimp Diagnosis. Integument setose; carapace with transverse sutures; carapace with short longitudinal suture above antennal spine. Telson with numerous small movable spines. Antennular flagella as long or longer than carapace. Pereopods 1–3 with elongate chelae, propodus long and fingers less than half length of propodus; pereopods 4 and 5 elongate, flagelliform, much longer than pereopod 3. Exopods on maxillipeds 1 and 3, pereopods 1–5. Maximum cl. 25 mm. Intertidal–shelf (0–40 m). Tropical Eastern Pacific. 1 species (Santamaría et al. 2018).

Rimapenaeus Pérez Farfante & Kensley, 1997 Diagnosis. Integument setose; carapace with transverse sutures; carapace with longitudinal suture conspicuously overreaching hepatic spine. Telson with 3 or 4 pairs of movable spines. Antennular flagella less than length of carapace. Maximum cl. 41 mm. Subtidal–slope (2–228 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 6 species (Pérez Farfante 1971: key to E Pacific species as members of Trachypenaeus).

Tanypenaeus Pérez Farfante, 1972 Diagnosis. Integument setose; carapace without transverse sutures; carapace with short longitudinal suture above antennal spine. Telson with 3 or 4 pairs of movable spines. Antennular flagella as long

36

Marine Decapod Crustacea

or longer than carapace. Pereopods 4 and 5 elongate, flagelliform, much longer than pereopod 3. Maximum cl. 19 mm. Shelf (155–157 m). Tropical W Atlantic. 1 species (Pérez Farfante 1972).

Trachypenaeopsis Burkenroad, 1934 Diagnosis. Integument setose; carapace without transverse sutures. Telson with pair of fixed subapical spines, with 2 or 3 pairs of movable spines on strong shoulder. Antennular flagella less than length of carapace. Maximum cl. 12 mm. Estuarine, subtidal, shelf (3–100 m). Tropical W Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific. 3 species (Thomas 1972: key to species).

Trachypenaeus Alcock, 1901 hardback shrimp Diagnosis. Integument setose; carapace without transverse sutures; with short longitudinal suture above antennal spine. Telson with 3 pairs of movable spines (on strong shoulder). Antennular flagella less than length of carapace. Thelycum with plate on thoracic sternite 8 long, not emarginate, without caudal extension. Maximum cl. 23 mm. Subtidal, shelf (12.5–52 m). Central Indo-Pacific. 1 species. Chanda (2018) rediagnosed Trachypenaeus but treated Megokris as a junior synonym.

Trachysalambria Burkenroad, 1934 Diagnosis. Integument setose; carapace with transverse sutures; carapace with short longitudinal suture above antennal spine. Telson with 3 pairs of movable spines. Antennular flagella less than length of carapace. Thylecum with plate on thoracic sternite 8 shallowly emarginate or slightly produced, without caudal extension. Maximum cl. 30 mm (Pl. 3h). Subtidal–slope (4–271 m). Temperate Northern Atlantic (Mediterranean), Temperate Northern W Pacific, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate Australia. 12 species. Species of Trachysalambria are thick-set pubescent prawns, some fished commercially. The genus was divided into three groups on the basis of the possession or not of epipods on pereopods 1–3 and the form of arthrobranch on thoracic somite 4 and the petasma by Sakaji and Hayashi (2003). They redescribed and presented a key to six species with an epipod on pereopods 1–3, the T. curvirostris (Stimpson, 1860) group. Chan et al. (2016) expanded on this, redescribing all species, adding new ones and provided a key to all species. They suggested on the basis of genetic data that the genus may be polyphyletic. Chanda (2018) rediagnosed Trachysalambria and redescribed Indian species.

Xiphopenaeus Smith, 1869 seabob Diagnosis. Carapace without transverse sutures; with longitudinal suture conspicuously overreaching hepatic spine. Antennular

flagella as long or longer than carapace. Pereopods 4 and 5 elongate, flagelliform, much longer than pereopod 3, dactyli multiarticulate. Maximum cl. 27 mm. Subtidal, shelf (1–70 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species. Xiphopenaeus is an important commercial shrimp in Brazil (see e.g. Reis et al. 2019). and elsewhere along the Atlantic and Pacific coasts of Central and South America. Two species separated by the Panama isthmus are recognised but the one in the Atlantic may involve a third as yet undescribed (Gusmão et  al. 2006). References Ahyong ST, Wilkens SL (2011) Aliens in the antipodes: non-indigenous marine crustaceans of New Zealand and Australia. In In the Wrong Place  – Alien Marine Crustaceans: Distribution, Biology and Impacts. (Eds Galil BS, Clark PF, Carlton JT) pp. 451–485. Springer Basel, Netherlands. Alcock A (1905) A revision of the “genus” Peneus, with diagnoses of some new species and varieties. Annals and Magazine of Natural History (ser. 7) 16, 508–532. Aydin I, Bakir K, Galil BS (2009) The first record of the jinga shrimp, Metapenaeus affinis (H. Milne Edwards, 1837) (Decapoda, Penaeidae) from the Mediterranean Sea. Crustaceana 82, 1091–1095. doi:10.1163/156854009X448907 Baldwin JD, Bass AL, Bowen BW, Clark WH (1998) Molecular phylogeny and biogeography of the marine shrimp, Penaeus. Molecular Phylogenetics and Evolution 10, 399–407. doi:10.1006/mpev.1998.0537 Burkenroad MD (1983) Natural classification of Dendrobranchiata, with a key to Recent genera. In: Schram FR (Ed.), Crustacean phylogeny. Crustacean Issues 6, 279–290. Chan T-Y (2012) A new genus of deep-sea solenocerid shrimp (Decapoda: Penaeoidea) from Papua New Guinea. Journal of Crustacean Biology 32, 489–495. doi:10.1163/193724012X626557 Chan T-Y, Tong J, Tam YK, Chu KH (2008) Phylogenetic relationships among the genera of the Penaeidae (Crustacea: Decapoda) revealed by mitochondrial 16S rRNA gene sequences. Zootaxa 1694, 38–50. doi:10.11646/zootaxa.1694.1.2 Chan T-Y, Cleva R, Chu KH (2016) On the genus Trachysalambria Burkenroad, 1934 (Crustacea, Decapoda, Penaeidae), with descriptions of three new species. Zootaxa 4150, 201–254. doi:10.11646/zootaxa.4150.3.1 Chan T-Y, Muchlisin ZA, Hurzaid A (2021) Verification of a pseudocryptic species in the commercially important tiger prawn Penaeus monodon Fabricius, 1798 (Decapoda: Penaeidae) from Aceh Province, Indonesia. Journal of Crustacean Biology 41, doi:10.1093/jcbiol/ruaa096. Chanda A (2018) A taxonomic study on trachypenaeid prawns with special reference to Indian records. Journal of Threatened Taxa 10, 11332–11338. doi:10.11609/jott.3765.10.2.11332-11338 Cheng J, Sha Z-l, Liu R-Y (2015) DNA barcoding of genus Metapenaeopsis (Decapoda: Penaeidae) and molecular phylogeny inferred from mitochondrial and nuclear DNA sequences. Biochemical Systematics and Ecology 61, 376–384. doi:10.1016/j.bse.2015.07.005 Cheng J, Chan T-Y, Zhang N, Sun S, Sha Z-l (2018) Mitochondrial phylogenomics reveals insights into taxonomy and evolution of Penaeoidea (Crustacea: Decapoda). Zoologica Scripta 47, 582–594. doi:10.1111/ zsc.12298 Cockcroft AC, Wooldridge T (1987) Reproduction and larval distribution of the penaeid prawn Macropetasma africanus (Balss) in Algoa Bay. African Zoology 22, 228–234. doi:10.1080/02541858.1987.11448050 Crosnier A (1986) Crustacés Décapodes: Penaeidae. Les espèces indoouest-pacifique du genre Parapeneaeus. In: Crosnier A (Ed.), Résultats

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des Campagnes MUSORSTOM I et II  – Philippines (1976, 1980). Tome 2. Mémoires du Muséum National d’Histoire Naturelle, Paris, ser. A (Zoologie) 133, 303–355. Crosnier A (1987) Les espèces indo-ouest-pacifiques d’eau profonde du genre Metapenaeopsis (Crustacea, Decapoda Penaeidae). Bulletin du Muséum National d’Histoire Naturelle Paris, 4e série, A 9, 409–453. Crosnier A (1991) Crustacea Decapoda: Les Metapenaeopsis indo-ouestpacifiques sans appareil stridulent (Penaeidae). Deuxième partie. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM. Vol. 9. Mémoires du Muséum National d’Histoire Naturelle, Paris, ser. A (Zoologie) 152, 155–297. Crosnier A (1994) Crustacea Decapoda: Les Metapenaeopsis indo-ouestpacifiques avec un appareil stridulant (Penaeidae). In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, Volume 12. Mémoires du Muséum National d’Histoire Naturelle, Paris 161, 255–337. Crosnier A (2006) Penaeopsis Bate, 1881 (Crustacea, Decapoda, Penaeidae) récoltées dans le Pacifique sud-ouest par les campagnes françaises depuis 1976. Description d’une espèce nouvelle. Zoosystema 28, 331–340. da Costa RC, Branco JO, Machado IF, Campos BR, Avila MG (2010) Population biology of shrimp Artemesia longinaris (Crustacea: Decapoda: Penaeidae) from the southern coast of Brazil. Journal of the Marine Biological Association of the United Kingdom 90, 663–669. doi:10.1017/ S002531540999124X Dall W (1957) A revision of the Australian species of Penaeinae (Crustacea Decapoda: Penaeidae). Australian Journal of Marine and Freshwater Research 8, 136–230. doi:10.1071/MF9570136 Dall W (2007) Recent molecular research on Penaeus sensu lato. Journal of Crustacean Biology 27, 380–382. doi:10.1651/S-2803.1 Dall W, Hall BJ, Rothlisberg P, Staples DJ (1990) The biology of the Penaeidae. Advances in Marine Biology 27, 1–489. Davie PJF (2002) Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells A, Houston WWK, Zoological Catalogue of Australia. Vol. 19.3A. CSIRO Publishing, Melbourne. De Grave S, Fransen CHJM (2011) Carideorum Catalogus: the Recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoölogische Mededeelingen 85, 195–589. FAO (2007–2019) Fisheries and Aquaculture Department. Fisheries and Aquaculture Fact Sheets. In: FAO Fisheries and Aquaculture Department [online]. Rome. Updated 22 May 2013. . Flegel TW (2007) The right to refuse revision in the genus Penaeus. Aquaculture 264, 2–8. doi:10.1016/j.aquaculture.2006.12.013 Flegel TW (2008) Confirmation of the right to refuse revision in the genus Penaeus. Aquaculture 280, 1–4. doi:10.1016/j.aquaculture.2008.04.029 Fujino T (1975) Funchalia sagamiensis sp. nov. from Central Japan, with discussion of the generic characters (Decapoda, Natantia, Penaeidae). Crustaceana 28, 200–210. doi:10.1163/156854075X01071 Grippa G (1987) A revision of gen. Pelagopenaeus Burkenroad, 1934 and Funchalia Johnson, 1867 (Crustacea, Decapoda, Penaeidea). Investigaciones Pesqueras 51(suppl. 1), 73–85. Gusmão J, Laoski C, Sole-Cava AM (2000) A new species of Penaeus (Crustacea: Penaeidae) revealed by allozyme and cytochrome oxidase I analysis. Marine Biology 137, 435–446. doi:10.1007/s002270000365 Gusmão J, Lazoski C, Monteiro FA, Solé-Cava AM (2006) Cryptic species and population structuring of the Atlantic and Pacific seabob shrimp species, Xiphopenaeus kroyeri and Xiphopenaeus riveti. Marine Biology 149, 491–502. doi:10.1007/s00227-005-0232-x Holthuis LB (1980) FAO species catalogue. Vol. 1. Shrimps and prawns of the world. An annotated catalogue of species of interest to fisheries. FAO Fisheries Synopsis 125, 1–261.

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Jory D (2019) Shrimps. In Aquaculture: Farming Aquatic Animals and Plants, 3rd Edition. (Eds Lucas JS, Southgate PC, Tucker CS) pp. 499–526. Wiley-Blackwell. Li X, Xu Y, Kou Q (2014) Molecular phylogeny of Parapenaeopsis Alcock, 1901 (Decapoda: Penaeidae) based on Chinese materials and 16S rDNA and COI sequence. Journal of Ocean University of China 13, 104–114. doi:10.1007/s11802-014-2272-7 Ma KY, Chan T-Y, Chu KH (2009) Phylogeny of penaeoid shrimps (Decapoda: Penaeoidea) inferred from nuclear protein-coding genes. Molecular Phylogenetics and Evolution 53, 45–55. doi:10.1016/j. ympev.2009.05.019 Ma KY, Chan T-Y, Chu KH (2011) Refuting the six-genus classification of Penaeus s.l. (Dendrobranchiata, Penaeidae): a combined analysis of mitochondrial and nuclear genes. Zoologica Scripta 40, 498–508. doi:10.1111/j.1463-6409.2011.00483.x Maggioni R, Rogers AD, Maclean N, D’Incao F (2001) Molecular phylogeny of Western Atlantic Farfantepenaeus and Litopenaeus shrimp based on mitochondrial 16S partial sequences. Molecular Phylogenetics and Evolution 18, 66–73. doi:10.1006/mpev.2000.0866 McLaughlin PA, Lemaitre R, Ferrari FD, Felder DL, Bauer RT (2008) A reply to T.W. Flegel. Aquaculture 275, 370–373. doi:10.1016/j. aquaculture.2007.12.020 Miquel JC (1982) Le genre Metapenaeus (Crustacea, Penaeidae): taxonomie, biologie et pêches mondiales. Zoölogische Verhandelingen 195, 1–137. Pérez Farfante I (1971) A key to the American Pacific shrimps of the genus Trachypenaeus (Decapoda, Penaeidae), with the description of a new species. Fishery Bulletin 69, 635–646. Pérez Farfante I (1972) Tanypenaeus caribeus, a new genus and species of the shrimp family Penaeidae (Crustacea, Decapoda) from the Caribbean sea. Bulletin of Marine Science 22, 185–195. Pérez Farfante I (1980) Revision of the penaeid shrimp genus Penaeopsis (Crustacea: Decapoda). Fishery Bulletin 77, 721–763. Pérez Farfante I, Kensley B (1997) Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175, 1–233. Primavera JH (1998) Mangroves as nurseries: shrimp populations in mangrove and non-mangrove habitats. Estuarine, Coastal and Shelf Science 46, 457–464. doi:10.1006/ecss.1997.0275 Racek AA, Dall W (1965) Littoral Penaeinae (Crustacea Decapoda) from northern Australia, New Guinea and adjacent waters. Verhandelingen der Koninklijke Akademie van Wetenschappen 56, 1–119, pls 1–13. Ramirez JL, Simbine L, Marques CG, Zelada-Mázmela E, Reyes-Flores LE, et al. (2021) DNA barcoding of Penaeidae (Decapoda; Crustacea): non-distance-based species delimitation of the most economically important shrimp family. Diversity 13, 460. doi:10.3390/d13100460 Reis JJDC, Freire KMF, Da Rosa LC, Barreto TMRDR, Pauly D (2019) Population dynamics of Atlantic seabob Xiphopenaeus kroyeri (Decapoda: Penaeidae) off the state of Sergipe, north-eastern Brazil. Journal of the Marine Biological Association of the United Kingdom 99, 143–153. doi:10.1017/S0025315417001916 Robalino J, Wilkins B, Bracken-Grissom HD, Chan T-Y, O’Leary MA (2016) The origin of large-bodied shrimp that dominate modern global aquaculture. PLoS One 11, e0158840. doi:10.1371/journal. pone.0158840 Sakai K, Shinomiya S (2011) Preliminary report on eight new genera formerly attributed to Parapenaeopsis Alcock, 1901, sensu lato (Decapoda, Penaeidae). Crustaceana 84, 491–504. doi:10.1163/​ 001121611X557037 Sakaji H, Hayashi K-I (2003) A review of the Trachysalambria curvirostris species group (Crustacea: Decapoda: Penaeidae) with description of a new species. Species Diversity 8, 141–174. doi:10.12782/specdiv.8.141

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Marine Decapod Crustacea

Santamaría J, Carbajal-Enzian P, Clemente S, Alemán S, Ordinola E (2018) Guía ilustrada para reconocimiento de langostinos y otros crustáceos con valor comercial en el Perú. Instituto del Mar del Perú (Imarpe), Lima. Shinomiya S, Sakai K (2006) Three new species of the genus Megokris (Decapoda, Penaeidae) from Hall’s and Motoh’s collections. Crustaceana 79, 1251–1268. doi:10.1163/156854006778859579 Shinomiya S, Sakai K (2016) A review of the genus Batepenaeopsis, with the description of a new species, B. moresbyensis sp. nov. (Decapoda, Penaeidae). Crustaceana 89, 495–513. doi:10.1163/15685403-00003537 Tavares C, Gusmão J (2016) Description of a new Penaeidae (Decapoda: Dendrobranchiata) species, Farfantepenaeus isabelae sp. nov. Zootaxa 4171, 505–516. doi:10.11646/zootaxa.4171.3.6 Thomas MM (1972) Trachypenaeopsis minicoyensis sp. nov. (Penaeidae, Decapoda) from the Laccadive Sea. Indian Journal of Fisheries 19, 116–121. Tong JG, Chan T-Y, Chu KH (2000) A preliminary phylogenetic analysis of Metapenaeopsis (Decapoda: Penaeidae) based on mitochondrial DNA sequences of selected species from the Indo-West Pacific. Journal of Crustacean Biology 20, 541–549. doi:10.1163/20021975-99990070 Wang T-W, Chan T-Y, Chan BKK (2013) Diversity and community structure of decapod crustaceans at hydrothermal vents and nearby deepwater fishing grounds off Kueishan Island, Taiwan: a high biodiversity deep-sea area in the NW Pacific. Bulletin of Marine Science 89, 505–528. doi:10.5343/bms.2012.1036 WoRMS Editorial Board (2022) World Register of Marine Species. Available from at VLIZ. Accessed May 2022. Yang C-H, Sha Z, Chan T-Y, Liu R (2015) Molecular phylogeny of the deep-sea penaeid shrimp genus Parapenaeus (Crustacea: Decapoda: Dendrobranchiata). Zoologica Scripta 44, 312–323. doi:10.1111/ zsc.12097

Sicyoniidae Ortmann, 1898 rock shrimp, ridgeback shrimp Figure 3.6, Plate 4b–g The rigid carapace is the most obvious feature distinguishing sicyoniids from all other dendrobranchiates. The common names ridgeback shrimp and rock shrimp are applied to some species. The legs are usually shorter and stronger than in other prawns and they lack endopods on pleopods 3–5. All are benthic species, the family ranging from intertidal to slope depths. Males are smaller than females and have relatively small chelipeds. Sicyonia is the only genus but Crosnier (2003) recognised eight groups of species among 31 that he dealt with from the Indo-West Pacific. His work contains colour photographs and a key to Indo-West Pacific species. Hanamura and Wadley (1998) also provided a key to Indo-West Pacific species. Diagnosis. Body typically shrimp-like; integument rigid and stony. Rostrum elongate, blade-like with dorsal and sometimes ventral teeth. Postorbital spine absent; with 3 or more postrostral teeth. With at least 3 branchiae on some

Fig. 3.6.  Sicyoniidae. Sicyonia carinata (Brünnich, 1768).

thoracic somites, at least 11 on each side. Prosartesma (mesial process on antennule article 1) reduced to setose boss. Sicyonia H. Milne Edwards, 1830 Maximum cl. 28 mm. Intertidal–slope (0–936 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Tropical Eastern Pacific, Temperate Australasia. 52 species. References Crosnier A (2003) Sicyonia (Crustacea, Decapoda, Penaeoidea, Sicyoniidae) de l’indo-ouest Pacifique. Zoosystema 25, 197–350. Hanamura Y, Wadley V (1998) A new species of the rock shrimp genus Sicyonia (Decapoda, Sicyoniidae) from south-eastern Australia, with a key to the Indo-West Pacific species. Crustaceana 71, 700–711. doi:10.1163/156854098X00699

Solenoceridae Wood-Mason in Wood-Mason & Alcock, 1891 Figure 3.7, Plate 4h–j Solenocerid shrimps are rarely found over the continental shelf but prefer waters over the slope, at depths of hundreds of metres. Some are mesopelagic or bathypelagic while others are benthic. Members of the family are distinguished from other dendrobranchiates by the possession of a postorbital spine and usually a complete cervical groove (Pérez Farfante and Kensley 1997). Half of the known species belong to the genus Solenocera (De Grave and Fransen 2011). Dall’s (1999) review of the Australian species diagnosed the family, discussed anatomical features, and provided a key to nine genera. He also reviewed the literature and gave keys to Indo-West Pacific species. Chan (2012) added another genus and species, and presented a molecular phylogeny. While solenocerids are typical dendrobranchiates, some species of Solenocera and one of Mesopenaeus have

3 – Dendrobranchiata – prawns and shrimps

dendrolamellate gills in which the secondary and tertiary processes are flattened (Tavares and Serejo 2019). Several species are of commercial interest. Diagnosis. Body typically shrimp-like; integument soft, stiff but not rigid. Rostrum elongate, blade-like with dorsal

39

and sometimes ventral teeth. Postorbital spine absent; with 2 or more postrostral teeth. With at least 3 branchiae on some thoracic somites, at least 11 on each side. Prosartesma (mesial process on antennule article 1) reduced to setose boss (well developed only in Maximiliaeus).

Key to genera of Solenoceridae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. – 9. –

Telson with small movable lateral spiniform setae (Fig. 3.7i, j)������������������������������������������������������������������������������������������������2 Telson without movable lateral spiniform setae (Fig. 3.7k)�������������������������������������������������������������������������������������������������������4 Postcervical spine present, close to hepatic spine (Fig. 3.7d)����������������������������������������������������������������������������������� Gordonella Postcervical spine absent�����������������������������������������������������������������������������������������������������������������������������������������������������������������3 Pleon lateral surface more or less covered with complex ridges. Telson with pair of fixed lateral spines, 3 pairs of movable lateral spines (Fig. 3.7i)����������������������������������������������������������������������������������������������������������������� Haliporus Pleon lateral surface weakly carinate (Fig. 3.7b). Telson without fixed lateral spines, with minute movable spines���������������������������������������������������������������������������������������������������������������������������������������������������������������������� Maximiliaeus Both antennular flagella subcylindrical���������������������������������������������������������������������������������������������������������������������������������������5 One or both antennular flagella flattened������������������������������������������������������������������������������������������������������������������������������������9 Pereopods 4 and 5 slightly longer than pereopod 3 (not flagelliform)������������������������������������������������������������������������������������6 Pereopod 5 flagelliform, longer than pereopod 3�����������������������������������������������������������������������������������������������������������������������8 Orbital spine present (Fig. 3.7g)��������������������������������������������������������������������������������������������������������������������������������������� Pleoticus Orbital spine absent�������������������������������������������������������������������������������������������������������������������������������������������������������������������������7 Suprahepatic spine present (Fig. 3.7a)���������������������������������������������������������������������������������������������������������������������� Haliporoides Suprahepatic spine absent (Fig. 3.7c)�������������������������������������������������������������������������������������������������������������������� Cryptopenaeus Pereopod 5 only flagelliform, much longer than pereopod 4���������������������������������������������������������������������������� Hadropenaeus Pereopods 4 and 5 both flagelliform, longer than pereopod 3�����������������������������������������������������������������������Hymenopenaeus Lower antennular flagellum flattened�������������������������������������������������������������������������������������������������������������������� Mesopenaeus Both antennular flagella flattened��������������������������������������������������������������������������������������������������������������������������������Solenocera

Fig. 3.7.  Solenoceridae. Habitus: a, Haliporides sibogae Kensley, Tranter & Griffin, 1987. Carapace: b, Maximiliaeus odoceros Chan, 2012; c, Cryptopenaeus crosnieri Pérez Farfante & Kensley, 1985; d, Gordonella kensleyi Crosnier, 1989; e, Haliporus thetis Faxon, 1893; f, Hymenopenaeus halli Bruce, 1966; g, Pleoticus muelleri (Bate, 1888); h, Solenocera faxoni De Man, 1907. Telson: i, Haliporus; j, Maximiliaeus; k, Solenocera.

40

Marine Decapod Crustacea

Cryptopenaeus de Freitas, 1979 Diagnosis. Lateral carapace with antennal, postantennal and branchiocardiac carinae. Rostrum short, triangular. Telson without lateral spiniform setae. Pereopods 4, 5 slightly longer than pereopod 3 (not flagelliform). Maximum cl. 48 mm. Slope (200–540 m). Western and Central Indo-Pacific, Temperate Australasia. 5 species (Dall 1999: key to 4 species).

Gordonella Tirmizi, 1960 Diagnosis. Postcervical spine present, close to hepatic spine; lateral carapace with antennal, postantennal, vertical cervical, hepatic, branchiocardiac, sublateral, subhepatic-branchiostegal carinae. Rostrum short, triangular. Telson with 4 pairs of lateral spiniform setae. Pereopods 4, 5 slightly longer than pereopod 3 (not flagelliform). Maximum cl. 58 mm. Bathyal-bathypelagic (750–2798 m). GOODS bathyal provinces: Indian Ocean, West Pacific Ocean, South Atlantic Ocean. 3 species (Crosnier 1989; Dall 1999).

Hadropenaeus Pérez Farfante, 1977 trident shrimp

Hymenopenaeus Smith, 1882 veiled shrimp Diagnosis. Lateral carapace with hepatic, long branchiocardiac, posthepatic and submarginal carinae. Telson without lateral spiniform setae. Pereopods 4, 5 both flagelliform, longer than pereopod 3. Maximum cl. 44 mm. Slope–abyssal (200–4792 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Tropical Eastern Pacific, Temperate Southern Africa. 17 species (Crosnier and Dall 2004; Dall 1999; Pérez Farfante 1977).

Maximiliaeus Chan, 2012 Diagnosis. Carapace middorsal ridge with 2 postcervical teeth; lateral carapace with sharp antennal, postantennal, nearly straight branchiocardiac, fused hepatic and posthepatic, branchiostegal and submarginal carinae. Telson with 1 pair of minute lateral spiniform setae. Pereopods 4, 5 slightly longer than pereopod 3 (not flagelliform). Maximum cl. 49 mm (Pl. 4i). Slope (403–485 m). Central Indo-Pacific. 1 species (Chan 2012).

Mesopenaeus Pérez Farfante, 1977

Diagnosis. Lateral carapace with hepatic carina, without branchiocardiac, posthepatic and submarginal carinae. Telson without lateral spiniform setae. Antennular both flagella subcylindrical, or lower flagellum flattened. Pereopod 5 flagelliform, much longer than pereopod 4. Maximum cl. 26 mm. Shelf, slope (100–698 m). Temperate Northern and Tropical Atlantic, Western and Central Indo-Pacific, Temperate Australasia. 4 species (Dall 1999: Indo-West Pacifc species; Pérez Farfante 1977: Atlantic species).

Diagnosis. Orbital spine present; lateral carapace with hepatic, without branchiocardiac, posthepatic and submarginal carinae. Telson without lateral spiniform setae. Antennular lower flagellum flattened. Pereopods 4 and 5 elongate, flagelliform, much longer than pereopod 3. Maximum cl. 25 mm. Subtidal–slope (11–915 m). Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific. 3 species (Crosnier 1986: Indo-West Pacific species; Pérez Farfante 1977: Atlantic species).

Haliporoides Stebbing, 1914

Pleoticus Bate, 1888

jack-knife shrimp, royal red prawn

red shrimp

Diagnosis. Suprahepatic spine minute, just posterior to cervical groove above hepatic spine; lateral carapace with hepatic, branchiocardiac and submarginal carinae. Telson without lateral spiniform setae. Pereopods 4, 5 slightly longer than pereopod 3 (not flagelliform). Maximum cl. 101 mm. Shelf, slope (100–1336 m). Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate South America, Temperate Australasia. 4 species. Some species are fished commercially. Of the four species, one has three subspecies and another two (Dall 1999; De Grave and Fransen 2011; Pérez Farfante 1977).

Diagnosis. Orbital spine present; lateral carapace with hepatic, with or without branchiocardiac, without posthepatic, with submarginal carinae. Telson without lateral spiniform setae. Pereopods 4 and 5 elongate, flagelliform, much longer than pereopod 3. Maximum cl. 62 mm. Subtidal–slope (2–1000 m). Tropical Atlantic, Western IndoPacific, Temperate South America. 3 species (Pérez Farfante 1977). One species, the royal red shrimp P. robustus (Smith, 1885) has been the target of a fishery in the Atlantic (Anderson and Lindner 1971).

Haliporus Bate, 1881 Diagnosis. Lateral carapace with antennal, hepatic, long branchiocardiac, submarginal carinae. Telson with 1 pair of fixed, 3 pairs of movable lateral spiniform setae. Pereopods 4, 5 slightly longer than pereopod 3 (not flagelliform). Maximum cl. 46 mm (Pl. 4h). Slope–abyssal (300–5700 m). GOODS abyssal provinces: Indian Ocean, E Pacific basins, Central Pacific Ocean, N Central Pacific. 3 species (Crosnier 1989: key to species; Dall 1999: key to species).

Solenocera Lucas, 1849 mud shrimp, Kolibri shrimp, ridgeback shrimp, flower shrimp, comb shrimp Diagnosis. Orbital spine present, or absent; lateral carapace with hepatic, with or without branchiocardiac, without posthepatic, with submarginal carinae. Telson without lateral spiniform setae. Antennular flagella both flattened. Pereopods 4 and 5 elongate, flagelliform, much longer than pereopod 3. Maximum cl. 54 mm (Pl. 4j).

3 – Dendrobranchiata – prawns and shrimps

Subtidal–slope (15–900 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Central and Eastern IndoPacific, Tropical Eastern Pacific, Temperate Southern Africa. 38 species of which 28 live in the Indo-West Pacific (Dall 1999: key to 22 Indo-West Pacific species; De Grave and Fransen 2011).

Species of Solenocera are benthic on muddy sediments of the continental shelf and slope. The respiratory tube fashioned from the modified antennules suggests the shrimps bury themselves in the mud with the pleon flexed upwards. As the variety of common names used indicates, several species are fished commercially and for this reason are well studied (e.g. Li et al. 2016; Villalobos-Rojas and Wehrtman 2018). References Anderson WW, Lindner MJ (1971) Contributions to the biology of the royal red shrimp, Hymenopenaeus robustus Smith. Fishery Bulletin 69, 313–336. Chan T-Y (2012) A new genus of deep-sea solenocerid shrimp (Decapoda: Penaeoidea) from Papua New Guinea. Journal of Crustacean Biology 32, 489–495. doi:10.1163/193724012X626557 Crosnier A (1986) Sur deux espèces du genre Mesopenaeus (Penaeoidea: Solenoceridae) de l’Océan Indien: M. brucei sp. nov. et M. mariae Pérez Farfante & Ivanov, 1982. Indo-Malayan Zoology 3, 19–25. Crosnier A (1989) Contribution à l’étude des genres Haliporus Bate, 1881 et Gordonella Tirmizi, 1960 (Crustacea Decapoda Penaeoidea). Description de deux espèces nouvelles. Bulletin du Muséum National d’Histoire Naturelle Paris, 4e série, A 10, 563–601. Crosnier A, Dall W (2004) Redescription of Hymenopenaeus obliquirostris (Crustacea, Decapoda, Penaeoidea, Solenoceridae) and descriptions of two new species of Hymenopenaeus from the Indo-West Pacific. Zootaxa 600, 1–26. doi:10.11646/zootaxa.600.1.1 Dall W (1999) Australian species of Solenoceridae (Penaeoidea: Decapoda). Memoirs of the Queensland Museum 43, 553–587. De Grave S, Fransen CHJM (2011) Carideorum Catalogus: the Recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoölogische Mededeelingen 85, 195–589. Li HY, Ling JZ, Yan LP, Cheng JH, Hu F (2016) Feeding ecology of the mud shrimp Solenocera melantho in the East China Sea. Marine Ecology (Berlin) 37, 380–391. doi:10.1111/maec.12290 Pérez Farfante I (1977) American solenocerid shrimps of the genera Hymenopenaeus, Haliporoides, Pleoticus, Hadropenaeus new genus, and Mesopenaeus new genus. Fishery Bulletin 75, 261–346. Pérez Farfante I, Kensley B (1997) Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175, 1–233. Tavares C, Serejo CS (2019) A new pattern of dendrobranch gill within the family Solenoceridae (Crustacea: Decapoda: Dendrobranchiata). Zootaxa 4571, 439–445. doi:10.11646/zootaxa.4571.3.12 Villalobos-Rojas F, Wehrtman I (2018) Reproductive biology of the commercially exploited kolibri shrimp, Solenocera agassizii (Decapoda: Solenoceridae), from the Pacific coast of Costa Rica, with considerations for its management. Revista de Biologia Tropical 66 (suppl.) 1, S92–S107.

Superfamily Sergestoidea Dana, 1852 There are two families: Luciferidae and Sergestidae. Diagnosis. Never more than 8 (sometimes with at most 2) branchiae (gills) on each side.

41

Luciferidae De Haan, 1849 Figure 3.8 These tiny prawns appear in plankton, sometimes in large numbers, from shallow water to the surface waters of all temperate and tropical oceans. Their elongate shape, long neck, and dominant eyes make them immediately recognisable. Because they lack so many basic dendrobranchiate features, like gills, their relationships to other families and even their inclusion in Dendrobranchiata itself have been questioned (Burkenroad 1983; Pérez Farfante and Kensley 1997). The presence of a petasma in males is the only feature uniting luciferids with dendrobranchiates but euphausiaceans possess this feature too. Loss of pereopods 4 and 5 is shared with some sergestids but is a dubious synapomorphy. Another unusual feature of luciferids is that females brood their eggs on pereopod 3; all other dendrobranchiates are broadcast spawners. The major taxonomic study of Hansen (1919) remained unchallenged until a significant revision by Vereshchaka et al. (2016). Their work took a phylogenetic approach based on morphology and reached the same conclusion as Hansen (1919), that the family can be divided in two. They rediagnosed Lucifer and introduced a new genus name for Hansen’s ‘Group B’. Hansen (1919) provided keys to males and females of all species while the keys to genera and species of Vereshchaka et al. (2016) relied on males. The latter authors provided detailed illustrations of the petasma of all species. Vereshchaka (2017) suggested that Acetes was more related to Luciferidae than to Sergestidae where the genus is usually placed. He also noted that these and Sicyonella (Sergestidae) are epipelagic (surface dwellers). Species of Belzebub occupy neritic waters (above the continental shelf) while those of Lucifer are in the open ocean. Vereshchaka’s family names are nomina nuda. Diagnosis. Body strongly laterally compressed; integument soft, stiff but not rigid. Rostrum absent. Postorbital spine absent. Without branchiae. Belzebub Vereshchaka, Olesen & Lunina, 2016 Diagnosis. Posterior ventral process on pleonite 6 of male nearly straight, tapering, apically subacute (Fig. 3.8a). Eyestalks of moderate length, subcylindrical, not reaching end of scaphocerite. Petasma sheath narrow, tapering, supported by strong chitinous ribs, armed with apical scales or ridged, with additional plate-like structures; processus ventralis spiniform, without apical pincer (Fig. 3.8c). Maximum tl. 11.6 mm. Neritic (0–100 m). Cosmopolitan in all tropical to temperate oceans. 5 species (Vereshchaka et al. 2016: key to species). Belzebub hanseni (Nobili, 1906) has entered the Mediterranean Sea from the Red Sea via the Suez Canal (Galil et al. 2002).

42

Marine Decapod Crustacea

Fig. 3.8.  Luciferidae. Pleonite 6, uropod and telson of male: a, Belzebub; c, Lucifer. Petasma (left), processus ventralis (right): b, Belzebub; d, Lucifer. e, Lucifer typus H. Milne Edwards, 1837.

Lucifer Thompson, 1829 Diagnosis. Posterior ventral process on pleonite 6 of male curved, not tapering, apically obtuse (Fig. 3.8b). Eyestalks elongated, conical, reaching near end of scaphocerite. Petasma sheath wide, entirely armed with transverse chitinous ribs along inner margin, without additional plate-like structures; processus ventralis lamellar, with apical pincer (Fig. 3.8d). Maximum tl. 12.4 mm. Oceanic (0–100 m). Cosmopolitan in all tropical to temperate oceans. 2 species (Vereshchaka et al. 2016: key to species). References Burkenroad MD (1983) Natural classification of Dendrobranchiata, with a key to Recent genera. In: Schram FR (Ed.), Crustacean phylogeny. Crustacean Issues 6, 279–290. Galil B, Froglia CPN, Noël P (2002) CIESM Atlas of exotic species in the Mediterranean. Volume 2. Crustaceans: decapods and stomatopods. CIESM Publishers, Monaco. Hansen HJ (1919) The Sergestidae of the Siboga-Expedition. SibogaExpéditie 38, 1–65, pls 1–7. Pérez Farfante I, Kensley B (1997) Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175, 1–233. Vereshchaka AL (2017) The shrimp superfamily Sergestoidea: a global phylogeny with definition of new families and an assessment of the pathways into principal biotopes. Royal Society Open Science. 4, 170221 (11 pp.). Vereshchaka AL, Olesen J, Lunina AA (2016) A phylogeny-based revision of the family Luciferidae (Crustacea: Decapoda). Zoological Journal of the Linnean Society 178, 15–32. doi:10.1111/zoj.12398

Sergestidae Dana, 1852 Figures 3.9–3.11, Plate 4a The combination of a short rostrum and reduced last pereopods serve to characterise sergestid shrimps. Sergestids differ from other dendrobranchiates in having swimming legs fringed with long setae, although the number of legs may be

fewer than the expected five pairs. Most are small pelagic species from offshore environments over and beyond the continental shelf. Only species of Acetes are found in bays and shallow water. Hansen (1919, 1922) pioneered the systematics of Sergestidae. Pérez Farfante and Kensley (1997) recognised six genera but Judkins and Kensley (2008) divided one, Sergestes, into six genera. Vereshchaka (2009) further divided Sergestes and Sergia into species groups and provided keys to all species. He detailed sergestid anatomy and characters of taxonomic importance and mapped the geographic distribution of the groups. Later, Vereshchaka et al. (2014) gave these groups generic status, genera with the -sergestes and -sergia suffixes, all of which they diagnosed in detail. That paper and Vereshchaka (2017) discussed cladistic relationships between the genera, discussed ecological and geographic distribution and related the possession of photophores and some other characters to the clades (genera). The key below and generic diagnoses are based on their work Species of genera related to Sergia, but not of Sergia itself, have the ability to produce light using dermal photophores which are distinct bioluminescent organs, sometimes with external lenses associated with an internal pigment layer (Nowel et al. 2002). Occurring on various parts of the body such as the carapace, pleon, scaphocerite and uropod, photophores appear in preserved specimens as small or minute spots. Photophores bearing cuticular lenses appear to have a convex bulge when viewed side-on. On the other hand, species of Sergestes and related genera use bioluminescence produced by the organs of Pesta as countershading to help them remain cryptic to visual predators (Latz 1995). Organs of Pesta are internal light-producing organs, pigmented and modified areas of the gastrohepatic gland appearing as

3 – Dendrobranchiata – prawns and shrimps

small, yellow pea-like structures that can be seen through the carapace. Preserved specimens of Sergia and related genera of Vereshchaka et al. (2014) are difficult to place to genus because of the reliance on the luminescent organs that are not obvious to the novice. Differences between genera, especially those related to Sergia and Sergestes, rely too in part on the structure of the petasma, the male genital structure comprising the enlarged and coupled endopods of pleopods 1 (Fig.  3.10l, m). It is variously lobed, comprising the lobus armatus (LA) typically well developed, lobus connectens (LC), lobus inermis (LI) typically straight and slender, lobus terminalis (LT) typically entire and well developed, processus uncifer (PU) typically with a hook, and processus ventralis (PV). The key

43

to genera below omits petasma characters but these are included in diagnoses, typical states excepted. Vereshchaka et al.’s (2014) ecological-bathymetric characterisation of these two groups of genera is followed here: benthopelagic (living above shelves, continental slopes, and seamounts); epipelagic (living in upper 200 m); interzonal mesopelagic migrants living in the low mesopelagic zone (400–800 m) during the day and ascending to the epipelagic zone (100–200 m) at night; and bathypelagic (800–3000 m). Diagnosis. Body typically shrimp-like; integument soft, stiff but not rigid. Rostrum short, erect blade, sometimes with few teeth. Postorbital spine present. With no more than 2 branchiae per thoracic somite, at most 8 on each side.

Key to genera of Sergestidae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. – 9. – 10.

Pereopods 4 and 5 absent�������������������������������������������������������������������������������������������������������������������������������������������������������Acetes Pereopods 4 and 5 present���������������������������������������������������������������������������������������������������������������������������������������������������������������2 Pereopods 4 and 5 with 1–5 or 7 articles��������������������������������������������������������������������������������������������������������������������������������������3 Pereopods 4 and 5 with 6 articles��������������������������������������������������������������������������������������������������������������������������������������������������4 Pereopod 4 of male with 3 articles, of female with 5 articles. Pereopod 5 of male with 1 article, of female with 3 articles (Fig. 3.9d)��������������������������������������������������������������������������������������������������������������������������������������������������� Peisos Pereopods 4 and 5 of 7 articles (Fig. 3.9e)��������������������������������������������������������������������������������������������������������������������� Sicyonella Maximum height of rostrum at midpoint (Fig. 3.9b)����������������������������������������������������������������������������������������������� Petalidium Maximum height of rostrum near tip�������������������������������������������������������������������������������������������������������������������������������������������5 Organ of Pesta present (Fig. 3.10a). Body semi-transparent in live specimen, without dermal photophores�����������������������������������������������������������������������������������������������������������������������������������������������Sergestes-group … 6 Organ of Pesta absent (Fig. 3.11a). Body opaque in live specimen, or, if semitransparent, with dermal photophores������������������������������������������������������������������������������������������������������������������������������������������������� Sergia-group … 12 Uropodal exopod outer margin with tooth, not setose along margin proximal to tooth�����������������������������������������������������7 Uropodal exopod outer margin without tooth, setose along at least part of margin of proximal article��������������������������9 Antennule article 1 elongate, ≥ 1.5 times as long as article 3 (Fig. 3.10b). Scaphocerite distal tooth not overreaching blade. Maxilliped 3 sexually dimorphic. Arthrobranch: posterior lobe above pereopod 3 dendritic���������������������������������������������������������������������������������������������������������������������������������������������������������������������Eusergestes Antennule article 1 not elongate, 1.5 times as long as article 3 (Fig. 3.10c). Scaphocerite distal tooth overreaching blade. Maxilliped 3 not sexually dimorphic. Arthrobranch: posterior lobe above pereopod 3 lamellar��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������8 Rostrum triangular, not reaching middle of eyestalk (Fig. 3.10c). Maxilliped 1 endopod of 3 articles��������������� Sergestes Rostrum elongate, considerably overreaching middle of eyestalk (Fig. 3.10e). Maxilliped 1 endopod of 2 articles��������������������������������������������������������������������������������������������������������������������������������������������������������������� Cornutosergestes Maxilliped 3 not elongate, less than twice as long as carapace. Pereopod 2 chela with long setae (Fig. 3.10k). Arthrobranch: posterior lobe above pereopod 3 dendritic�������������������������������������������������������������������������������Deosergestes Maxilliped 3 elongate, more than twice as long as carapace. Pereopod 2 chela without long setae. Arthrobranch: posterior lobe above pereopod 3 lamellar��������������������������������������������������������������������������������������������������10 Rostrum with vertical frontal margin and beak-like terminal tooth (Fig. 3.10d). Ocular papilla prominent. Maxilliped 3 2.8 times as long as carapace. Pereopod 2 without distally curved hooks on ischium, without protrusion on merus, chela with unequal fingers (Fig. 3.10j). Pereopod 3 with strong curved spines proximal to tufts of long setae on propodus. Pereopod 5 with distal article setose along both margins���������������������������������������������������������������������������������������������������������������������������������������������������������������������Allosergestes

44

Marine Decapod Crustacea

Fig. 3.9.  Sergestidae. Carapace: a, Acetes sibogae Hansen, 1919; b, Petalidium foliaceum Bate, 1881; c, d, Peisos petrunkevitchi Burkenroad, 1945 (with female pereopod 5); e–g, Sicyonella inermis (Paul’son, 1875) (habitus, anterior dorsal, tailfan).

Fig. 3.10.  Sergestidae. Sergestes and related genera. a, Parasergestes armatus (Krøyer, 1855). Eyes, antennule, antenna: b, Eusergestes; c, Sergestes. Rostrum, anterior carapace of type species: d, Allosergestes sargassi (Ortmann, 1893); e, Cornutosergestes cornutus (Krøyer, 1855); f, Deosergestes corniculum (Krøyer, 1855); g, Neosergestes edwardsii (Krøyer, 1855); h, Parasergestes armatus (Krøyer, 1855); i, Sergestes atlanticus H. Milne Edwards, 1830. Chela of pereopod 2: j, Allosergestes; k, Deosergestes. Petasma (see text for abbreviations of lobes): l, Deosergestes; m, Eusergestes. Uropodal exopod: n, Eusergestes; o, Deosergestes; p, Neosergestes.

3 – Dendrobranchiata – prawns and shrimps

45



Rostrum with oblique frontal margin, without beak-like terminal tooth (Fig. 3.10a, g, h). Ocular papilla ill-defined. Maxilliped 3 2.0–2.8 times as long as carapace. Pereopod 2 with distally curved hooks on ischium and protrusion on merus, chela with subequal fingers. Pereopod 3 without strong curved spines proximal to tufts of long setae on propodus. Pereopod 5 with distal article setose along 1 margin (Fig. 3.10a)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������11 11. Maxilliped 3 dactylus subdivided into 4. Pereopod 1 with strong movable spines on ischium. Uropodal exopod outer margin partly setose����������������������������������������������������������������������������������������������������������������������Parasergestes – Maxilliped 3 dactylus subdivided into 6. Pereopod 1 without strong movable spines on ischium. Uropodal exopod outer margin entirely setose��������������������������������������������������������������������������������������������������������������������Neosergestes 12. Integument membranous, dermal photophores absent����������������������������������������������������������������������������������������������������Sergia – Integument firm, dermal photophores present�������������������������������������������������������������������������������������������������������������������������13 13. Dermal photophores without lens, visible as opaque spots (Fig. 3.11c, e, g)�������������������������������������������������������������������������14 – Dermal photophores with lens (Fig. 3.11b, d, f, h)���������������������������������������������������������������������������������������������������������������������16

Fig. 3.11.  Sergestidae. Sergia and related genera. a, Prehensilosergia prehensilis (Bate, 1881). Carapace, scaphocerite, uropod of type species: b, Challengerosergia challengeri (Hansen, 1903); c, Gardinerosergia gardineri (Kemp, 1913); d, Lucensosergia lucens (Hansen, 1922); e, Phorcosergia phorca (Faxon, 1893); f, Prehensilosergia prehensilis (Bate, 1881); g, Robustosergia robusta (Smith, 1882); h, Scintillosergia scintillans (Burkenroad, 1940); i, Sergia tenuiremis (Krøyer, 1855).

46

14. – 15. – 16. – 17. – 18. –

Marine Decapod Crustacea

Photophores large, partly fused organs, arranged in 2 rows on scaphocerite and triangular patch on uropodal exopod (Fig. 3.11e)�������������������������������������������������������������������������������������������������������������������������������� Phorcosergia Photophores small, not fused, arranged in row on scaphocerite, and row (sometimes 1 organ) on uropodal exopod�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������15 Ocular papilla developed (>0.3 times as long as wide). Photophores on uropodal exopod close to inner margin (Fig. 3.11c)�������������������������������������������������������������������������������������������������������������������������������������������� Gardinerosergia Ocular papilla rudimentary ( 5 times as long as short palm, fringed with long setae (Fig. 6.53j). Maxilliped 2 with last 2 articles side by side (Fig. 6.53i)���������������������Stylodactyloidea … Stylodactylidae – Pereopods 1 and 2 fingers not extremely long. Maxilliped 2 with last 2 articles in sequence (Fig. 6.21b)�������������������������6 6. Pereopod 1 with both fingers movable (Fig. 6.52b)��������������������������������������������������������Psalidopodoidea … Psalidopodidae – Pereopod 1 with only dactylus movable���������������������������������������������������������������������������������������������������������������������������������������7 7. Pereopods with epipods, ending in a naked appendix extending well into branchial cavity (Fig. 6.29i). Pereopods 1 and 2 similar�������������������������������������������������������������������������������������������������������������������������Oplophoroidea … 8 – Pereopodal epipods, if present, not ending in a naked appendix. Pereopods 1 and 2 similar or dissimilar���������������������9 8. Mandibles similar, molar weakly separate from incisor, consisting of channel flanked by thin walls opposing similar structure on other of pair (Fig. 6.30e), or reduced to small process (Fig. 6.30d)���������������������������� Oplophoridae – Mandibles dissimilar, molar distinct from incisor, with distal surface triangular on right member, compressed, sublinear on left (Fig. 6.29h)����������������������������������������������������������������������������������������������� Acanthephyridae 9. Pereopods 1 and 2 with similar chelae, fingers ending in dense brush of setae (Fig. 6.14b). Estuarine or freshwater��������������������������������������������������������������������������������������������������������������������������� Atyoidea … Atyidae – Pereopods 1 and 2 with fingers not ending in dense brush of setae. Marine or estuarine, rarely freshwater�����������������10 10. Pereopods 1 stronger and heavier, though often shorter than pereopods 2�������������������������������������������������������������������������11 – Pereopods 1 usually more slender (rarely subequal) than pereopods 2�������������������������������������������������������������������������������� 20 11. Pereopods without strap-like epipods. Mandibular molar conical, laminar or vestigial (Fig. 6.17c) ���������������������������������������������������������������������������������������������������������������������������������������������������������������������������Bresilioidea … 12 – Pereopods 1–3 at least with strap-like epipods (Fig. 6.48w). Mandibular molar blunt, with grinding surface (Fig. 6.26c, 6.27h)���������������������������������������������������������������������������������������������������������������Nematocarcinoidea … 16 12. Pereopod 1 chela dactylus semicircular, disc-like with sharp edge (Fig. 6.18f–h)����������������������������������������������Disciadidae – Pereopod 1 chela dactylus linear��������������������������������������������������������������������������������������������������������������������������������������������������13 13. Pereopods 1, 2 chelate, pereopods 3–5 pseudochelate, dactylus opposing long strong seta (Fig. 6.19b) ��������Pseudochelidae – Pereopods not as above������������������������������������������������������������������������������������������������������������������������������������������������������������������14 14. Antennal spine present. Eyes with complete cornea (Fig. 6.17a, e)��������������������������������������������������������������������������Bresiliidae – Antennal spine absent. Eyes with weakly or unpigmented cornea (Fig. 6.16)����������������������������������������������������������������������15 15. Pereopod 1 palm attached to the distal end of the carpus (Fig. 6.16l). Telson rounded, with spiniform setae and/or plumose setae (Fig. 6.16p–s)��������������������������������������������������������������������������������������������������������������Alvinocarididae – Pereopod 1 carpus articulating with chela at middle of propodus below base of dactylus, propodus projecting (Fig. 6.15a). Telson with 4 or 5 pairs of dorsal spiniform setae, posterior margin with 1 pair of long spiniform setae, 2 or 3 pairs of fine setae (Fig. 6.15b)�������������������������������������������������������������������� Agostocarididae 16. Rostrum grossly dentate, hinged and movable (Fig. 6.28a, c)�������������������������������������������������������������������� Rhynchocinetidae – Rostrum finely or grossly dentate, fixed�������������������������������������������������������������������������������������������������������������������������������������17 17. Rostrum grossly dentate. Carapace with 3 longitudinal lateral ridges (Fig. 6.25). Pereopods 1, 2 fingers with lateral and distal spines forming basket-like cage������������������������������������������������������������������������������������ Eugonatonotidae

6 – Caridea – shrimps

– 18. – 19. – 20. – 21. – 22. – 23. – 24. – 25. – 26. – 27. – 28. – 29. – 30. – 31. – 32.

63

Rostrum finely dentate. Carapace laterally smooth. Pereopods 1, 2 fingers without distal spines�����������������������������������18 Freshwater. Maxilliped 2 last article longer than broad, longer than penultimate article, attached by transverse suture����������������������������������������������������������������������������������������������������������������������������������������������� Xiphocarididae Marine, oceanic. Maxilliped 2 last article a narrow strip on submesial-distal margin of broader penultimate article (Figs 6.26b, 6.27i)�����������������������������������������������������������������������������������������������������������������������������������������������������������.19 Pereopods 3–5 twice length of pereopod 1, carpus several times length of propodus, articulation between ischium and merus thickened (Fig. 6.27n)�����������������������������������������������������������������������������������������������Nematocarcinidae Pereopods 3–5 1.5 times length of pereopod 1, carpus shorter than propodus, articulation between ischium and merus not thickened (Fig. 6.26a)������������������������������������������������������������������������������������������������������������������������Lipkiidae Pereopods 1, 2 both slender. Pereopod 2 chela with palm short, quarter length of fingers (Fig. 6.54). Anchialine caves�������������������������������������������������������������������������������������������������������������������������������������������Anchialocarididae Pereopod 2 more robust than pereopod 1. Pereopod 2 chela with palm longer than fingers. Usually marine���������������21 Pereopods 1–4 with arthrobranchs. Antennule with dorsal flagellum simple (Fig. 6.21)������������Campylonotoidea … 22 Pereopods without arthrobranchs. Antennule with dorsal flagellum of 2 branches (Fig. 6.40h, j) �������������������������������������������������������������������������������������������������������������������������������������������������������������������� Palaemonoidea … 23 Pereopods 2 subequal. Maxilliped 2 last article a narrow strip on mesial margin of penultimate (Fig. 6.21b). Epipods present�������������������������������������������������������������������������������������������������������������������������� Campylonotidae Pereopods 2 very strongly unequal. Maxilliped 2 last article forming oblique suture with penultimate article (Fig. 6.20e). Epipods absent����������������������������������������������������������������������������������������������������� Bathypalaemonellidae Freshwater only. Maxilliped 2 dactylus ovate, propodus with convex mesial margin (appearing bilobed). Maxilliped 1 with palp ovate������������������������������������������������������������������������������������������������������������������������� Desmocarididae Marine or freshwater. Maxilliped 2 dactylus not bilobed. Maxilliped 1 palp not ovate���������������������������������������������������� 24 Marine or freshwater. Maxilliped 1 exopod caridean lobe low, rounded (Fig. 6.34g–i)��������������������������������Palaemonidae Freshwater only. Maxilliped 1 exopod caridean lobe acutely produced, elongated������������������������������������������������������������ 25 Carapace postantennal longitudinal suture complete. Third antennular flagellum partially fused with dorsal antennular flagellum�������������������������������������������������������������������������������������������������������������������������� Typhlocarididae Carapace postantennal longitudinal suture absent. All antennular flagella free����������������������������������������Euryrhynchidae Carapace merging anteriorly into an inflated, massive rostrum (Fig. 6.50). Pereopod 2 with fixed finger curved around short dactylus������������������������������������������������������������������������������� Physetocaridoidea … Physetocarididae Rostrum, if present, not as above. Pereopod 2 with conventional chela������������������������������������������������������������������������������� 27 Right pereopod 1 chelate, left usually simple (Fig. 6.51l–o). Maxilliped 1 with exopod abutting endite, displacing palp (endopod) (Fig. 6.51k)�������������������������������������������������������������������������������������Processoidea … Processidae Pereopods 1 both simple or chelate. Maxilliped 1 with exopod far removed from endite (Fig. 6.44g)��������������������������� 28 Pereopod 1 with chelae microscopic or simple (Fig. 6.48k–m)������������������������������������������������������������������ Pandaloidea … 29 Pereopod 1 distinctly chelate (Figs 6.3, 6.8, 6.10, 6.11, 6.13)�������������������������������������������������������������������������Alpheoidea … 30 Rostrum usually well developed (Fig. 6.47). Pereopod 2 carpus variable, never of 3 segments. Telson with dorsolateral spiniform setae on dorsolateral ridges (Fig. 6.48s, t)���������������������������������������������������������������������Pandalidae Rostrum absent (Fig. 6.46b–d), or ventral margin with no or few teeth (Fig. 6.46a). Pereopod 2 carpus of 3 segments (Fig. 6.46j). Telson with dorsolateral spiniform setae adjacent to lateral margins (Fig. 6.46e)�������������������������������������������������������������������������������������������������������������������������������������������������������� Chlorotocellidae Eyestalks unusually elongate, as long as antennular peduncle. Pereopods 1 and 2 equally strong (Fig. 6.12)�������Ogyrididae Eyestalks of normal length. Pereopod 1 stronger than 2���������������������������������������������������������������������������������������������������������31 Eyes usually covered partly or entirely by carapace (Figs 6.1, 6.2). Pereopods 1 subequal or unequal, larger one usually swollen (Fig. 6.3)������������������������������������������������������������������������������������������������������������������������������������Alpheidae Eyes not covered. Pereopods 1 usually equal, not swollen�������������������������������������������������������������������������������������������������������32 Rostrum little longer than eyestalk, with 2 or 3 dorsal teeth, unarmed ventrally (Fig. 6.11a). Uropod exopod article 1 with anterior margin produced alongside article 2 (Fig. 6.11b). Semiterrestrial on mangrove prop roots������������������������������������������������������������������������������������������������������������������������������������������������������������������ Merguiidae

64

Marine Decapod Crustacea



Rostrum at least as long as eyestalk, usually armed dorsally and ventrally (Fig. 6.6–6.10). Uropod exopod article 1 with anterior margin not produced alongside article 2. Fully aquatic��������������������������������������������������������������33 33. Maxilliped 1 exopodal flagellum short (Fig. 6.7g). Maxilliped 3 last article flat (Fig. 6.7i, j). Mandible without palp or incisor. Deep-water��������������������������������������������������������������������������������������������������������������� Bythocarididae – Maxilliped 1 exopodal flagellum long (Fig. 6.10i). Maxilliped 3 last article linear (Fig. 6.10j). Mandible with palp or incisor or both, sometimes with neither���������������������������������������������������������������������������������������������������������������.34 34. Antennular peduncle article 2 with sharp lateral tooth (or at least tubercle) (Fig. 6.13l); dorsal flagellum usually thick, stout, with proximal articles annulated, bearing many closely-spaced setae, appearing brush-like (Fig. 6.13k). Pereopod 2 carpus with 6 or 7 segments (Fig. 6.13p)����������������������������������������������������� Thoridae – Antennular peduncle without lateral tubercle; dorsal flagellum not brush-like. Pereopod 2 carpus usually with fewer or more than 6 or 7 segments������������������������������������������������������������������������������������������������������������������������������35 35. Inhabiting anchialine caves or coastal saltwater pools (Fig. 6.6)������������������������������������������������������������������������ Barbouriidae – Inhabiting marine environments������������������������������������������������������������������������������������������������������������������������������������������������ 36 36. Antennular lateral flagellum distally biramous, or with thicker proximal section ending in short spur or swelling (Fig. 6.10f–h). Pereopod 2 carpus of 12–29 segments������������������������������������������������������������������������� Lysmatidae – Antennular lateral flagellum uniramous (Fig. 6.8). Pereopod 2 carpus of 2–17, exceptionally 90 (usually 3) segments����������������������������������������������������������������������������������������������������������������������������������������������������������������� Hippolytidae References Bracken HD, De Grave S, Toon A, Felder DL, Crandall KA (2010) Phylogenetic position, systematic status, and divergence time of the Procarididea (Crustacea: Decapoda). Zoologica Scripta 39, 198–212. doi:10.1111/j.​ 1463-6409.2009.00410.x Bracken HD, De Grave S, Felder DL (2009a) Phylogeny of the Infraorder Caridea based on mitochondrial and nuclear genes (Crustacea: Decapoda). In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 281–305. CRC Press, Boca Raton. Bracken HD, Toon A, Felder DL, Martin JW, Finley M, et al. (2009b) The decapod tree of life: compiling the data and moving toward a consensus of decapod evolution. Arthropod Systematics & Phylogeny 67, 99–116. Burukovsky RN (2012) Deep sea shrimps of the family Nematocarcinidae (history of study, systematics, distribution, and biology) Prospekt Nauki, St Petersberg. Chace FA (1992) On the classification of the Caridea (Decapoda). Crustaceana 63, 70–80. doi:10.1163/156854092X00299 Christoffersen ML (1987) Phylogenetic relationships of hippolytid genera, with an assignment of new families for the Crangonoidea and Alpheoidea (Crustacea, Decapoda, Caridea). Cladistics 3, 348–362. doi:10.1111/j.1096-0031.1987.tb00898.x Christoffersen ML (1990) A superfamily classification of the Caridea (Crustacea: Pleocyemata) based on phylogenetic pattern. Journal of Zoological Systematics and Evolutionary Research 28, 94–106. doi:10.1111/j.1439-0469.1990.tb00369.x D’Udekem D’Acoz C, De Grave S (2018) A new genus and species of largebodied caridean shrimp from the Crozet Islands, Southern Ocean (Crustacea, Decapoda, Lipkiidae) with a checklist of Antarctic and sub-Antarctic shrimps. Zootaxa 4392, 201–240. doi:10.11646/zootaxa.4392.2.1 De Grave S, Fransen CHJM (2011) Carideorum Catalogus: the Recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoölogische Mededeelingen 85, 195–589. De Grave S, Moosa MK (2004) A new species of the enigmatic shrimp genus Pseudocheles (Decapoda: Bresiliidae) from Sulawesi (Indonesia), with the designation of a new family Pseudochelidae. Crustacean Research 33, 1–9. doi:10.18353/crustacea.33.0_1

De Grave S, Li CP, Tsang LM, Chu KH, Chan T-Y (2014) Unweaving hippolytoid systematics (Crustacea, Decapoda, Hippolytidae): resurrection of several families. Zoologica Scripta 43, 496–507. doi:10.1111/zsc.12067 Guerao G, Reuschel S, Anger K, Schubart CD (2014) On the presumed phylogenetic position of the Xiphocarididae (Decapoda, Caridea) based on the larval morphology of Xiphocaris elongata. In: Yeo D, Cumberlidge N, Klaus S (Eds) Advances in freshwater decapod systematics and biology. Crustaceana Monographs 19, 233–244. Holthuis LB (1955) The Recent genera of the caridean and stenopodidean shrimps (Class Crustacea, Order Decapoda, Supersection Natantia) with keys for their determination. Zoologische Verhandelingen 26, 1–153. Holthuis LB (1993) The Recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda) with an appendix on the Order Amphionidacea. revised edn. Nationaal Natuuristorisch Museum, Leiden. Komai T, Segonzac M (2003) Review of the hydrothermal vent shrimp genus Mirocaris, redescription of M. fortunata and reassessment of the taxonomic status of the family Alvinocarididae (Crustacea: Decapoda: Caridea). Cahiers de Biologie Marine 44, 199–215. Mejía-Ortíz L, Yañez G, López-Mejía M (2017) Anchialocarididae, a new family of anchialine decapods and a new species of the genus Agostocaris from Cozumel Island, Mexico. Crustaceana 90, 381–398. doi:10.1163/15685403-00003657 Wang Y, Ma KY, Tsang LM, Wakabayashi K, Chan T-Y, et al. (2021) Confirming the systematic position of two enigmatic shrimps, Amphionides and Procarididae (Crustacea: Decapoda). Zoologica Scripta 50, 812–823. doi:10.1111/zsc.12509 Wicksten MK (2010) Infraorder Caridea Dana, 1852. In Treatise on Zoology – Anatomy Taxonomy Biology. The Crustacea Volume 9 Part A. Eucarida: Euphausiacea, Amphionidacea, and Decapoda (partim). (Eds Schram FR, Charmantier-Duares M, Von Vaupel Klein JC, Forest J). pp. 165–206. Brill, Leiden. Wicksten M, De Grave S, France S, Kelley C (2017) Presumed filter-feeding in a deep-sea benthic shrimp (Decapoda, Caridea, Stylodactylidae), with records of the deepest occurrence of carideans. ZooKeys 646, 17–23. doi:10.3897/zookeys.646.10969 Williamson DI (1973) Amphionides reynaudii (H. Milne Edwards), representative of a proposed new order of eucaridean Malacostraca. Crustaceana 25, 35–50. doi:10.1163/156854073X00470

6 – Caridea – shrimps

Superfamily Alpheoidea Rafinesque, 1815 Alpheoids are a diverse assemblage of eight families, most in shallow marine waters. The superfamily is diagnosed with a multiarticulate carpus on pereopod 2 (at least in most genera) and two pairs of chelate pereopods. They differ from pandaloids, the only other superfamily with all its members having a multiarticulate carpus, in the last article of maxilliped 2 being a narrow strip along the mesial margin of the penultimate article, instead of having these articles in sequence. The monophyly of Alpheoidea has not been questioned. However, the arrangement of families within the so-called ‘hippolytoid’ group of families, all except Alpheidae and Ogyrididae, has been more fluid (Bracken et al. 2009; De Grave et al. 2014). Holthuis (1993) recognised five families in his keys, De Grave and Fransen (2011) catalogued species in four families, but molecular evidence has found these not to be well supported. This evidence led De Grave et al. (2014) to advocate six hippolytoid families plus the more easily recognised Alpheidae and Ogyrididae. This schema is followed here but morphological diagnosis of the families is difficult, as De Grave et al. (2014) acknowledged. Diagnosis. Maxilliped 1 exopod far removed from endite (Fig. 6.7g). Maxilliped 2 last article a narrow strip on terminal half of mesial margin of penultimate article (Fig. 6.7h). Pereopods 1, 2 similar, chelate, or dissimilar, each pair symmetrical, first more robust than second, or first pair asymmetrical, propodus swollen on one side. Pereopod 2 carpus multiarticulate. Exopods present on maxilliped 3, pereopods 1–5. References Bracken HD, De Grave S, Felder DL (2009) Phylogeny of the Infraorder Caridea based on mitochondrial and nuclear genes (Crustacea: Decapoda). In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 281–305. CRC Press, Boca Raton. De Grave S, Fransen CHJM (2011) Carideorum Catalogus: the Recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoölogische Mededeelingen 85, 195–589. De Grave S, Li CP, Tsang LM, Chu KH, Chan T-Y (2014) Unweaving hippolytoid systematics (Crustacea, Decapoda, Hippolytidae): resurrection of several families. Zoologica Scripta 43, 496–507. doi:10.1111/zsc.12067 Holthuis LB (1993) The Recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda) with an appendix on the Order Amphionidacea. revised edn. Nationaal Natuuristorisch Museum, Leiden.

Alpheidae Rafinesque, 1815 snapping shrimps, pistol shrimps Figures 6.1–6.5, Plate 6

65

The family Alpheidae includes the snapping shrimps renowned for the sharp clicking noise made with the larger chela, but not all alpheids have this capability (Anker et al. 2006a). Individuals are frequently seen under rocks, in burrows and in sponges near low water level and subtidally. Alpheids are probably the most abundant family of caridean shrimps subtidally in temperate environments and their clicking noise can frequently be heard by divers (Chitre et al. 2012). The click generates a shock wave that stuns prey (small fish and other crustaceans) before being captured (Qin et al. 2019). Alpheidae are characterised by the first pereopods usually having an enlarged claw (with a snapping mechanism in some genera), the posterior margin of the carapace having a well developed cardiac notch (Chace and Kensley 1992), and the eyestalks being dorsally at least partly, and often completely covered by the anterior margin of the carapace (Anker and Komai 2004; Chace 1988). A phylogenetic hypothesis of relationships among 36 genera has shed light on the evolution of this family (Anker et al. 2006a). Alpheidae are certainly monophyletic but the largest genera, Alpheus, with over 300 species, and other large genera are not. Several functionally significant features of the cheliped evolved independently several times, such as asymmetry, folding, inverted orientation, sexual dimorphism, adhesive plaques that enhance claw cocking, and tooth-cavity systems on opposing claw fingers, a preadaptation for snapping. The most recent key to genera (Holthuis 1993) included 31 genera  – there are now 50  – and today seems quite removed from any phylogenetic progression. Nevertheless, it relies largely on relatively easy dichotomous choices and forms the basis for this new key. The cornea is well exposed in some genera, completely covered from dorsal view in others  – intermediate conditions may prove problematic. Diagnosis. Rostrum small, obsolete, or absent. Eyestalks compact, completely or partially covered by anterior margin of carapace. Antennular lateral flagellum with accessory branch (usual). Mandible palp apically bifid (sometimes). Pereopods 1, 2 usually dissimilar, if so first pair usually asymmetrical, propodus swollen on one side. Implicit generic attributes. Orbital hoods present, eyestalks concealed from dorsal view, visible from anterior view. Mandible with palp. Chelipeds usually unequal, usually extended forwards; dactylus without molar-shaped tooth fitting into socket. Uropodal exopod suture with spiniform seta at anterior end. At least pereopods 1, 2 with epipods.

66

Marine Decapod Crustacea

Key to genera of Alpheidae 1.

Carapace and pleon strongly compressed dorsoventrally; carapace with anterolateral projections; dorsal integument setose (Fig. 6.1c)�������������������������������������������������������������������������������������������������������������������������������������Pterocaris – Carapace and pleon not strongly compressed dorsoventrally, typically shrimp-like, usually glabrous (Fig. 6.1a, b)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 2. Pleonites 3–5 pleura with 2–4 small spines along posterior margin���������������������������������������������������������������� Crosnierocaris – Pleonites 3–5 pleura unarmed along posterior margin��������������������������������������������������������������������������������������������������������������3 3. Pereopods without epipods�������������������������������������������������������������������������������������������������������������������������������������������������������������4 – At least pereopods 1, 2 with epipods (absent in 1 species of Rugathanas)�������������������������������������������������������������������������������9 4. Major cheliped dactylus without molar-shaped tooth. Mandible without palp���������������������������������������������������������������������5 – Major cheliped dactylus with molar-shaped tooth fitting into socket (Fig. 6.4d). Mandible with palp�����������������������������7 5. Pereopod 2 fingers less than quarter length of palm (Fig. 6.4h)������������������������������������������������������������������������������������ Batella – Pereopod 2 fingers more than half length of palm���������������������������������������������������������������������������������������������������������������������6 6. Mandible without molar; incisor widened distally, with asymmetrical dentition (Fig. 6.4k). Rostrum acute (Fig. 6.2x). Pleonite 6 pleuron with movable plate on posteroventral angle (Fig. 6.4p, q). Uropodal exopod suture toothed along anterior half (Fig. 6.5i)�������������������������������������������������������������������������������������� Prionalpheus – Mandible with molar; incisor not widened distally, simply dentate. Rostrum rounded (Fig. 6.1s). Pleonite 6 pleuron without movable plate on posteroventral angle. Uropodal exopod suture with 1 spiniform seta at anterior margin (Fig. 6.5a, b)������������������������������������������������������������������������������������������������������������������������������� Bannereus 7. Maxilliped 3 ischium-merus operculiform (Fig. 6.4o)�����������������������������������������������������������������������������������������Pomagnathus – Maxilliped 3 ischium-merus pediform (Fig. 6.4m)��������������������������������������������������������������������������������������������������������������������8 8. Rostrum, orbital spines prominent (Fig. 6.1a). Uropodal exopod suture sinuous (Fig. 6.5m)���������������������������Synalpheus – Rostrum, orbital spines absent, front sinuous (Fig. 6.2B). Uropod exopod suture with deep incision and triangular tooth near posterior margin (Fig. 6.5k)�������������������������������������������������������������������������������������������� Richalpheus 9. Pleonite 6 pleuron with movable plate on posteroventral angle (absent in 1 species of Nennalpheus) (Fig. 6.4p, q)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������10 – Pleonite 6 pleuron without movable plate on posteroventral angle (Fig. 6.1a)���������������������������������������������������������������������32 10. Rostrum absent, mid-anterior margin of carapace sinuous, concave or evenly convex (Figs 6.1x, 6.2u, i, o, E)��������������������������������������������������������������������������������������������������������������������������������������������������������������11 – Rostrum present, with acute, triangular or rounded tip (Figs 6.1f, p, q, 6.2y)���������������������������������������������������������������������17 11. Uropodal exopod with terminal segmented appendage, suture with ~9 lobe-like crenellations (Fig. 6.5l)�������������������������������������������������������������������������������������������������������������������������������������������������������������Stenalpheops – Uropodal exopod without terminal segmented appendage, suture not crenellated����������������������������������������������������������� 12 12. Front concave at midpoint������������������������������������������������������������������������������������������������������������������������������������������������������������13 – Front convex at midpoint��������������������������������������������������������������������������������������������������������������������������������������������������������������14 13. Telson apex produced as triangular projection (Fig. 6.4t). Cheliped merus with 2 distal spines (Fig. 6.3t). Front with sharp extraorbital teeth���������������������������������������������������������������������������������������������������������������������� Parabetaeus – Telson apex not produced as triangular projection. Cheliped merus without distal spines. Chelipeds subequal, not flexed, chela inverted, dactylus below fixed finger (Fig. 6.3h). Front sinuous������������������������������ Betaeus 14. Antennular peduncle article 2 longer than both article 1 and 3 (Fig. 6.2i). Major cheliped fingers slender, not gaping, evenly toothed (Fig. 6.3l, m)��������������������������������������������������������������������������������������������������������� Fenneralpheus – Antennular peduncle article 1 as long or longer than article 2 (Fig. 6.2o). Major cheliped fingers often somewhat twisted, gaping, unevenly toothed, or reflexed (Fig. 6.3n)������������������������������������������������������������������������������15 15. Uropodal exopod suture with 2 strong spiniform setae near anterior margin (Fig. 6.5g)������������������������������� Pachelpheus – Uropodal exopod suture without or with 1 spiniform seta near anterior margin (Fig. 6.5c, d)����������������������������������������16 16. Antennule article 1 with strong mesioventral tooth. Uropodal exopod suture with deep incision and tooth at midpoint (Fig. 6.5c). Minor chela fingers longer than palm, slender (Fig. 6.4f)����������������������������������������Leptalpheus

6 – Caridea – shrimps

– 17. – 18. – 19. – 20. – 21. – 22. – 23. – 24. – 25. – 26. – 27. – 28. – 29. – 30. – 31.

67

Antennule article 1 without mesioventral tooth. Uropodal exopod suture incomplete (Fig. 6.5d). Minor chela fingers shorter than palm (Fig. 6.4g)���������������������������������������������������������������������������������������������������������Leptathanas Eyes entirely (rarely nearly so) concealed from dorsal view, visible from anterior view (Figs 6.1f, g, p, 6.2f, h, u, y)�������������������������������������������������������������������������������������������������������������������������������������������������������18 Eyes entirely (or substantially) visible in dorsal view (Figs 6.1k, l, n, 6.2a, n, C)����������������������������������������������������������������� 22 Carapace with small marginal tooth dorsal to pterygostomial angle (Fig. 6.2z). Major cheliped cutting edges serrated on proximal two-thirds, followed by broad hiatus and large subdistal tooth (Fig. 6.3u)����������������������������������������������������������������������������������������������������������������������������������������������������������� Pseudalpheopsis Carapace without marginal tooth dorsal to pterygostomial angle. Major cheliped otherwise�����������������������������������������19 Rostrum distally rounded, with broad vertical ventral lamella (Fig. 6.1p, q)������������������������������������������������������� Athanopsis Rostrum distally acute, with narrow ventral ridge at most���������������������������������������������������������������������������������������������������� 20 Rostrum alone triangular or narrowly acute; extraorbital teeth sharp or sometimes absent (Fig. 6.1f, g). Carapace without anterolateral suture���������������������������������������������������������������������������������������������Alpheopsis Rostrum and ocular hoods together broadly triangular; extraorbital teeth absent or invisible from dorsal view. Carapace with anterolateral suture (Fig. 6.2g)�����������������������������������������������������������������������������������������������������������21 Rostrum with 2 anteriorly-directed long setae (Fig. 6.2f, g). Chelipeds equal, symmetrical, carried extended, chelae robust, carpus with mesial rows of setae (Figs 6.3j, 6.4c)���������������������������������������������� Coutieralpheus Rostrum without anteriorly-directed long setae (Fig. 6.2h). Chelipeds unequal, asymmetrical, carried flexed, chelae cylindrical, carpus without mesial rows of setae (Fig. 6.3k)��������������������������������������������������������� Deioneus Uropodal exopod suture with row of strong movable spines (Fig. 6.5j)����������������������������������������������������������� Pseudathanas Uropodal exopod suture without row of strong movable spines������������������������������������������������������������������������������������������� 23 Eyestalks each with conspicuous acute spine (Fig. 6.1d, e)�������������������������������������������������������������������������������������Acanthanas Eyestalks without conspicuous acute spine, sometimes with tubercle��������������������������������������������������������������������������������� 24 Cheliped carried extended; dactylus in lower position (Fig. 6.3d)�����������������������������������������������������������������������������Aretopsis Chelipeds carried flexed or extended; dactylus in upper or lateral position����������������������������������������������������������������������� 25 Male chelipeds with oval chelae, palm with rugose margins (Fig. 6.3v). Supraorbital teeth present (Fig. 6.2C). Mandible with expanded incisor (Fig. 6.4l)�������������������������������������������������������������������������������������Rugathanas Male chelipeds with elongate or oval chelae, palm with smooth margins. Supraorbital teeth present or absent. Mandible without expanded incisor������������������������������������������������������������������������������������������������������������������������ 26 Extraorbital teeth present. Rostrum elongate, tapering (Figs 6.1k, n, o, 6.2a, w)���������������������������������������������������������������� 27 Extraorbital teeth absent. Rostrum short, triangular or minute (Fig. 6.2m, n, s)��������������������������������������������������������������� 30 Uropodal exopod suture with numerous crenellations along anterior two-thirds (Fig. 6.5h)�������������������� Potamalpheops Uropodal exopod suture without crenellations������������������������������������������������������������������������������������������������������������������������ 28 Chelipeds carpus and merus with mesial rows of short setae (Fig. 6.4b). Pereopods 2–4 coxae with blunt process alongside basis (Fig. 6.4a)���������������������������������������������������������������������������������������������������������������������������� Bruceopsis Chelipeds carpus and merus without mesial rows of short setae. Pereopods 2–4 coxae without blunt process alongside basis������������������������������������������������������������������������������������������������������������������������������������������������������������ 29 Pereopod 2 carpus with 4 segments in adults. Pereopods 1−2 with epipod. Cheliped ischium usually with strong mesial process (Fig. 6.3c). Supraorbital teeth absent (Fig. 6.1k)���������������������������������������������������������������������� Arete Pereopod 2 carpus with 5 segments in adults. Pereopods 1−3 with epipod. Cheliped ischium without mesial process. Supraorbital teeth absent or present (Fig. 6.1n, o)�������������������������������������������������������������������������������������Athanas Chelipeds equal, symmetrical, carpus with mesial rows of short setae (Fig. 6.4c), dactylus tapering, above fixed finger. Carapace with horizontal anterolateral suture (Fig. 6.2m, n)��������������������������������������������������� Jengalpheops Chelipeds unequal, symmetrical or asymmetrical, carpus without mesial rows of short setae, dactylus wide, laminar, inverted, below fixed finger (Fig. 6.3q, r). Carapace without horizontal anterolateral suture������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������31 Major cheliped chela with finely serrate cutting edges, dactylus broad, laminar (Fig. 6.3r)������������������������������Notalpheus

68

– 32. – 33. – 34. – 35. – 36. – 37. – 38. – 39. – 40. – 41. – 42. – 43. – 44. – 45. – 46. –

Marine Decapod Crustacea

Major cheliped chela with rounded fitting teeth along proximal half of cutting edges, dactylus with distal hook (Fig. 6.3q)�������������������������������������������������������������������������������������������������������������������������������������������������������Nennalpheus Eyestalks entirely or partially visible in dorsal view (Figs 6.1r, v, 6.2c, e, H, M)������������������������������������������������������������������33 Eyestalks completely covered in dorsal view, sometimes exposed in anterior or lateral view (Figs 6.1h–j, w, 6.2k, p–r, t, A, D, J)������������������������������������������������������������������������������������������������������������������������������������������������������������������� 39 Chelipeds equal or subequal, symmetrical (Fig. 6.3g, i, x)����������������������������������������������������������������������������������������������������� 34 Chelipeds unequal, asymmetrical (Figs 6.1a, 6.3f, s, w)�����������������������������������������������������������������������������������������������������������37 Rostrum triangular, longer than eyestalks. Chelipeds of minor form, chelae minute��������������������������������������������������������35 Rostrum minute or broadly rounded. Chelipeds with chelae about long as carapace������������������������������������������������������� 36 Rostrum elongate, tapering, without extraorbital teeth, without teeth in midline (Fig. 6.2l, m, L, M). Telson with rectangular projection beyond subdistal spiniform setae (Fig. 6.4u)����������������������������������������������������� Yagerocaris Rostrum triangular, with extraorbital teeth, elevated in midline, triangular in cross-section, with 1 or 2 postrostral teeth (Fig. 6.2c, d). Telson with deeply cleft apex (Fig. 6.4s)���������������������������������������������������������� Caligoneus Carapace posterodorsal margin with pair of submedian high bluntly-rounded crests in adult. Rostrum minute (Fig. 6.2F, G). Chelipeds with dactylus laminar, carried above fixed finger (Fig. 6.3x)�������������������Thuylamea Carapace smooth dorsally. Rostrum broadly rounded (Fig. 6.1v). Chelipeds with dactylus narrow, as long as palm, carried below fixed finger (Fig. 6.3g)������������������������������������������������������������������������������������������������Bermudacaris Carapace with mediodorsal spine, surface behind orbit usually with posthepatic spine. Rostrum prominent, acute, with subrostral tooth (Fig. 6.2H, I)������������������������������������������������������������������������������������������������������� Triacanthoneus Carapace smooth dorsally, rarely with mediodorsal spine. Rostrum absent, rounded or triangular����������������������������� 38 Cornea reduced, weakly pigmented, eyestalks adjacent, without (rarely with) mesiodistal tubercle. Rostrum usually short, acute, sometimes rounded (Fig. 6.1r)���������������������������������������������������������������������������������������������� Automate Cornea spherical, pigmented, eyestalks divergent, with mesiodistal tubercle. Rostrum broadly triangular, as long as eyestalk (Fig. 6.2e)������������������������������������������������������������������������������������������������������������������������������ Coronalpheus Body strongly laterally compressed. Carapace and pleon with high median dorsal carina (Fig. 6.2A)�����������������Racilius Body not laterally compressed. Carapace and pleon without carina������������������������������������������������������������������������������������ 40 Uropodal endopod more than 2.5 times as long as exopod, ending in ribbon-like extension (Fig. 6.5f). Carapace with 1 median and 2 submedian granular ridges in anterior half (Fig. 6.2r). Cheliped dactylus flat, discoid (Fig. 6.4e)����������������������������������������������������������������������������������������������������������������������������������������������� Mohocaris Uropodal endopod about as long as exopod. Carapace dorsally smooth or with spines. Cheliped dactylus typical, slender, distally curved�����������������������������������������������������������������������������������������������������������������������������������������������41 Major cheliped dactylus with molar tooth fitting into socket in fixed finger or with swelling on dactylus fitting into shallow depression on fixed finger (Fig. 6.4d)������������������������������������������������������������������������������������������������ 42 Major cheliped dactylus without molar tooth�������������������������������������������������������������������������������������������������������������������������� 44 Maxilliped 3 ischium-merus broad, flat, suboperculiform (Fig. 6.4n)����������������������������������������������������������������� Metalpheus Maxilliped 3 ischium-merus pediform (Fig. 6.4m)����������������������������������������������������������������������������������������������������������������� 43 Eyestalks concealed from dorsal, not anterior view (Fig. 6.1j). Chelipeds folded under body��������������������� Amphibetaeus Eyestalks concealed from dorsal and anterior views (Fig. 6.1a, h, i). Chelipeds held forwards, not folded under body�����������������������������������������������������������������������������������������������������������������������������������������������������������������������Alpheus Rostrum absent or front sinuous������������������������������������������������������������������������������������������������������������������������������������������������ 45 Rostrum prominent���������������������������������������������������������������������������������������������������������������������������������������������������������������������� 47 Ocular hoods hemispherical, separated by notch (Fig. 6.1w). Uropodal endopod with 2 distal strong hooked spiniform setae (Fig. 6.5a)�������������������������������������������������������������������������������������������������������������������������� Betaeopsis Carapace frontal margin sinuous (Fig. 6.2p, t). Uropodal endopod without distal spiniform setae������������������������������� 46 Chelipeds with chelae robust, inflated (Fig. 6.3p)��������������������������������������������������������������������������������������������������� Leslibetaeus Chelipeds with chelae elongate (Fig. 6.3s)������������������������������������������������������������������������������������������������������������Orygmalpheus

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69

Fig. 6.1.  Alpheidae. a, Alpheus richardsoni Yaldwyn, 1971; b, Synalpheus stimpsonii (De Man, 1888); c, Pterocaris typica Heller, 1862. Front of carapace: d, e, Acanthanas; f, Alpheopsis keiji Anker, 2017; g, A. trigona (Rathbun, 1901); h, Alpheus socialis Heller, 1862; i, A. stephensoni Banner & Smalley, 1969; j, Amphibetaeus; k, Arete; l, m, Aretopsis; n, Athanas djiboutensis Coutière, 1897; o, A. ivoiriensis Anker & Ahyong, 2007; p, q, Athanopsis; r, Automate; s, Bannereus; t, Batella parvimanus (Bate, 1888); u, B. praecipua De Grave, 2004; v, Bermudacaris; w, Betaeopsis; x, Betaeus.

47. Carapace with prominent pterygostomial tooth. Ocular hood unarmed (Fig. 6.2k, l)���������������������������������Harperalpheus – Carapace pterygostomial angle rounded. Ocular hood with anterior tooth or spine (Fig. 6.2q, D, J, K)����������������������� 48 48. Mandible with palp, incisor with conspicuous dark-brown spot visible in external view of mouthparts (Fig. 6.4j). Uropodal endopod with tooth near end of anterior margin (Fig. 6.5e)�������������������������������������� Metabetaeus – Mandible incisor unpigmented. Uropodal endopod with smooth anterior margin���������������������������������������������������������� 49 49. Major chelipeds equal, symmetrical, carried extended forwards, inverted, dactylus below fixed finger (Fig. 6.3y). Mandible without palp��������������������������������������������������������������������������������������������������������������������������� Vexillipar – Major cheliped unequal, asymmetrical, carried flexed, not inverted, dactylus above fixed finger (Fig. 6.3w). Mandible with palp����������������������������������������������������������������������������������������������������������������������������������������������������Salmoneus

70

Marine Decapod Crustacea

Fig. 6.2.  Alpheidae. Front of carapace: a, b, Bruceopsis; c, d, Caligoneus; e, Coronalpheus; f, g, Coutieralpheus; h, Deioneus; i, j, Fenneralpheus (with antennules, antennae); k, l, Harperalpheus; m, n, Jengalpheops; o, Leptalpheus (with antennules, antennae); p, Leslibetaeus; q, Metabetaeus; r, Mohocaris; s, Nennalpheus; t, Orygmalpheus; u, Parabetaeus; v, Pomagnathus; w, Potamalpheops; x, Prionalpheus; y, z, Pseudalpheopsis; A, Racilius; B, Richalpheus; C, Rugathanas; D, Salmoneus; E, Stenalpheops; F, G, Thuylamea; H, I, Triacanthoneus; J, K, Vexillipar; L, M, Yagerocaris.

Diagnosis. Rostrum elongate, tapering; orbital hoods present, eyestalks partly visible in dorsal view; extraorbital teeth acute; cornea spherical, pigmented, with spiny projection. Pterygostomial angle angular. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds equal, symmetrical or subsymmetrical; chela robust, inflated (male). Maximum cl. 2.3 mm. Subtidal (7–24 m). Central and Eastern Indo-Pacific. 1 species (Anker et al. 2006b).

movable plate on posteroventral angle, or without movable plate on posteroventral angle (rarely). Chelipeds asymmetrical; chela robust, inflated or cylindrical. Maximum cl. estimated 10 mm (Pl. 6a, b). Intertidal–slope (0–786 m). Cosmopolitan except polar seas and Temperate North Atlantic. 26 species. Alpheopsis is a diverse genus that has been divided into groups of which the ten species of the A. trispinosa (Stimpson, 1860)-group has received the most attention (Anker 2017a; Anker et al. 2005). Some species have variously carinate carapaces (Anker 2017b). The most recent key to all species dealt with 20 species (Wang and Sha 2016b).

Alpheopsis Coutière, 1897

Alpheus Fabricius, 1798

Diagnosis. Rostrum triangular, or acute; extraorbital teeth absent, or long, acute. Pterygostomial angle angular. Pleonite 6 pleuron with

Diagnosis. Rostrum triangular; orbital hoods inflated, completely covered by carapace dorsally and partly anteriorly;

Acanthanas Anker, Poddoubtchenko & Jeng, 2006

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Fig. 6.3.  Alpheidae. Major cheliped: a, b, Amphibetaeus; c, Arete; d, Aretopsis; e, Athanopsis; f, Automate; g, Bermudacaris; h, Betaeus; i, Caligoneus; j, Coutieralpheus; k, Deioneus; l, m, Fenneralpheus; n, Leptalpheus; o, Leptathanas; p, Leslibetaeus; q, Nennalpheus; r, Notalpheus; s, Orygmalpheus; t, Parabetaeus; u, Pseudalpheopsis; v, Rugathanas; w, Salmoneus; x, Thuylamea; y, Vexillipar.

orbital hoods with or without anterior supraocular spine. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds asymmetrical; chela robust, inflated; dactylus with molar-shaped tooth fitting into socket. Maximum cl. 18 mm (Pl. 6c, d, m, n). Intertidal–slope (0–640 m). Cosmopolitan except polar seas. 327 species. Species of Alpheus are the typical snapping shrimps, or pistol shrimps, with a molar-like projection on the cheliped dactylus fitting into a cavity on the fixed finger. Considerable research has been done on the physics of the use of this mechanism in stunning prey (Qin et al. 2019; Versluis et al. 2000). The genus has been divided into seven species groups (see Banner and Banner 1982), some of which are now thought to be polyphyletic (Williams et al. 2001). The genus is certainly paraphyletic (Anker et al. 2006a). Keys to species are confined to regional faunas, such as 85 species from Australia (Banner and Banner 1982), 28 species from southern Australia (Poore 2004), and 83 species from the Philippines (Chace 1988) but many more species have been described since.

Amphibetaeus Coutière, 1897 Diagnosis. Rostrum absent, front sinuous. Carapace with horizontal anterolateral suture. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds asymmetrical, flexed; dactylus swollen at midpoint, fitting shallow depression on fixed finger. Maximum cl. 11 mm. Intertidal, subtidal. Western Indo-Pacific. 1 species (Anker and Jeng 2006; Marin et al. 2014).

Arete Stimpson, 1860 Diagnosis. Rostrum elongate, tapering; orbital hoods absent, eyestalks mostly visible in dorsal view; extraorbital teeth long, acute. Pterygostomial angle rounded, or angular. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds subequal or equal, subsymmetrical, flexed; major cheliped ischium usually with strong mesial process; chela cylindrical. Pereopod 2 carpus with 4 segments in adults. Pereopods 1−2 with epipod.

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Fig. 6.4.  Alpheidae. Major cheliped coxa: a, Bruceopsis. Major cheliped, merus, carpus, setal brush: c, Bruceopsis; d, Coutieralpheus. Major cheliped, fingers: d, Metalpheus; e, Mohocaris. Minor cheliped: f, Leptalpheus; g, Leptathanas. Pereopod 2, propodus, dactylus: h, Batella. Mandible: i, Bruceopsis; j, Metabetaeus; k, Prionalpheus (pair); l, Rugathanas. Maxilliped 3: m, Richalpheus; n, Metalpheus; o, Pomagnathus. Pleonite 6, with articulated plate: p, Bruceopsis; q, Coutieralpheus. Telson: r, Alpheopsis; s, Caligoneus; t, Parabetaeus; u, Yagerocaris.

Fig. 6.5.  Alpheidae. Uropod: a, Betaeopsis (with telson); b, Jengalpheops; c, Leptalpheus; d, Leptathanas; e, Metabetaeus; f, Mohocaris; g, Pachelpheus; h, Potamalpheops; i, Prionalpheus; j, Pseudathanas; k, Richalpheus; l, Stenalpheops; m, Synalpheus.

Intertidal–shelf (0–50 m, obligate symbionts of sea urchins). Indo-West Pacific, Temperate Australasia. 4 species (Anker and Jeng 2007).

Aretopsis De Man, 1910 Diagnosis. Rostrum tip rounded, with broad vertical ventral lamella; orbital hoods absent, eyestalks mostly visible in dorsal

view. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds asymmetrical; chela robust, inflated, chela inverted, dactylus below fixed finger; fingers widely gaping, with uneven teeth. Maximum cl. 5 mm. Intertidal–slope (0–550 m, inside shells occupied by diogenid hermit crabs, Dardanus spp.). Temperate Northern Pacific, Western and Central Indo-Pacific 3 species (Ashrafi et al. 2021; Banner and Banner 1973; Marin 2010: illustrations).

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Athanas Leach, 1814

Batella Holthuis, 1955

Diagnosis. Rostrum elongate, tapering; orbital hoods absent, eyestalks mostly visible in dorsal view; extraorbital teeth acute. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds subequal, asymmetrical or symmetrical, flexed or extended forwards; chela robust, inflated or cylindrical. Pereopod 2 carpus with 5 segments in adults. Pereopods 1−3 with epipod. Maximum cl. 5 mm. Intertidal–slope (0–450 m; coral reef crevices, or living commensally in burrows or associated with echinoderms; 0–450 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Temperate Southern Africa, Temperate Australasia. 41 species. Anker and Jeng (2007) rediagnosed the genus and discussed similarities to others with a triangular articulated plate on pleonite 6 and a well developed acute rostrum. Athanas has been divided into two species-groups based on the way the cheliped are held, and within that into ecological groups; all are associated with other invertebrates (Anker and Ahyong 2007a).

Diagnosis. Rostrum triangular; orbital hoods with or without anterior supraocular spine. Pterygostomial angle produced. Pleonite 6 pleuron without movable plate on posteroventral angle. Mandible without palp. Chela robust, inflated. Pereopod 2 chela fingers less than quarter length of palm, shrouded in setae. Pereopods without epipods. Maximum cl. 6 mm. Subtidal–slope (15–450 m; associated with hexactinellid and other sponges). Central Indo-Pacific. 3 species (De Grave 2004).

Athanopsis Coutière, 1897 Diagnosis. Rostrum tip rounded, with broad vertical ventral lamella; orbital hoods with blunt supraocular tooth; extraorbital teeth absent, or triangular. Pterygostomial angle angular, or rounded, protruding. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds unequal, asymmetrical, flexed; chela swollen at midpoint, narrow at base of angled fingers, folding against laterally expanded merus. Maximum cl. 8 mm (Pl. 6e). Intertidal, subtidal (0–12 m, possibly associated with burrows of echiurans). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 8 species (Anker and Ahyong 2007b: key to 5 species; Marin et al. 2014).

Automate De Man, 1888 Diagnosis. Rostrum triangular, or absent, or minute; orbital hoods absent, eyestalks mostly visible in dorsal view (contiguous); cornea weakly pigmented, eyestalks adjacent. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds asymmetrical; chela wide, flat. Maximum cl. 9 mm. Intertidal–slope (0–250 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific. 13 species (Anker and Komai 2004: informal species-groups).

Bannereus Bruce, 1988 Diagnosis. Rostrum rounded. Pleonite 6 pleuron without movable plate on posteroventral angle. Mandible without palp. Chelipeds asymmetrical; chela robust, inflated. Pereopods without epipods. Maximum cl. 5.7 mm. Slope (345–360 m; associated with hexactinellid sponges). Central Indo-Pacific. 1 species (Bruce 1988).

Bermudacaris Anker & Iliffe, 2000 Diagnosis. Rostrum rounded; orbital hoods absent, eyestalks mostly visible in dorsal view; cornea weakly pigmented, eyestalks adjacent. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds equal, symmetrical; with mesial carpal rows of short setae; chela wide, flat, chela inverted, dactylus below fixed finger; fingers as long as palm. Maximum cl. 4.5 mm. Intertidal, marine anchialine caves (0–38 m). Temperate Northern and Tropical Atlantic, Western and Central Indo-Pacific. 3 species (Anker et al. 2006c: key to species).

Betaeopsis Yaldwyn, 1971 Diagnosis. Rostrum absent, ocular hoods separated by deep notch. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds equal, subsymmetrical; chela inverted, dactylus below fixed finger. Uropodal endopod with 2 distal strong hooked spiniform setae. Maximum cl. 7.7 mm. Low supratidal–subtidal (0–18 m). Western and Central IndoPacific, Temperate Australasia. 3 species (Anker and Jeng 2002: key to species).

Betaeus Dana, 1852 Diagnosis. Rostrum absent, mid-front of carapace concave; orbital hoods broadly rounded, forming part of sinuous anterior margin of carapace. Carapace with horizontal anterolateral suture. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds unequal, asymmetrical; chela robust, inflated, chela inverted, dactylus below fixed finger; fingers widely gaping, with uneven teeth (variable). Maximum cl. 16 mm (Pl. 6f). Intertidal–shelf (0–107 m). Temperate Northern Pacific, Temperate South America, Temperate Southern Africa, Temperate Australasia. 16 species (Anker and Ahyong 2007b: antitropical species; Hart 1964: key to 7 American species; Nomura and Komai 2000: differences between 14 species tabulated).

Bruceopsis Anker, 2010 Diagnosis. Rostrum elongate, tapering; orbital hoods present, eyestalks partly visible in dorsal view; extraorbital teeth triangular. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds unequal or equal, asymmetrical or subsymmetrical; with carpo-meral rows of setae; chela cylindrical.

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Pereopods 1–4 coxae with blunt process alongside basis. Maximum cl. 7.7 mm. Subtidal (3–7 m). Central Indo-Pacific (Guam). 2 species (Anker 2010a).

Caligoneus Komai & Fujita, 2018 Diagnosis. Rostrum triangular, longer than eyestalks, with elevated midline, triangular in cross-section; orbital hoods absent, eyestalks mostly visible in dorsal view; extraorbital teeth acute. Carapace posterodorsal margin with 1 or 2 postrostral median spines. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds equal, symmetrical; chela both of minor form. Telson posterior margin deeply cleft. Maximum cl. 5.8 mm. Marine anchialine caves (12–17 m). Central Indo-Pacific (Ryukyu Is, Japan). 1 species (Komai and Fujita 2018).

Coronalpheus Wicksten, 1999 Diagnosis. Rostrum triangular, as long as wide, reaching distal margin of cornea; orbital hoods absent, eyestalks mostly visible in dorsal view (divergent); cornea spherical, pigmented, with distal tubercle. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds asymmetrical; chela wide, flat. Maximum cl. 6.6 mm (Pl. 6g). Subtidal (15 m, under rocks). Tropical Eastern Pacific (Galapagos Is). 1 species (Wicksten 1999).

Coutieralpheus Anker & Felder, 2005 Diagnosis. Rostrum triangular, as long as wide, reaching distal margin of cornea; orbital hoods absent, eyestalks mostly visible in dorsal view (divergent); cornea spherical, pigmented, with distal tubercle. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds asymmetrical; chela wide, flat. Maximum cl. 6.6 mm. Intertidal (with callianassoid ghost shrimp burrows). Tropical Atlantic. 1 species (Anker and Felder 2005).

Crosnierocaris Anker, 2022 Diagnosis. Rostrum spiniform, usually about half carapace length. Pleonites 3–5 pleura with 2–4 small spines along posterior margin. Maximum cl. 5.3 mm. Slope (240–255 m). Western and Central Indo-Pacific. 1 species (Anker 2022).

Deioneus Dworschak, Anker & Abed-Navandi, 2000 Diagnosis. Rostrum triangular; extraorbital teeth absent (or invisible from dorsal view). Carapace with horizontal anterolateral suture. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds subequal, asymmetrical, flexed; chela cylindrical. Maximum cl. 6.3 mm. Intertidal (with callichirid ghost shrimp burrows). Temperate Northern Atlantic (Cape Verde Is). 1 species (Dworschak et  al. 2000).

Fenneralpheus Felder & Manning, 1986 Diagnosis. Rostrum absent, front evenly convex; orbital hoods broadly rounded, forming part of sinuous anterior margin of carapace. Carapace with horizontal anterolateral suture; pterygostomial angle rounded, or produced. Pleonite 6 pleuron with movable plate on posteroventral angle. Antennular peduncle article 1 as long or longer than article 2. Chelipeds asymmetrical, flexed; chela cylindrical; fingers slender, not gaping, with rounded, cuspate, obliquely oriented ridges, intermeshing near mid-length. Maximum cl. 13 mm. Intertidal–shelf (0–31 m). Tropical W Atlantic. 2 species (Anker 2011c; Felder and Manning 1986).

Harperalpheus Felder & Anker, 2007 Diagnosis. Rostrum triangular. Pterygostomial angle produced. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds equal, symmetrical; with mesial carpal rows of short setae; chela cylindrical. Uropodal exopod suture weakly defined, marginal spiniform seta at end of anterior margin. Maximum cl. 7.8 mm. Intertidal, subtidal (0–21 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Anker and Lazarus 2017; Felder and Anker 2007).

Jengalpheops Anker & Dworschak, 2007 Diagnosis. Rostrum broadly triangular; orbital hoods absent, eyestalks mostly visible in dorsal view. Carapace with horizontal anterolateral suture. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds equal, symmetrical; with mesial carpal rows of short setae; chela elongate, linear. Maximum cl. 5.5 mm. Intertidal (in burrows of callichirid ghost-shrimp, Glypturus armatus (A. Milne-Edwards, 1870)). Central Indo-Pacific. 1 species (Anker and Dworschak 2007).

Leptalpheus Williams, 1965 Diagnosis. Rostrum absent, front evenly convex, or minute. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds asymmetrical, flexed; chela cylindrical; fingers strongly curved or forceps-shaped, sometimes gaping, unevenly toothed. Minor chela fingers long, slender. Uropodal exopod suture with deep incision such that posterior margin ends in strong tooth. Maximum cl. 8.2 mm (Pl. 6h). Intertidal, subtidal (0–5 m, in burrows of laomediid, upogebiid or callichirid shrimps, or stomatopods). Tropical W Atlantic, Indo-West Pacific, Tropical Eastern Pacific. 18 species (Vera Caripe et al. 2021).

Leptathanas De Grave & Anker, 2008 Diagnosis. Rostrum minute. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds asymmetrical, flexed; chela swollen at midpoint, narrow at base of angled fingers, folding against laterally expanded merus. Minor chela fingers short,

6 – Caridea – shrimps

stout. Uropodal exopod suture incomplete, with spiniform seta at anterior end. Maximum cl. 3.4 mm. Subtidal, estuarine (in burrows of upogebiid shrimps). Tropical E Atlantic (Niger Delta). 1 species (De Grave and Anker 2008).

Leslibetaeus Anker, Poddoubtchenko & Wehrtmann, 2006 Diagnosis. Rostrum absent, front sinuous. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds equal, symmetrical; carpus with mesial rows of short setae; chela robust, inflated. Maximum cl. 3.4 mm (Pl. 6j). Intertidal, subtidal (0–1 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Anker 2011a).

Metabetaeus Borradaile, 1899 Diagnosis. Rostrum triangular; orbital hoods with anterior supraocular spine; cornea weakly pigmented. Pleonite 6 pleuron without movable plate on posteroventral angle. Mandible incisor with conspicuous dark-brown spot (visible in external view of mouthparts). Chelipeds equal, symmetrical or subsymmetrical; major cheliped ischium with row of spiniform setae on upper margin; chela cylindrical. Uropodal endopod with sharp tooth at end of anterior margin. Uropodal exopod suture with strong tooth near anterior margin and spiniform seta. Maximum cl. 8 mm. Intertidal, subtidal (0–17 m), marine anchialine caves, marine and brackish pools. Indo-West Pacific. 3 species. (Anker 2010b: key to species)

Metalpheus Coutière, 1908 Diagnosis. Rostrum triangular; orbital hoods inflated, completely covered by carapace dorsally and partly anteriorly. Pleonite 6 pleuron without movable plate on posteroventral angle. Maxilliped 3 propodus broad, flat, suboperculiform. Chelipeds asymmetrical; dactylus with molar-shaped tooth fitting into socket. Maximum cl. 4 mm. Intertidal, subtidal (0–20 m). Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific, Temperate Australasia. 3 species (Banner and Banner 1982: key to species).

Mohocaris Holthuis, 1973 Diagnosis. Rostrum absent, front sinuous. Carapace posterodorsal margin with 1 median and 2 submedian granular ridges in anterior half. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds equal, symmetrical; dactylus flat, discoid, with narrow tapering apex. Uropodal endopod more than 2.5 times as long as exopod, ending in ribbon-like extension. Maximum cl. 8 mm. Shelf (64–131 m). Tropical W Atlantic. 1 species (Holthuis 1973).

Nennalpheus Banner & Banner, 1981 Diagnosis. Rostrum narrow, acute; orbital hoods present, eyestalks partly visible in dorsal view; extraorbital teeth absent. Pleonite 6 pleuron with or without movable plate on posteroventral

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angle. Chela cylindrical, inverted, dactylus below fixed finger; fingers with rounded fitting teeth along proximal half of cutting edges, dactylus with distal hook. Maximum cl. 10 mm. Intertidal–shelf (0–208 m). Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific. 3 species (Anker 2019; Chace 1988).

Notalpheus Méndez G. & Wicksten, 1982 Diagnosis. Rostrum triangular; orbital hoods present, eyestalks partly visible in dorsal view. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds asymmetrical; chela elongate, dilating, dactylus flat, longer and wider than palm, chela inverted, dactylus below fixed finger; fingers with finely serrate cutting edges, dactylus broad, laminar. Maximum cl. 11 mm. Shelf (143 m). Temperate South America (Peru). 1 species (Méndez G. and Wicksten 1982).

Orygmalpheus De Grave & Anker, 2000 Diagnosis. Rostrum absent, front sinuous. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds equal, subsymmetrical; chela elongate, linear; fingers slender. Maximum cl. 3.5 mm. Subtidal (10 m, in burrow with goby). Central Indo-Pacific. 1 species (De Grave and Anker 2000).

Pachelpheus Anker, 2020 Diagnosis. Rostrum absent, front evenly convex. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds equal, symmetrical; chela cylindrical. Uropodal exopod suture with 2 strong spiniform setae at anterior end. Maximum cl. 3.5 mm. Subtidal (1–1.5 m, in burrow). Tropical Eastern Pacific. 1 species (Anker 2020a).

Parabetaeus Coutière, 1897 Diagnosis. Rostrum absent, front with acute extraorbital teeth. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds equal, symmetrical, flexed; major chela merus flexor margin with 2 sharp distal teeth; chela cylindrical; fingers serrated. Telson posterior margin produced as median triangle. Maximum cl. 8.2 mm. Intertidal–shelf (0–36 m). Tropical Atlantic, Indo-West Pacific. 4 species (Anker 2015; Nomura and Anker 2001: diagnosis, species). Neoalpheopsis Banner, 1953 is a junior synonym.

Pomagnathus Chace, 1937 Diagnosis. Rostrum acute; orbital hoods inflated, completely covered by carapace dorsally and partly anteriorly. Pleonite 6 pleuron without movable plate on posteroventral angle. Maxilliped 3 ischium-merus operculiform. Chelipeds asymmetrical; chela robust, inflated; dactylus with molar-shaped tooth fitting into socket. Pereopods without epipods. Maximum cl. 5 mm. Subtidal (5 m, obligate symbiont of pocilloporid corals). Tropical Eastern Pacific. 1 species (Chace 1937).

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Potamalpheops Powell, 1979 Diagnosis. Rostrum acute; orbital hoods absent, eyestalks mostly visible in dorsal view; extraorbital teeth acute. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds subequal, subsymmetrical, flexed or extended forwards; with mesial carpal rows of short setae; chela robust, inflated or elongate, linear. Uropodal exopod suture with numerous crenellations along anterior two-thirds of length. Maximum cl. 6 mm. Intertidal, subtidal (0–6 m), freshwater, estuarine, marine anchialine caves (rivers, mangroves, peat flats). Tropical Atlantic, Central Indo-Pacific. 16 species. Potamalpheops is the only alpheid genus with species from fresh water (Anker 2008). Other species occur in brackish and marine environments, some in anchialine caves (Anker 2005; Christodoulou et  al. 2019; Marin 2021a; Yeo and Ng 1997).

Prionalpheus Banner & Banner, 1960 Diagnosis. Rostrum triangular. Pterygostomial angle produced. Pleonite 6 pleuron with movable plate on posteroventral angle. Mandible without molar; incisor widened distally, with asymmetrical dentition; without palp. Chelipeds asymmetrical; chela robust, inflated. Uropodal exopod suture toothed along anterior half. Pereopods without epipods. Maximum cl. 2.2 mm. Subtidal, shelf (1–143 m). Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific. 7 species (Alvarez et al. 1996; Banner and Banner 1971).

Pseudalpheopsis Anker, 2007 Diagnosis. Rostrum elongate, tapering; extraorbital teeth triangular. Pterygostomial angle rounded, with small marginal tooth dorsal to angle. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds equal, symmetrical; chela robust, inflated; fingers cutting edges serrated on proximal two-thirds, followed by broad hiatus and large subdistal tooth. Maximum cl. 5.2 mm. Subtidal (9 m). Tropical Atlantic (British Virgin Is). 1 species (Anker 2007).

Pseudathanas Bruce, 1983 Diagnosis. Rostrum triangular; orbital hoods present, eyestalks partly visible in dorsal view. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds asymmetrical, flexed; chela robust, inflated. Uropodal exopod suture with numerous long movable spines. Maximum cl. 3 mm. Intertidal. Central Indo-Pacific (NT, Australia). 1 species (Bruce 1983).

Pterocaris Heller, 1862 Diagnosis. Rostrum triangular; orbital hoods absent, eyestalks mostly visible in dorsal view. Carapace and pleon strongly compressed dorsoventrally; carapace with anterolateral projections;

dorsal integument setose. Chelipeds symmetrical; chela cylindrical. Maximum cl. 8 mm. Subtidal (uncertain). Central Indo-Pacific. 1 species. Pterocaris typica Heller, 1862, is an exceptionally unusual flat setose shrimp, never seen since its discovery (Anker and Dworschak 2001).

Racilius Paul’son, 1875 Diagnosis. Rostrum triangular; orbital hoods inflated, completely covered by carapace dorsally and partly anteriorly; orbital hoods with anterior supraocular spine. Carapace and pleon strongly laterally compressed, with high median dorsal carina. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds subequal, subsymmetrical; chela wide, flat; dactylus with molar-shaped tooth fitting into socket. Maximum cl. 5 mm. Intertidal, subtidal (0–3 m, associated with corals, Galaxea). Western and Central Indo-Pacific. 1 species (Banner and Banner 1973).

Richalpheus Anker & Jeng, 2006 Diagnosis. Rostrum absent; orbital hoods broadly rounded, forming part of sinuous anterior margin of carapace. Pleonite 6 pleuron with more or less marked suture separating posteroventral triangular area. Chelipeds asymmetrical, flexed; chela elongate, linear; dactylus swollen at midpoint, fitting shallow depression on fixed finger. Uropodal exopod suture with deep incision and triangular tooth near posterior margin. Pereopods without epipods. Maximum cl. 5.2 mm. Intertidal, subtidal (0–4 m, associated with burrows of callianassoid ghost shrimps). Indo-West Pacific. 4 species (Anker 2011b: key to 3 species).

Rugathanas Anker & Jeng, 2007 Diagnosis. Rostrum acute; orbital hoods absent, eyestalks mostly visible in dorsal view; orbital hoods with anterior supraocular spine; extraorbital teeth long, acute. Pterygostomial angle angular. Pleonite 6 pleuron with movable plate on posteroventral angle. Mandible with expanded incisor. Chelipeds unequal or subequal, symmetrical (female) or subsymmetrical (male), flexed; chela robust, inflated (tuberculate below in male). At least pereopods 1, 2 with epipods, or pereopods without epipods. Maximum cl. 5.4 mm. Intertidal–shelf (0–36 m, with corals). Western and Central Indo-Pacific. 2 species (Anker and Jeng 2007: key to species and to 7 similar genera)

Salmoneus Holthuis, 1955 Diagnosis. Rostrum triangular; orbital hoods with anterior supraocular spine. Carapace with horizontal anterolateral suture; posterodorsal margin unornamented, or with 1 or 2 postrostral median spines (rare). Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds asymmetrical, flexed; chela

6 – Caridea – shrimps

robust, inflated; fingers slender. Telson posterior margin truncate, or with shallow notch. Maximum cl. 9 mm (Pl. 6i). Intertidal–shelf (0–32 m, sometimes associated with Upogebiidae). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Tropical Eastern Pacific, Temperate Southern Africa, Temperate Australasia. 62 species. Salmoneus is rich in species, varied and widespread, especially in the Indo-West Pacific (Anker 2003; Wang and Sha 2016c; Marin 2021b). Seven species groups are recognised (Anker and Marin 2006; Komai et al. 2021).

Stenalpheops Miya, 1997 Diagnosis. Rostrum triangular, or absent; orbital hoods absent, eyestalks mostly visible in dorsal view. Pleonite 6 pleuron with movable plate on posteroventral angle. Chelipeds equal, symmetrical; with carpo-meral rows of setae; chela cylindrical. Uropodal exopod suture with ~9 crenellations along entire length (exopod with segmented appendage). Maximum cl. 6.6 mm. Intertidal, estuarine (in callianassoid and upogebiid burrows). Temperate Northern Pacific, Central Indo-Pacific. 2 species (Aldea et al. 2021; Anker et al. 2001; Miya 1997).

Synalpheus Bate, 1888 Diagnosis. Rostrum triangular, or acute; orbital hoods produced anteriorly, or with blunt supraocular tooth. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds asymmetrical, extended forwards; dactylus with molar-shaped tooth fitting into socket. Pereopods without epipods. Maximum cl. 13 mm (Pl. 6l). Intertidal–shelf (0–250 m). Cosmopolitan in tropical to temperate seas (associated with crinoids, corals or sponges; many at high densities). 165 species. Synalpheus is a large widespread genus that has been divided into species groups (Coutière 1908). Three of six species groups are probably monophyletic (Hultgren et al. 2014). The taxonomy of some of these groups has received special attention (Anker and Tóth 2008; Dardeau 1984; Hermoso Salazar et al. 2005; Hultgren and Brandt 2015). Keys to species are limited to regional treatments, e.g. the Philippines (Chace 1988), southern Australia (Poore 2004), all Australia (Banner and Banner 1975), China (Wang and Sha 2015). Some species of Synalpheus are noted for their eusociality, occurring in large numbers in host sponges (Didderen et al. 2006; Duffy et al. 2000; Macdonald et al. 2006).

Thuylamea Nguyên, 2001 Diagnosis. Rostrum minute; orbital hoods absent, eyestalks mostly visible in dorsal view. Carapace and pleon subcylindrical, elongate; posterodorsal margin posterodorsal margin with pair of submedian high bluntly-rounded crests in adult. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds subequal, symmetrical; chela cylindrical; dactylus laminar, broader than fixed finger, twisted, cutting edge simple. Maximum cl. 11.5 mm. Subtidal, shelf, marine, estuarine (5–29 m). Temperate Northern Pacific, Central Indo-Pacific. 2 species. The genus was initially

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characterised by the prominent posterior tubercle on the carapace (Nguyên 2001) but this is less obvious in juveniles (Wang and Sha 2016a) and in T. shiranui (see Komai 2018).

Triacanthoneus Anker, 2010 Diagnosis. Rostrum prominent, acute, with subrostral tooth; orbital hoods present, eyestalks partly visible in dorsal view; extraorbital teeth acute. Carapace posterodorsal margin with dorsal median spine, surface behind orbit usually with posthepatic spine. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds asymmetrical, flexed; chela cylindrical; fingers slender. Telson posterior margin truncate, or with shallow notch. Maximum cl. 4.3 mm. Intertidal–shelf (0–40 m), marine anchialine caves. Tropical W Atlantic, Tropical Eastern Pacific. 6 species (Alvarez et al. 2014; Anker 2010c; 2020b: rediagnosis, key to species).

Vexillipar Chace, 1988 Diagnosis. Rostrum triangular; orbital hoods with anterior supraocular spine. Pleonite 6 pleuron without movable plate on posteroventral angle. Mandible without palp. Chelipeds equal, symmetrical; chela cylindrical, chela inverted, dactylus below fixed finger; fingers slender. Maximum cl. 14 mm. Slope (296–875 m, with hexactinellid sponges). Temperate Northern W Pacific (Japan), Central Indo-Pacific (Philippines). 1 species (Chace 1988).

Yagerocaris Kensley, 1988 Diagnosis. Rostrum elongate, tapering; orbital hoods absent, eyestalks mostly visible in dorsal view; cornea weakly pigmented. Pterygostomial angle produced. Pleonite 6 pleuron without movable plate on posteroventral angle. Chelipeds equal, symmetrical; with mesial carpal rows of short setae; chela both of minor form. Maximum cl. 7.2 mm (Pl. 6k). Marine anchialine caves (9–12 m). Tropical Atlantic (Mexico). 1 species (Kensley 1988). References Aldea KQ, Henmi Y, Itani G (2021) Ontogenetic changes in cheliped and uropod morphology of the symbiotic shrimp Stenalpheops anacanthus Miya, 1997 (Decapoda: Caridea: Alpheidae): implications for the taxonomy of the genus. Journal of Crustacean Biology 41, doi:10.1093/ jcbiol/ruab029. Alvarez F, Camacho ME, Villalobos JL (1996) The first species of Prionalpheus from the Eastern Pacific, and new records of caridean shrimp (Crustacea: Decapoda: Caridea) from the western coast of Mexico. Proceedings of the Biological Society of Washington 109, 715–724. Alvarez F, Iliffe TM, Villalobos JL (2014) A new species of the alpheid shrimp genus Triacanthoneus Anker, 2010 (Crustacea: Alpheidae) from Belize. Zootaxa 3768, 88–94. doi:10.11646/zootaxa.3768.1.6 Anker A (2003) Alpheid shrimps from the mangroves and mudflats of Singapore. Part I. Genera Salmoneus, Athanas and Potamalpheops, with the description of two new species (Crustacea: Decapoda: Caridea). Raffles Bulletin of Zoology 51, 283–314.

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Anker A (2005) Presence of the alpheid shrimp genus Potamalpheops Powell, 1979 (Crustacea: Decapoda: Caridea) in south Asia, with description of a new species from Sri Lanka. Raffles Bulletin of Zoology Supplement 12, 31–37. Anker A (2007) Pseudalpheopsis guana gen. nov., sp. nov. (Crustacea: Decapoda), a new alpheid shrimp from the British Virgin Islands, Caribbean Sea. Zoological Studies 46, 428–440. Anker A (2008) A worldwide review of stygobiotic and stygophilic shrimps of the family Alpheidae (Crustacea, Decapoda, Caridea). Subterranean Biology 6, 1–16. Anker A (2010a) Description of a new genus and two new species of alpheid shrimps from Guam (Crustacea, Decapoda). Zootaxa 2372, 389–404. doi:10.11646/zootaxa.2372.1.31 Anker A (2010b) Metabetaeus Borradaile, 1899 revisited, with description of a new marine species from French Polynesia (Crustacea: Decapoda: Alpheidae). Zootaxa 2552, 37–54. doi:10.11646/zootaxa.2552.1.2 Anker A (2010c) A new genus and three new species of alpheid shrimps (Crustacea, Decapoda, Caridea) from the tropical American coasts. Zootaxa 2652, 47–63. doi:10.11646/zootaxa.2652.1.4 Anker A (2011a) The alpheid shrimp genus Leslibetaeus Anker, Poddoubtchenko & Wehrtmann, 2006 in the Western Atlantic, with description of a new species from Tobago (Crustacea, Decapoda). Zootaxa 2734, 63–68. doi:10.11646/zootaxa.2734.1.5 Anker A (2011b) Description of a third species in the alpheid genus Richalpheus Anker & Jeng, 2006 (Crustacea, Decapoda). Zootaxa 3059, 59–68. doi:10.11646/zootaxa.3059.1.4 Anker A (2011c) Six new species and three new records of infaunal alpheid shrimps from the genera Leptalpheus Williams, 1965 and Fenneralpheus Felder & Manning, 1986 (Crustacea, Decapoda). Zootaxa 3041, 1–38. doi:10.11646/zootaxa.3041.1.1 Anker A (2015) Description of a new distinctive species of Parabetaeus Coutière, 1897 (Decapoda: Caridea: Alpheidae) from the Indo-West Pacific. Zootaxa 3957, 585–595. doi:10.11646/zootaxa.3957.5.7 Anker A (2017a) A new deep-water species of Alpheopsis Coutiere, 1897 from the Indo-West Pacific (Decapoda: Alpheidae). Zootaxa 4226, 571–580. doi:10.11646/zootaxa.4226.4.7 Anker A (2017b) Strongly carinate species of Alpheopsis Coutiere, 1897 of the tropical Atlantic and eastern Pacific, with redescription of A. trigona (Rathbun, 1901) and description of three new species (Malacostraca: Decapoda: Alpheidae). Zootaxa 4277, 199–227. doi:10.11646/ zootaxa.4277.2.2 Anker A (2019) The alpheid shrimp genus Nennalpheus Banner & Banner, 1981 in the tropical eastern Atlantic, with description of a new species from Gabon and new records of N. sibogae (De Man, 1910) in the IndoWest Pacific (Malacostraca: Decapoda: Caridea). Zootaxa 4646, 87–100. doi:10.11646/zootaxa.4646.1.5 Anker A (2020a) A remarkable burrow-dwelling alpheid shrimp, new genus and new species, from the tropical eastern Pacific (Malacostraca: Decapoda: Caridea). Zootaxa 4731, 75–88. doi:10.11646/zootaxa.​ 4731.1.5 Anker A (2020b) Taxonomic remarks on the alpheid shrimp genus Triacanthoneus Anker, 2010, with description of a second eastern Pacific species (Malacostraca: Decapoda). Zootaxa 4772, 450–468. doi:10.11646/​ zootaxa.4772.3.2 Anker A (2022) Crosnierocaris athanasoides gen. et sp. nov., a new deepwater alpheid shrimp from the Mozambique Channel (Malacostraca: Decapoda: Caridea). Zootaxa 5105, 269–280. doi:10.11646/zootaxa.​ 5105.2.6 Anker A, Ahyong S (2007a) Description of two species in the alpheid shrimp genus Athanas Leach, 1814, with remarks on A. amazone Holthuis, 1951 (Decapoda, Caridea). Zootaxa 1563, 17–30. doi:10.11646/​ zootaxa.1563.1.2

Anker A, Ahyong S (2007b) A rediagnosis of Athanopsis australis Banner & Banner, 1982, a rare alpheid shrimp from southern Australia, with a phylogeny of Athanopsis Coutière, 1897 and remarks on antitropical distributions in the Alpheidae (Decapoda, Caridea). Crustaceana 80, 685–697. doi:10.1163/156854007781360612 Anker A, Dworschak PC (2001) Redescription and systematic position of Pterocaris Heller, 1862 (Crustacea; Decapoda: Caridea). Raffles Bulletin of Zoology 49, 73–82. Anker A, Dworschak P (2007) Jengalpheops rufus gen. nov., sp. nov., a new commensal alpheid shrimp from the Philippines (Crustacea: Decapoda). Zoological Studies 46, 290–302. Anker A, Felder DL (2005) Description of Coutieralpheus setirostris, new genus, an infaunal alpheid shrimp (Crustacea: Decapoda) from Florida, U.S.A. Crustacean Research 34, 40–52. doi:10.18353/crustacea.​ 34.0_40 Anker A, Jeng M-S (2002) Redefinition of Betaeopsis Yaldwyn, 1971, and invalidation of Hamalpheus Bruce & Iliffe, 1991 (Crustacea: Decapoda: Alpheidae). Proceedings of the Biological Society of Washington 115, 570–584. Anker A, Jeng M-S (2006) Richalpheus palmeri, n. gen., n. sp., an infaunal alpheid shrimp from the Philippines, with redescription of Amphibetaeus jousseaumei (Coutière, 1896) (Decapoda: Caridea). Journal of Crustacean Biology 26, 379–391. doi:10.1651/S-2647.1 Anker A, Jeng M-S (2007) Establishment of a new genus for Arete borradailei Coutière, 1903 and Athanas verrucosus Banner and Banner, 1960, with redefinitions of Arete Stimpson, 1860 and Athanas Leach, 1814 (Crustacea: Decapoda: Alpheidae). Zoological Studies 46, 454–472. Anker A, Komai T (2004) Descriptions of two new species of alpheid shrimps from Japan and Australia, with notes on taxonomy of Automate De Man, Coronalpheus Wicksten and Bermudacaris Anker and Iliffe (Crustacea: Decapoda: Caridea). Journal of Natural History 38, 1895–1914. doi:10.1080/0022293031000156312 Anker A, Lazarus JF (2017) First finding of the shrimp genus Harperalpheus Felder & Anker, 2007 in the eastern Pacific, with description of a new species from Bahia Malaga, Colombia (Malacostraca: Decapoda: Alpheidae). Zootaxa 4329, 196–200. doi:10.11646/zootaxa.4329.2.7 Anker A, Marin I (2006) New records and species of Alpheidae (Crustacea: Decapoda) from Vietnam. Part I. Genus Salmoneus Holthuis, 1955. Raffles Bulletin of Zoology Supplement 54, 295–319. Anker A, Tóth E (2008) A preliminary revision of the Synalpheus paraneptunus Coutière, 1909 species complex (Crustacea: Decapoda: Alpheidae). Zootaxa 1915, 1–28. Anker A, Jeng M-S, Chan T-Y (2001) Two unusual species of Alpheidae (Decapoda: Caridea) associated with upogebiid mudshrimps of Taiwan and Vietnam. Journal of Crustacean Biology 21, 1049–1061. doi:10.1163/20021975-99990197 Anker A, d’Udekem d’Acoz C, Poddoubtchenko D (2005) Description of a new species of Alpheopsis from the Azores, with remarks on A. africana Holthuis, 1952 and other species of the A. trispinosa (Stimpson, 1860) group (Crustacea, Decapoda, Caridea, Alpheidae). Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, Biologie 75, 97–110. Anker A, Ahyong ST, Noël P, Palmer AR (2006a) Morphological phylogeny of alpheid shrimps: parallel preadaptation and the origin of a key morphological innovation, the snapping claw. Evolution 60, 2507–2528. doi:10.1111/j.0014-3820.2006.tb01886.x Anker A, Poddoubtchenko D, Jeng M-S (2006b) Acanthanas pusillus, new genus, new species, a miniature alpheid shrimp with spiny eyes from the Phillipines (Crustacea: Decapoda). Raffles Bulletin of Zoology 54, 341–348. Anker A, Poddoubtchenko D, Marin IN (2006c) On the presence of the alpheid shrimp genus Bermudacaris Anker & Iliffe, 2000 (Crustacea:

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Decapoda: Caridea) in the Pacific Ocean, with description of a new species from Vietnam. Journal of Natural History 40, 1675–1686. doi:10.1080/00222930600937734 Ashrafi H, Ďuriš Z, Anker A (2021) Aretopsis sandybrucei, a new deepwater shrimp (Malacostraca: Decapoda: Caridea: Alpheidae) from the Coral Sea. European Journal of Taxonomy 774, 42–57. doi:10.5852/ ejt.2021.774.1527 Banner AH, Banner DM (1971) Contributions to the knowledge of the alpheid shrimp of the Pacific Ocean. Part XIV. A review of Prionalpheus (Decapoda, Alpheidae) with the description of two new species. Crustaceana 20, 263–270. doi:10.1163/156854071X00058 Banner DM, Banner AH (1973) The alpheid shrimp of Australia. Part I: the lower genera. Records of the Australian Museum 28, 291–382. doi:1 0.3853/j.0067-1975.28.1973.407 Banner DM, Banner AH (1975) The alpheid shrimp of Australia. Part 2: the genus Synalpheus. Records of the Australian Museum 29, 267–389. doi:10.3853/j.0067-1975.29.1975.389 Banner DM, Banner AH (1982) The alpheid shrimp of Australia. Part III: the remaining alpheids, principally the genus Alpheus, and the family Ogyrididae. Records of the Australian Museum 34, 1–357. doi:10.385 3/j.0067-1975.34.1982.434 Bruce AJ (1983) Pseudathanas darwiniensis, New Genus, New Species, an Alpheid Shrimp from the Northern Territory, Australia. Journal of Crustacean Biology 3, 463–471. doi.org/10.1163/193724083X00111 Bruce AJ (1988) Bannereus anomalus, new genus, new species, a deep-sea alpheid shrimp from the Coral Sea. Pacific Science 42, 139–149. Chace FA (1937) The Templeton Crocker Expedition. VII. Caridean decapod Crustacea from the Gulf of California and the west coast of Lower California. Zoologica 22, 109–138. Chace FA (1988) The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 5: family Alpheidae. Smithsonian Contributions to Zoology 466, 1–99. Chace FA, Kensley B (1992) The cardiac notch in decapods. Journal of Crustacean Biology 12, 442–447. doi:10.2307/1549038 Chitre M, Legg M, Koay T-B (2012) Snapping shrimp dominated natural soundscape in Singapore waters. Contributions in Marine Science 2012, 127–134. Christodoulou M, Iliffe TM, De Grave S (2019) A new anchialine cave dwelling species of Potamalpheops Powell, 1979 from the Solomon Islands (Crustacea, Decapoda, Alpheidae). Crustacean Research 48, 11–21. doi:10.18353/crustacea.48.0_11 Coutière H (1908) Sur les Synalphées américaines. Comptes Rendus Hebdomadaires des Séances de l’Académie des Sciences 146, 710–712. Dardeau MR (1984) Synalpheus shrimps (Crustacea: Decapoda: Alpheidae). 1. The Gambarelloides group, with a description of a new species. Memoirs of the Hourglass Cruises 7, 1–125. De Grave S (2004) A new species of Batella (Crustacea, Decapoda, Alpheidae) from New Caledonia. Zoosystema 26, 503–509. De Grave S, Anker A (2000) A new genus and species of alpheid shrimp (Crustacea: Decapoda) from Papua New Guinea. Raffles Bulletin of Zoology 48, 249–256. De Grave S, Anker A (2008) Leptathanas powelli gen. nov., sp. nov, a new infaunal alpheid shrimp associated with upogebiid mudshrimps in Nigeria (Crustacea, Decapoda). Zootaxa 1750, 43–52. doi:10.11646/ zootaxa.1750.1.4 Didderen K, Fransen CHJM, Voogd NJd (2006) Observations on sponge-dwelling colonies of Synalpheus (Decapoda, Alpheidae) of Sulawesi, Indonesia. Crustaceana 79, 961–975. doi:10.1163/ 156854006778815937 Duffy JE, Morrison CL, Rios R (2000) Multiple origins of eusociality among sponge-dwelling shrimps (Synalpheus). Evolution 54, 503–516. doi:10.1111/j.0014-3820.2000.tb00053.x

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Dworschak PC, Anker A, Abed-Navandi D (2000) A new genus and three new species of alpheids (Decapoda: Caridea) associated with thalassinids. Annalen des Naturhistorischen Museums in Wien, B 102, 301–320. Felder DL, Anker A (2007) Description of Harperalpheus pequegnatae, new genus, new species, from the Gulf of Mexico and Atlantic coast of the southeastern USA (Crustacea, Decapoda, Alpheidae). Systematics and Biodiversity 5, 455–463. doi:10.1017/S1477200007002563 Felder DL, Manning RB (1986) A new genus and two new species of alpheid shrimps (Decapoda: Caridea) from south Florida. Journal of Crustacean Biology 6, 497–508. doi:10.2307/1548189 Hart JFL (1964) Shrimps of the genus Betaeus on the Pacific Coast of North America with descriptions of three new species. Proceedings of the United States National Museum 115, 431–466. doi:10.5479/ si.00963801.115-3490.431 Hermoso Salazar M, Wicksten MK, Morrone JJ (2005) Redescriptions and taxonomic notes on species of the Synalpheus townsendi Coutière, 1909 complex (Decapoda: Caridea: Alpheidae). Zootaxa 1027, 1–26. doi:10.11646/zootaxa.1027.1.1 Holthuis LB (1973) Biological results of the University of Miami deep-sea expeditions. 99. Mohocaris, a new genus of alpheid shrimps from the Caribbean region (Crustacea, Decapoda, Natantia). Bulletin of Marine Science 23, 489–495. Holthuis LB (1993) The Recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda) with an appendix on the Order Amphionidacea. revised edn. Nationaal Natuuristorisch Museum, Leiden. Hultgren KM, Brandt A (2015) Taxonomy and phylogenetics of the Synalpheus paraneptunus-species-complex (Decapoda: Alpheidae), with a description of two new species. Journal of Crustacean Biology 35, 547–558. doi:10.1163/1937240X-00002354 Hultgren KM, Hurt C, Anker A (2014) Phylogenetic relationships within the snapping shrimp genus Synalpheus (Decapoda: Alpheidae). Molecular Phylogenetics and Evolution 77, 116–125. doi:10.1016/j. ympev.2014.03.008 Kensley B (1988) New species and records of cave shrimps from the Yucatan Peninsula (Decapoda: Agostocarididae and Hippolytidae). Journal of Crustacean Biology 8, 688–699. doi:10.2307/1548704 Komai T (2018) A new species of the alpheid shrimp genus Thuylamea Nguyên, 2001 (Crustacea: Decapoda: Caridea) from Yatsushiro Sea, Kyushu, Japan. Zootaxa 4378, 387–396. doi:10.11646/zootaxa.​ 4378.3.7 Komai T, Fujita Y (2018) A new genus and new species of alpheid shrimp from a marine cave in the Ryukyu Islands, Japan, with additional record of Salmoneus antricola Komai, Yamada & Yunokawa, 2015 (Crustacea: Decapoda: Caridea). Zootaxa 4369, 575–586. doi:10.11646/ zootaxa.4369.4.7 Komai T, Maenosono T, Naruse T (2021) A new species of alpheid shrimp tentatively assigned to Salmoneus Holthuis, 1955 (Decapoda: Caridea) from the Ryukyu Islands, Japan. Zootaxa 4920, 287–296. doi:10.11646/ zootaxa.4920.2.9 Macdonald KS, III, Ríos R, Duffy JE (2006) Biodiversity, host specificity, and dominance by eusocial species among sponge-dwelling alpheid shrimp on the Belize Barrier Reef. Diversity & Distributions 12, 165–178. doi:10.1111/j.1366-9516.2005.00213.x Marin I (2010) On the presence of the symbiotic alpheid shrimp Aretopsis amabilis in the coastal waters of Nhatrang Bay, Vietnam. Marine Biodiversity Records 3, e74. doi:10.1017/S1755267210000643 Marin IN (2021a) A new species of the genus Potamalpheops (Crustacea: Decapoda: Alpheidae) from the intertidal mangrove swamps of South Vietnam. Arthropoda Selecta 30, 179–191. Marin IN (2021b) A new infaunal species of the alpheid shrimp genus Salmoneus Holthuis, 1955 (Crustacea: Decapoda: Alpheidae) and a

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new crustacean association from anoxic mangrove habitats in southern Vietnam. Arthropoda Selecta 30, 369–385. Marin I, Sheibani R, Sari A (2014) A re-description of rare alpheid shrimp Amphibetaeus jousseaumei (Coutière, 1896) and description of a new species of the genus Athanopsis Coutière, 1897 (Crustacea: Decapoda: Alpheidae) from Iranian coast of the Persian Gulf. Zootaxa 3846, 398–410. doi:10.11646/zootaxa.3846.3.4 Méndez GM, Wicksten MK (1982) Notalpheus imarpe: a new genus and species of snapping shrimp from western South America (Decapoda: Alpheidae. Proceedings of the Biological Society of Washington 95, 709–713. Miya Y (1997) Stenalpheops anacanthus, new genus, new species (Crustacea, Decapoda, Alpheidae) from the Seto Inland Sea and the Sea of Ariake, South Japan. Bulletin of the Faculty of Liberal Arts. Nagasaki University. Nature and Science 38, 145–161. Nguyên VX (2001) A new alpheid shrimp (Crustacea: Decapoda: Alpheidae) from South Vietnam. Zoölogische Mededeelingen 75, 217–228. Nomura K, Anker A (2001) A redescription of Parabetaeus culliereti Coutière and status of the genus Neoalpheopsis Banner (Decapoda: Caridea: Alpheidae). Crustacean Research 30, 43–54. doi:10.18353/crustacea.30.0_43 Nomura K, Komai T (2000) A new alpheid shrimp of the genus Betaeus from the Pacific coast of central Japan (Crustacea: Decapoda: Caridea). Crustacean Research 29, 45–57. doi:10.18353/crustacea.29.0_45 Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Qin S, Yang Y, Qin J, Di C (2019) Research on the cavitation in the snapping shrimp: a review. IOP Conference Series. Earth and Environmental Science 310(052057), 1–6. Vera Caripe J, Pereda L, Anker A (2021) A new species and two new records of symbiotic infaunal alpheid shrimps from the genera Leptalpheus Williams, 1965 and Fenneralpheus Felder & Manning, 1986 (Decapoda: Caridea) from Venezuela. Zootaxa 5061, 177–184. doi: 10.11646/ zootaxa.5061.1.10 Versluis M, Schmitz B, von der Heydt A, Lohse D (2000) How snapping shrimp snap: through cavitating bubbles. Science 289, 2114–2117. doi:10.1126/science.289.5487.2114

Wang Y-R, Sha Z-L (2015) A review of the genus Synalpheus (Crustacea: Decapoda: Caridea: Alpheidae) from China seas. Zoological Systematics 40, 357–435. Wang Y-L, Sha Z-L (2016a) Redescription of the type species of the genus Thuylamea Nguyên, 2001 (Decapoda, Caridea, Alpheidae), based on a female specimen. Crustaceana 89, 1185–1191. doi:10.1163/​1568540300003572 Wang Y-L, Sha Z-L (2016b) A review of the genus Alpheopsis (Decapoda, Caridea, Alpheidae) from the seas of China. Crustaceana 89, 1099– 1113. doi:10.1163/15685403-00003571 Wang Y, Sha Z (2016c) Taxonomic notes on the genus Salmoneus Holthuis, 1955 (Decapoda, Caridea, Alpheidae) from the South China Sea. Crustaceana 89, 1585–1609. doi:10.1163/15685403-00003609 Wicksten MK (1999) A new genus and species of alpheid shrimp (Caridea: Alpheidae) from the Galapagos Islands. Crustacean Research 28, 104–111. doi:10.18353/crustacea.28.0_104 Williams ST, Knowlton N, Weigt LA, Jara JA (2001) Evidence for three major clades within the snapping shrimp genus Alpheus inferred from nuclear and mitochondrial gene sequence data. Molecular Phylogenetics and Evolution 20, 375–389. doi:10.1006/mpev.2001.0976 Yeo DCJ, Ng PKL (1997) The alpheid shrimp genus Potamalpheops Powell, 1979, (Crustacea: Decapoda: Caridea: Alpheidae) from Southeast Asia, with descriptions of three new species. Journal of Natural History 31, 163–190. doi:10.1080/00222939700770111

Barbouriidae Christoffersen, 1987 Figure 6.6 Barbouriidae is one of the hippolytoid families of alpheoid shrimps confined largely to anchialine pools and caves in the Caribbean Sea and eastern Mexico. One exception is Calliasmata pholidota Holthuis, 1973, widespread in coastal saltwater pools in the Indo-West Pacific (Ditter et al. 2020). The composition of the family was justified on the basis of molecular relationships by De Grave et  al.

Fig. 6.6.  Barbouriidae. Carapace: a, Barbouria; b, c, Calliasmata pholidota Holthuis, 1973 (with dorsal view of front, eyestalks); d, e, Parhippolyte antiguensis (Chace, 1972); f, Parhippolyte uveae Borradaile, 1900. Pereopod 2: g, Parhippolyte. Pereopod 3: h, Barbouria. Pereopod 5: i, Parhippolyte.

6 – Caridea – shrimps

(2014) but they were unable to find morphological attributes to separate it from other hippolytoids. De Grave et al. (2014) accepted the monospecific genus Janicea Manning & Hart, 1984, but this was synonymised with Parhippolyte by Ditter et al. (2020). These authors introduced two subfamilies, now formalised as Calliasmatinae Ditter, Mejía-Ortíz & Bracken-Grissom, 2021, for Calliasmata and Barbouriinae for the remaining genera (Ditter et  al. 2021). Both

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subfamilies were diagnosed but the family remained, until now, without a diagnosis. Diagnosis. Rostrum as long as or twice as long as eyestalk, with dorsal teeth, or shorter than eyestalk, unarmed. Eyestalks compact, exposed, or immovable, cornea obsolete. Mandible incisor absent, palp absent or 3-articled. Pereopods 1, 2 dissimilar, each pair symmetrical, first more robust than second. Pereopod 2 merus (usually), carpus and propodus multiarticulate (Fig. 6.6g–i).

Key to genera of Barbouriidae 1. – 2. –

Rostrum usually if present shorter than eyestalk, unarmed. Carapace without antennal spine. Eyestalks flat, immovable, cornea obsolete (Fig. 6.6b, c). Mandibular palp absent���������������������� Calliasmatinae … Calliasmata Rostrum as long as or longer than eyestalk, dentate. Carapace with antennal spine. Eyestalks cylindrical, cornea dark (Fig. 6.6a, d–f). Mandibular palp 3-articled�����������������������������������������������������������������������Barbouriinae … 2 Cornea narrower than eyestalk (Fig. 6.6a)�������������������������������������������������������������������������������������������������������������������Barbouria Cornea wider than eyestalk (Fig. 6.6d, f)�����������������������������������������������������������������������������������������������������������������Parhippolyte Barbouria Rathbun, 1912

Diagnosis. Rostrum with 3–7 dorsal teeth (2–4 postorbital), 3–9 ventral teeth. Cornea narrower than eyestalk. Mandible with palp. Maximum cl. 21 mm. Marine anchialine caves. Tropical W Atlantic. 1 species (Ditter et  al. 2019: variation; Ditter et  al. 2020: synonymy of 2 species; Manning and Hart 1984: rediagnosis; Mejía et al. 2008: compared 2 species).

Calliasmata Holthuis, 1973 Diagnosis. Rostrum shorter than eyestalk, unarmed. Cornea obsolete, eyestalk immovable. Mandible without palp. Maximum cl. 14 mm. Marine anchialine pools and caves, and coastal saltwater pools (0–22 m). Tropical W Atlantic, Indo-West Pacific. 3 species (Ditter et al. 2020: key to species, distribution; Escobar-Briones et al. 1997: compared species; Holthuis 1973).

Parhippolyte Borradaile, 1900 Diagnosis. Rostrum as long as or longer than eyestalk, with 1–5 dorsal teeth (1 or 2 postorbital), 1–4 ventral teeth. Cornea broader than eyestalk. Mandible with palp. Maximum cl. 27 mm. Marine anchialine pools and caves (0–40 m). Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific, Tropical Eastern Pacific. 6 species (Chace 1997: rediagnosis, key to 6 species; Ditter et al. 2020: key to species; Wicksten 1996). References Chace FA (1997) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907–1910, Part 7: Families

Atyidae, Eugonatonotidae, Rhynchocinetidae, Bathypalaemonellidae, Processidae, and Hippolytidae. Smithsonian Contributions to Zoology 587, 1–106. De Grave S, Li CP, Tsang LM, Chu KH, Chan T-Y (2014) Unweaving hippolytoid systematics (Crustacea, Decapoda, Hippolytidae): resurrection of several families. Zoologica Scripta 43, 496–507. doi:10.1111/ zsc.12067 Ditter RE, Erdman RB, Goebel AM, Bracken-Grissom HD (2019) Widespread phenotypic hypervariation in the enigmatic anchialine shrimp Barbouria cubensis (Decapoda: Barbouriidae). Zootaxa 4648, 1–26. doi:10.11646/zootaxa.4648.1.1 Ditter RE, Mejía-Ortíz LM, Bracken-Grissom HD (2020) Anchialine adjustments: an updated phylogeny and classification for the family Barbouriidae Christoffersen, 1987 (Decapoda: Caridea). Journal of Crustacean Biology 40, 401–411. doi:10.1093/jcbiol/ruaa037 Ditter RE, Mejía-Ortíz LM, Bracken-Grissom HD (2021) Diagnosis of a new subfamily within Barbouriidae (Decapoda: Caridea). Journal of Crustacean Biology 41, ruab054. doi:10.1093/jcbiol/ruab054 Escobar-Briones E, Camacho ME, Alcocer J (1997) Calliasmata nohochi, new species (Decapoda: Caridea: Hippolytidae), from anchialine cave systems in continental Quintana Roo, Mexico. Journal of Crustacean Biology 17, 733–744. doi:10.2307/1549376 Holthuis LB (1973) Caridean shrimps found in land-locked saltwater pools at four Indo-West Pacific localities (Sinai Peninsula, Funafuti Atoll, Maui and Hawaii Islands), with the description of one new genus and four new species. Zoologische Verhandelingen 128, 1–48, pl. 1–7. Manning RB, Hart CW (1984) The status of the hippolytid shrimp genera Barbouria and Ligur (Crustacea: Decapoda): a reevaluation. Proceedings of the Biological Society of Washington 97, 655–665. Mejía L, Zarza E, López M (2008) Barbouria yanezi sp. nov., a new species of cave shrimp (Decapoda, Barbouriidae) from Cozumel Island, Mexico. Crustaceana 81, 663–672. doi:10.1163/​156854008784513474 Wicksten MK (1996) Parhippolyte cavernicola, new species (Decapoda: Caridea: Hippolytidae) from the tropical Eastern Pacific, with taxonomic remarks on the genera Somersiella and Koror. Journal of Crustacean Biology 16, 201–207. doi:10.2307/1548941

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Marine Decapod Crustacea

Bythocarididae Christoffersen, 1987 Figure 6.7 Bythocarididae are a family of shrimps from cold or deep water, all except for one species confined to the North Atlantic or Arctic. Bathyhippolyte yaldwyni Hayashi & Miyake, 1970 is from New Zealand. De Grave et al. (2014) recognised the family on the basis of molecular relationships of two of its genera and discussed its synapomorphies.

Diagnosis. Rostrum dentate, or shorter than eyestalk, unarmed. Mandible without palp. Maxilliped 1 exopodal flagellum short (Fig.  6.7g). Maxilliped 3 dactylus flat, dilated (Fig. 6.7i), or curved (Fig. 6.7j). Pereopods 1, 2 dissimilar, each pair symmetrical, first more robust than second. Pereopod 2 carpus multiarticulate, carpus of 7–13 segments. Epipod of maxilliped 1 with reduced dorsal lobe, absent from maxilliped 2. Implicit generic attributes. Eyestalk not concealed by carapace. Mandibular incisor absent.

Key to genera of Bythocarididae 1. – 2. – 3. –

Rostrum usually shorter than eyestalks, triangular, with pair of lateral supraorbital spines (Fig. 6.7d, e)�����������Bythocaris Rostrum markedly longer than eyestalks, tapering��������������������������������������������������������������������������������������������������������������������2 Rostrum spike-like, unarmed. Carapace with supraorbital spine (Fig. 6.7b, c)������������������������������������������������ Bythocarides Rostrum with dorsal teeth, usually with 1 ventral tooth. Carapace without supraorbital spine�����������������������������������������3 Rostrum with 15 dorsal teeth. Eyestalk concealed by carapace (Fig. 6.7a). Mandible without incisor������������������������������������������������������������������������������������������������������������������������������������������������������������������ Bathyhippolyte Rostrum with 7 dorsal teeth. Eyestalk not concealed by carapace (Fig. 6.7f). Mandible with incisor���������������������������������������������������������������������������������������������������������������������������������������������������������������������� Cryptocheles Bathyhippolyte Hayashi & Miyake, 1970

Diagnosis. Rostrum straight, with 15 dorsal teeth (7 post orbital), without or with 1 ventral tooth. Carapace without supraorbital spine. Eyestalk concealed by carapace. Antennule peduncle article 3 without sharp dorsodistal tooth. Pereopod 2 carpus with 10 segments. Maximum cl. 8 mm. Shelf, slope (219–1110 m depth). Temperate Australasia (New Zealand). 1 species (Hayashi and Miyake 1970).

Bythocarides Sokolov, 2002 Diagnosis. Rostrum more than half a long as carapace, spike-like, unarmed. Carapace with supraorbital spine. Antennule peduncle

article 3 with sharp dorsodistal tooth. Pereopod 2 carpus with 13 segments. Maximum cl. 8 mm. Slope (945 m depth). Arctic. 1 species (Sokolov 2002).

Bythocaris G.O. Sars, 1870 Diagnosis. Rostrum short, triangular, with pair of supraorbital lateral spines. Antennule peduncle article 3 without sharp dorsodistal tooth. Pereopod 2 carpus with 8–11 segments. Maximum cl. 19 mm. Shelf–bathyal (50–3965 m). Arctic, Temperate Northern Atlantic. 17 species. Three keys to species are available (Abele and Martin 1989; Fransen 1993; Sokolov 2000). Some species are sexually dimorphic (Sokolov and Spiridonov 2006).

Fig. 6.7.  Bythocarididae. Carapace: a, Bathyhippolyte yaldwyni Hayashi & Miyake, 1970; b, c, Bythocarides menshutkinae Sokolov, 2002 (with dorsal view of front, eyestalks); d, e, Bythocaris (with dorsal view of front, eyestalks); f, Cryptocheles. Maxilliped 1: g, Bathyhippolyte. Maxilliped 2: h, Bathyhippolyte. Maxilliped 3: i, Bythocaris; j, Bathyhippolyte.

6 – Caridea – shrimps

Cryptocheles G.O. Sars, 1870 Diagnosis. Rostrum straight, with 7 dorsal teeth (2 post orbital), with 1 ventral tooth. Carapace without supraorbital spine. Mandibular incisor present. Antennule peduncle article 3 with sharp dorsodistal tooth. Pereopod 2 carpus with 7 segments. Maximum cl. 5 mm. Shelf, slope (165–640 m). Arctic, Temperate Northern Atlantic. 1 species. Holthuis (1993) reproduced a habitus drawing from Sars (1912). References Abele LG, Martin JW (1989) American species of the deep-sea shrimp genus Bythocaris (Crustacea, Decapoda, Hippolytidae). Bulletin of Marine Science 45, 26–51. De Grave S, Li CP, Tsang LM, Chu KH, Chan T-Y (2014) Unweaving hippolytoid systematics (Crustacea, Decapoda, Hippolytidae): resurrection of several families. Zoologica Scripta 43, 496–507. doi:10.1111/zsc.12067 Fransen CHJM (1993) Notes on the genus Bythocaris G.O. Sars, with description of a new species. Zoölogische Mededeelingen 67, 567–599. Hayashi K, Miyake S (1970) Bathyhippolyte yaldwyni n. gen., n. sp., a deepsea hippolytid (Decapoda, Natantia) from New Zealand. Transactions of the Royal Society of New Zealand 12, 41–47. Holthuis LB (1993) The Recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda) with an appendix on the Order Amphionidacea. revised edn. Nationaal Natuuristorisch Museum, Leiden. Sars GO (1912) On the genera Cryptocheles and Bythocaris G. O. Sars with description of the type species of each genus. Archiv för Mathematik og Naturvidenskab 32, 1–19. Sokolov V (2000) Deep-sea shrimps of the genus Bythocaris G.O. Sars in the collections of Russian museums, with the description of a new species (Crustacea: Decapoda: Hippolytidae). Zoölogische Mededeelingen 74, 403–468. Sokolov VI (2002) Description of Bythocarides menshutkinae gen. nov., sp. nov. (Decapoda, Hippolytidae). Crustaceana 75, 137–144. doi:10.1163/​ 15685400252902353

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Sokolov VI, Spiridonov VA (2006) Development of external sexual characters of deep sea Bythocaris shrimps (Crustacea: Decapoda: Hippolytidae). Journal of the Marine Biological Association of the United Kingdom 86, 349–359. doi:10.1017/S0025315406013208

Hippolytidae Bate, 1888 Figures 6.8, 6.9, Plate 7a–n Hippolytidae were once a large family of ~40 genera (Chace 1997; Holthuis 1993) that has been variously subdivided. Christoffersen (1987) based his classification on a cladogram using morphological data, recognising five families within the traditional Hippolytidae. His Hippolytidae s.s. contained nine genera. The most recent phylogenetic analysis based on molecular data recognised six families, including Hippolytidae s.s., with 17 genera (De Grave et al. 2014). These authors noted some anomalous genera within this group (Alope, Saron, Leontocaris, Merhippolyte, Eumanningia) but did not suggest a morphological synapomorphy or diagnostic character for Hippolytidae s.s. Wicksten and Hancock (2021) showed Paralatreutes Kemp, 1925 to be a synonym of Latreutes. Diagnosis. Rostrum prominent, dentate dorsally and usually ventrally. Antennular lateral flagellum usually with accessory branch (Fig.  6.8a, g). Pereopods 1, 2 dissimilar, each pair symmetrical, first more robust than second, or dissimilar, second pair asymmetrical, propodus swollen on one side (Leontocaris only). Pereopod 2 carpus multiarticulate, with 2–17 segments (exceptionally 90, usually 3). Implicit generic attributes. Pleonite 6 pleuron without articulated plate at posteroventral angle. Mandible with incisor; mandible without palp. Maxilliped 3 with exopod. Telson with 3–6 pairs of dorsolateral spiniform setae. Pereopods 1–4 without arthrobranchs.

Key to the genera of Hippolytidae 1. – 2. – 3. – 4. – 5. – 6. – 7. –

Pereopods 2 strongly asymmetrical, carpus 4-segmented. Major pereopod 2 with elongate, robust propodus and enlarged, chopper-shaped dactylus (Fig. 6.8f)�������������������������������������������������������������������������������������� Leontocaris Pereopods 2 more or less symmetrical (Fig. 6.9p–r)�������������������������������������������������������������������������������������������������������������������2 Pereopods 1–4 with arthrobranchs�����������������������������������������������������������������������������������������������������������������������������������������������3 Pereopods 1–4 without arthrobranchs�����������������������������������������������������������������������������������������������������������������������������������������6 Pleonite 6 pleuron with articulated plate at posteroventral angle (Fig. 6.8g)�������������������������������������������������������������������������4 Pleonite 6 pleuron without articulated plate at posteroventral angle (Fig. 6.9c)��������������������������������������������������������������������5 Mandible with incisor (Fig. 6.9n)������������������������������������������������������������������������������������������������������������������������������������������ Saron Mandible without incisor (Fig. 6.9o)���������������������������������������������������������������������������������������������������������������������������� Nauticaris Carapace with fixed postrostral spines (Fig. 6.9g). Pereopod 2 carpus of 13–16 segments���������������������������� Merhippolyte Carapace with 3 strong movable postrostral spines (Fig. 6.9c). Pereopod 2 carpus of ~90 segments����������Eumanningia Mandible with palp (Fig. 6.9n)�������������������������������������������������������������������������������������������������������������������������������������������������������7 Mandible without palp (Fig. 6.9o)������������������������������������������������������������������������������������������������������������������������������������������������10 Carapace without supraorbital spine (Fig. 6.9a, b)����������������������������������������������������������������������������������������������������������������������8 Carapace with supraorbital spines (Figs 6.8a, b, d, 6.9f, l)���������������������������������������������������������������������������������������������������������9

84

8. – 9. – 10. – 11. – 12. – 13. – 14. –

Marine Decapod Crustacea

Pereopod 2 carpus of 2 segments, merus undivided����������������������������������������������������������������������������������������������������Caridion Pereopod 2 carpus of 9–12 segments, merus of 3 segments (Fig. 6.9p)������������������������������������������������������������������Chorismus Carapace smooth (Fig. 6.8a). Mandibular palp of 3 articles����������������������������������������������������������������������������������������������Alope Carapace spinulose (Fig. 6.9l). Mandibular palp of 2 articles�������������������������������������������������������������������������������� Trachycaris Mandible with incisor (Fig. 6.9n)�������������������������������������������������������������������������������������������������������������������������������������������������11 Mandible without incisor (Fig. 6.9o)�������������������������������������������������������������������������������������������������������������������������������������������13 Rostrum triangular, downturned, unarmed (Fig. 6.9f). Pereopod 2 carpus of 2 segments��������������������������������Phycocaris Rostrum laminar, dentate (Figs 6.8d, 6.9j). Pereopod 2 carpus of 3 or 4 segments������������������������������������������������������������ 12 Carapace with supraorbital spine (Fig. 6.8d). Pereopod 2 carpus of 3 segments (Fig. 6.9q)���������������������������������Hippolyte Carapace without supraorbital spine (Fig. 6.9j). Pereopod 2 carpus of 4 segments��������������������������������������������������Thorella Pereopods 3–5 dactyli with 4 large spines (Figs 6.8c, 6.9s). Scaphocerite lateral margin with row of spiniform setae (Fig. 6.9m)������������������������������������������������������������������������������������������������������������������������������������ Gelastocaris Pereopods 3–5 dactyli simple. Scaphocerite lateral margin without spiniform setae���������������������������������������������������������14 Maxilliped 3 with exopod���������������������������������������������������������������������������������������������������������������������������������Latreutes (in part) Maxilliped 3 without exopod�������������������������������������������������������������������������������������������������������������������������������������������������������15

Fig. 6.8.  Hippolytidae. a, b, Alope orientalis (De Man, 1890) (with dorsal view of front); c, Gelastocaris paronae (Nobili, 1905); d, Hippolyte australiensis (Stimpson, 1860); e, Latreutes compressus (Stimpson, 1860) female; f, Leontocaris yarramundi Taylor & Poore, 1998; g, Nauticaris marionis Bate, 1888.

6 – Caridea – shrimps

85

Fig. 6.9.  Hippolytidae. Carapace: a, Caridion gordoni (Bate, 1856); b, Chorismus antarcticus (Pfeffer, 1887); c, Eumanningia pliarthron Crosnier, 2000 (with pleon); d, e, Gelastreutes crosnieri Bruce, 1990 (with ventral view of left carapace anterolateral angle); f, Phycocaris simulans Kemp, 1916; g, Merhippolyte chacei Kensley, Tranter & Griffin, 1987; h, Saron marmoratus (Olivier, 1811); i, Latreutes bicornis (Kemp, 1925); j, Thorella cobourgi Bruce, 1982; k, Tozeuma tomentosum (Baker, 1904); l, Trachycaris rugosa (Bate, 1888) female. Scaphocerite: m, Gelastocaris. Mandible: n, Chorismus (with incisor and 3-articled palp); o, Gelastreutes (without incisor and palp). Pereopod 2: p, Chorismus; q, Hippolyte; r, Merhippolyte. Pereopod 3, propodus, dactylus: s, Gelastocaris.

15. – 16. –

Carapace anterolateral angle smooth (Fig. 6.9k). Pereopods 1–4 without epipods�������������������������������������������������Tozeuma Carapace anterolateral angle denticulate (Fig. 6.9d, e, i). Pereopods 1–4 with epipods������������������������������������������������������16 Anterior carapace strongly flattened ventrally, anterior lateral angle produced as large flattened tooth. Antennal spine fixed (Fig. 6.9d, e)������������������������������������������������������������������������������������������������������������������������ Gelastreutes Anterior carapace not strongly flattened ventrally, anterior lateral angle not produced. Antennal spine movable (Fig. 6.9i)�����������������������������������������������������������������������������������������������������������������������������������������Latreutes (in part) Alope White, 1847

Caridion Goës, 1864

Diagnosis. Rostrum spike-like. Carapace abruptly depressed alongside supraorbital tooth, with supraorbital spines. Mandibular palp of 3 articles. Pereopod 2 carpus of 10–17 segments. Telson with 2 pairs of dorsolateral spiniform setae. Maximum cl. 9 mm (Pl. 7a). Intertidal, shallow subtidal. Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 2 species. Alope orientalis (De Man, 1890) is widespread while A. spinifrons (H. Milne Edwards, 1837) is confined to New Zealand (Poore 2004).

Diagnosis. Rostrum about half length of carapace, slightly downturned, with 5–8 dorsal teeth, usually 1 ventral tooth. Pterygostomial spine absent. Mandibular palp of 3 articles. Pereopod 2 carpus of 2 segments. Pereopods 1–4 without epipods. Maximum cl. 6 mm. Subtidal–slope (0–400 m). Temperate Northern Atlantic. 2 species (Smaldon et al. 1993: illustrations of 2 species).

86

Marine Decapod Crustacea

Chorismus Bate, 1888 Diagnosis. Rostrum curving slightly dorsally, with 7–9 dorsal teeth, 6–9 ventral teeth. Pterygostomial spine present. Mandibular palp of 3 articles. Pereopod 2 carpus of 11 segments, merus of 3 segments. Telson with 3 pairs of dorsolateral spiniform setae. Maximum cl. 17 mm. Subtidal–slope (15–915 m). Southern Ocean. 2 species (Komai et  al. 1996: illustrations of C. antarcticus (Pfeffer, 1887), species compared).

Eumanningia Crosnier, 2000 Diagnosis. Rostrum as long as carapace, distally elevated, with 1 subdistal dorsal tooth, 5 ventral teeth. Carapace middorsal line with 3 strong movable postrostral spines, pterygostomial spine present. Pleonite 3 produced posteriorly into long, sharp, curved tooth. Mandibular palp of 3 articles. Pereopod 2 carpus of ~90 segments, merus of 40 segments. Telson with 3 pairs of dorsolateral spiniform setae. Pereopods 1–4 with arthrobranchs, without epipods. Maximum cl. 18 mm. Slope (900 m). Central Indo-Pacific. 1 species (Crosnier 2000).

obligate or facultative symbionts of other organisms, such as gorgonians and crinoids (Gan and Li 2019). These authors reviewed the taxonomic literature. Hippolyte was rediagnosed by Chace (1997). Atlantic species can be identified using the keys of d’Udekem d’Acoz (2007) or Fransen and De Grave (2019). Species from the Indo-West Pacific have not been reviewed. Molecular phylogenies of some species from the Indo-West Pacific (Gan and Li 2019) and of the genus as a whole (Terossi et  al. 2017) reveal species groups well linked to biogeography.

Latreutes Stimpson, 1860 Diagnosis. Rostrum with elevated dorsal margin, with conspicuous ventral blade projecting posteroventrally (sexually dimorphic). Branchiostegal or hepatic tooth present, pterygostomial spine prominent. Mandible without incisor. Maxilliped 3 with or without exopod. Pereopod 2 carpus of 3 segments. Pereopods 1–4 with epipods. Maximum cl. 8 mm (Pl. 7d). Intertidal–shelf (0–110 m). Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Temperate Southern Africa, Temperate Australasia (and on floating oceanic algae). 19 species (Chace 1997: diagnosis, key to females of 15 species; Terossi et al. 2019: key to Atlantic species).

Gelastocaris Kemp, 1914 Diagnosis. Rostrum with single movable subdistal spine, with strong unarmed ventral blade. Carapace middorsal line with smooth middorsal carina, pterygostomial spine prominent. Scaphocerite lateral margin with row of spiniform setae. Mandible without incisor. Maxilliped 3 without exopod. Pereopod 2 carpus of 3 segments. Pereopods 3–5 dactyli with 4 strong spines, 1 proximal, 1 distal, pair lateral. Telson with 2 pairs of dorsolateral spiniform setae. Maximum cl. 6 mm. Subtidal, shelf (1–30 m, associated with sponges). Western and Central Indo-Pacific. 1 species (Chace 1997).

Leontocaris Stebbing, 1905

Diagnosis. Rostrum prominent, deep, with dorsal distal tooth, with smooth ventral blade. Carapace anterolateral angle denticulate, middorsal line with high toothed carina over anterior half, pterygostomial spine prominent. Mandible without incisor. Maxilliped 3 without exopod. Pereopod 2 carpus of 3 segments. Pereopods 1–4 with epipods. Maximum cl. 6 mm. Shelf (64–120 m). Central Indo-Pacific. 1 species (Bruce 1990).

Diagnosis. Rostrum half or as long as carapace, with numerous dorsal and ventral teeth. Scaphocerite lateral margin denticulate. Mandibular palp of 1 article, or mandibular palp of 2 articles. Maxilliped 3 without exopod. Pereopods 2 strongly asymmetrical, carpus 4-segmented. Major pereopod 2 with elongate, robust propodus and enlarged, chopper-shaped dactylus. Maximum cl. 11 mm (Pl. 7e). Slope–bathyal (240–2182 m, associated with deep-sea corals). Temperate Northern and Tropical Atlantic, Western Indo-Pacific, Temperate South America, Temperate Southern Africa, Temperate Australasia. 9 species. Species of Leontocaris are instantly recognised by the asymmetrical second pereopod. Poore (2009) speculated on the interaction between the carpus, propodus and dactylus of the major pereopod 2 without concluding what it was used for. Relationships between the species have been addressed by morphological phylogenies (Fransen 2001; Komai and Chan 2010; Poore 2009). The species can be identified from keys (­Cardoso and Fransen 2012; Taylor and Poore 1998) but one species is doubtful (Chan et al. 2016).

Hippolyte Leach, 1814

Nauticaris Bate, 1888

Diagnosis. Rostrum variable, with ventral blade not conspicuously developed. Carapace middorsal line with smooth middorsal carina, with supraorbital spines, branchiostegal or hepatic tooth present, pterygostomial spine prominent. Pereopod 2 carpus of 3 segments. Telson with 2 pairs of dorsolateral spiniform setae. Maximum cl. 15 mm (Pl. 7b, c). Intertidal–slope (0–220 m). Cosmopolitan, including Arctic. 39 species. Most species inhabit seaweed beds, but some are

Diagnosis. Rostrum dentate dorsally, with strong ventral dentate blade. Pterygostomial spine prominent. Pleonite 6 pleuron with articulated plate at posteroventral angle. Mandible without incisor; mandibular palp of 3 articles. Maxilliped 3 without exopod. Pereopod 2 carpus of 13–16 segments. Pereopods 1–4 with arthrobranchs. Maximum cl. 30 mm (Pl. 7f). Subtidal, shelf intertidal–slope (0–746 m). Temperate South America, Temperate Southern Africa, Temperate Australasia,

Gelastreutes Bruce, 1990

6 – Caridea – shrimps

Southern Ocean. 3 species. All species are from subantarctic islands and southern continents (Holthuis 1952; Poore 2004). The ecology of N. marionis Bate, 1888 is well studied (Haley et al. 2017; Pakhomov et al. 2000).

Merhippolyte Bate, 1888 Diagnosis. Rostrum narrow, long, with 3–6 dorsal teeth, 5–10 ventral teeth. Pterygostomial spine absent. Mandibular palp of 3 articles. Pereopod 2 carpus of 13–16 segments. Telson with 3 pairs of dorsolateral spiniform setae. Pereopods 1–4 with arthrobranchs, with epipods. Maximum cl. 13 mm. Shelf, slope (75–1000 m). Temperate Northern and Tropical Atlantic, Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 7 species (Crosnier and Forest 1973: differences between 5 species; Kensley et al. 1987; Ledoyer 1990).

Phycocaris Kemp, 1916 Diagnosis. Rostrum triangular, downturned, unarmed. Carapace middorsal line with smooth middorsal carina, with supraorbital spines, pterygostomial spine prominent. Pereopod 2 carpus of 2 segments. Telson with 2 pairs of dorsolateral spiniform setae. Maximum cl. 2 mm. Intertidal, subtidal (0–20 m). Temperate Northern Pacific, Western and Central Indo-Pacific. 1 species. The only species Phycocaris simulans Kemp, 1916 is sometimes called the hairy shrimp due to its camouflage that closely resembles the small turf of algae within which it can often be found. The species was figured by Gan and Li (2017) and Saponari et al. (2018).

Thorella Bruce, 1982 Diagnosis. Rostrum prominent, with dorsal and ventral teeth. Branchiostegal or hepatic tooth present, pterygostomial spine present. Pereopod 2 carpus of 4 segments. Maximum cl. 1.8 mm. Intertidal, subtidal (0–2 m). Central Indo-Pacific (N Australia). 1 species (Bruce 1982).

Tozeuma Stimpson, 1860 Diagnosis. Rostrum prominent, slender, sometimes longer than slender body, smooth dorsally, with toothed ventral blade. Pterygostomial spine present. Mandible without incisor. Maxilliped 3 without exopod. Pereopod 2 carpus of 3 segments. Pereopods 1–4 without epipods. Maximum cl. 10 mm (Pl. 7g). Intertidal–shelf (0–135 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Western and Central IndoPacific, Temperate Southern Africa, Temperate Australasia. 11 species (Chace 1997: key to species; Poore 2004: illustrations of 4 species).

Trachycaris Calman, 1906 Diagnosis. Rostrum of females deep, truncated, of males elongate, with 4–6 dorsal teeth, 3 ventral teeth. Carapace spinulose, with

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supraorbital spines, pterygostomial spine prominent. Mandibular palp of 2 articles. Pereopod 2 carpus of 2 segments. Maximum cl. 13 mm (Pl. 7h). Subtidal, shelf (8–100 m). Tropical W Atlantic. 2 species. Criales (1992) redescribed and illustrated Trachycaris rugosa (Bate, 1888), pointing out the sexual dimorphism, and distinguished the two species.

Saron Thallwitz, 1891 Diagnosis. Rostrum dentate dorsally and ventrally, with strong ventral blade or series of strong teeth. Branchiostegal or hepatic tooth present, pterygostomial spine present. Pleonite 6 pleuron with articulated plate at posteroventral angle. Mandibular palp of 3 articles. Pereopod 2 carpus of 10–17 segments. Pereopods 1–4 with arthrobranchs. Maximum cl. 12 mm. Intertidal–shelf (0–40 m, coral reefs). Indo-West Pacific. 4 species (Chace 1997: key to species). Saron marmoratus (Olivier, 1811) has been introduced to the Mediterranean Sea, possibly via the Suez Canal (Rothman et al. 2013). References Bruce AJ (1982) Thorella cobourgi, new genus, new species, a hippolytid shrimp from the Northern Territory, Australia. Journal of Crustacean Biology 2, 451–458. doi:10.2307/1548059 Bruce AJ (1990) Crustacea Decapoda: Gelastreutes crosnieri gen. nov., sp. nov. (Hippolytidae) from New Caledonia. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 6. Mémoires du Muséum National d’Histoire Naturelle, Paris 145, 137–147. Cardoso I, Fransen C (2012) A new species of the deepwater shrimp genus Leontocaris (Hippolytidae: Caridea) from the South Mid-Atlantic Ridge. Journal of the Marine Biological Association of the United Kingdom 92, 1083–1088. doi:10.1017/S0025315412000458 Chace FA (1997) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907–1910, Part 7: Families Atyidae, Eugonatonotidae, Rhynchocinetidae, Bathypalaemonellidae, Processidae, and Hippolytidae. Smithsonian Contributions to Zoology 587, 1–106. Chan T-Y, Van Den Beld IMJ, De Grave S (2016) A further record of the rare hippolytid shrimp Leontocaris lar Kemp, 1906 (Decapoda, Caridea) from the Celtic Sea, off north-western France. Crustaceana 89, 251–257. doi:10.1163/15685403-00003515 Christoffersen ML (1987) Phylogenetic relationships of hippolytid genera, with an assignment of new families for the Crangonoidea and Alpheoidea (Crustacea, Decapoda, Caridea). Cladistics 3, 348–362. doi:10.1111/j.1096-0031.1987.tb00898.x Criales MM (1992) Redescription of the hippolytid shrimp Trachycaris rugosa (Bate) (Crustacea: Decapoda: Caridea) from the Western Atlantic, with notes on sexual dimorphism. Proceedings of the Biological Society of Washington 105, 562–570. Crosnier A (2000) A new genus and species of hippolytid shrimp (Decapoda, Caridea) from Wallis Island, south-west Pacific. Journal of Crustacean Biology 20, 109–115. doi:10.1163/1937240X-90000013 Crosnier A, Forest J (1973) Les crevettes profondes de l’Atlantique oriental tropical. Faune Tropicale 19, 1–409. d’Udekem d’Acoz C (2007) New records of Atlantic Hippolyte, with the description of two new species, and a key to all Atlantic and Mediterranean species (Crustacea, Decapoda, Caridea). Zoosystema 29, 183–207.

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De Grave S, Li CP, Tsang LM, Chu KH, Chan T-Y (2014) Unweaving hippolytoid systematics (Crustacea, Decapoda, Hippolytidae): resurrection of several families. Zoologica Scripta 43, 496–507. doi:10.1111/ zsc.12067 Fransen CHJM (2001) Leontocaris vanderlandi, a new species of hippolytid shrimp (Crustacea: Decapoda: Caridea) from the Seychelles, with an analysis of phylogenetic relations within the genus. Zoölogische Verhandelingen 334, 57–76. Fransen CHJM, De Grave S (2019) Two new species of Hippolyte from the Tropical Central and East Atlantic (Crustacea, Decapoda, Caridea). Zootaxa 4550, 201–220. doi:10.11646/zootaxa.4550.2.3 Gan Z, Li X (2017) New record of the genus Phycocaris Kemp, 1916 (Decapoda: Caridea: Hippolytidae) from Hainan Island, China. Chinese Journal of Oceanology and Limnology 35, 664–667. doi:10.1007/​ s00343-017-6059-8 Gan ZB, Li XZ (2019) Recognizing two new Hippolyte species (Decapoda, Caridea, Hippolytidae) from the South China Sea based on integrative taxonomy. PeerJ 7, e6605. doi:10.7717/peerj.6605 Haley C, von der Meden C, Atkinson L, Reed C (2017) Habitat associations and distribution of the hyperbenthic shrimp, Nauticaris marionis, around the sub-Antarctic Prince Edward Islands. Deep Sea Research. Part I, Oceanographic Research Papers 127, 41–48. doi:10.1016/j.dsr.​ 2017.07.005 Holthuis LB (1952) Reports of the Lund University Chile Expedition 1948–49, 5. The Crustacea Decapoda Macrura of Chile. Lunds Universitets Årsskrift N.F. Avd. 2(47), 1–109. Holthuis LB (1993) The Recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda) with an appendix on the Order Amphionidacea. revised edn. Nationaal Natuuristorisch Museum, Leiden. Kensley B, Tranter HA, Griffin DJG (1987) Deepwater decapod Crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39, 263–331. doi:10.3853/j.0067-1975.39.1987.171 Komai T, Chan T-Y (2010) Two new pandalid shrimps and the discovery of the second specimen of the rare hippolytid shrimp Leontocaris bulga Taylor & Poore, 1998 (Crustacea, Decapoda) from the Mozambique MAINBAZA Cruise. Zoosystema 32, 625–641. doi:10.5252/z2010n4a6 Komai T, Takeuchi I, Takeda M (1996) Deep-sea shrimp (Crustacea: Decapoda: Caridea) from the Antarctic Sea collected during the JARE-35 cruise. Proceedings of the National Institute of Polar Research, Tokyo, Symposium on Polar Biology 9, 179–206. Ledoyer M (1990) Mysidacés et caridés benthiques (Crustacea) de la Campagne MD 50/Jasus aux Iles Saint-Paul et Amsterdam (Océan Indien). Mésogée 50, 45–51. Pakhomov EA, Kuun P, McQuaid CD (2000) Biology of the bottomdwelling shrimp Nauticaris marionis Bate, 1888 at the sub-Antarctic Prince Edward Islands. Polar Biology 23, 522–530. doi:10.1007/ s003000000116 Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Poore GCB (2009) Leontocaris alexander, a new deepwater hippotylid shrimp from Tasmanian seamounts with a phylogeny of the genus (Crustacea: Caridea). Crustaceana 82, 951–967. doi:10.1163/156854009X427478 Rothman SBS, Shlagman A, Galil BS (2013) Saron marmoratus, an IndoPacific marble shrimp (Hippolytidae: Decapoda: Crustacea) in the Mediterranean Sea. Marine Biodiversity Records 6, e129. doi:10.1017/ S1755267213000997 Saponari L, Galli P, Montano S (2018) First record of Phycocaris simulans (Decapoda; Hippolytidae) from the Republic of Maldives. Galaxea, Journal of Coral Reef Studies 20, 11–12. doi:10.3755/galaxea.20.1_11

Smaldon G, Holthuis LB, Fransen CHJM (1993) Synopses of the British Fauna (New Series) No. 15. Coastal shrimps and prawns. Keys and notes for identification of the species (Second Edition). In: Barnes RSK, Crothers J, Field Studies Council, Shrewsbury. Taylor J, Poore GCB (1998) A review of the genus Leontocaris (Crustacea: Caridea: Hippolytidae) with descriptions of three species from southeastern Australian seamounts and slope. Memoirs of the Museum of Victoria 57, 57–69. doi:10.24199/j.mmv.1998.57.03 Terossi M, De Grave S, Mantelatto FL (2017) Global biogeography, cryptic species and systematic issues in the shrimp genus Hippolyte Leach, 1814 (Decapoda: Caridea: Hippolytidae) by multimarker analyses. Scientific Reports 7, 6697. doi:10.1038/s41598-017-06756-1 Terossi M, Almeida AO, Mantelatto FL (2019) Morphology and DNA data reveal a new shrimp species of genus Latreutes Stimpson, 1860 (Decapoda: Hippolytidae) from the Western Atlantic. Zoological Science 36, 440–447. doi:10.2108/zs190016 Wicksten MK, Hancock ZB (2021) Synonymy of the caridean shrimp genus Paralatreutes Kemp, 1925 with Latreutes Stimpson, 1860 (Decapoda: Hippolytidae). Proceedings of the California Academy of Sciences 67, 247–252.

Lysmatidae Dana, 1852 Figure 6.10, Plate 7m Lysmatidae comprise four genera with one or few species plus the much better known and more speciose Lysmata. Species of Exhippolysmata and Lysmata are unusual in that they are protandrous simultaneous hermaphrodites, that is, they mature and reproduce initially as males, and then later in life become functional simultaneous hermaphrodites (Baeza 2010; Bauer 2006). As with other alpheoid families supported by molecular phylogenies, Lysmatidae are difficult to diagnose morphologically (De Grave et  al. 2014). Christoffersen (1987) included a suite of four genera different from those included here, linked by two synapomorphies: the antennular lateral flagellum is biramous; and the infraorbital angle is depressed and inconspicuous, lying below the antennal tooth. However, the antennule is more variable than Christoffersen (1987) stated. The antennular lateral flagellum is usually biramous (Fig.  6.10f) but in some species of Lysmata a thicker proximal section ends in a short spur (Fig.  6.10g) (Fiedler et  al. 2010) or swelling (Fig.  6.10h) in Mimocaris heterocarpoides Nobili, 1903 (Nguyen 2000). Maxilliped 1 has a particularly long exopod (Fig. 6.10i); maxilliped 3 has an elongate last article (Fig.  6.10j) unlike Barbouridae. De Grave et  al. (2014) noted that the larval stages can have elongate appendages with flattened areas – in Lysmata pereopod 5 of the zoeae is long with a paddle-like propodus. They also suggested that the arrangement of seta of the carpo-propodal brush may be diagnostic (see De Grave and Goulding 2011: Fig. 8E, F). Diagnosis. Rostrum prominent, dentate dorsally and ventrally (usually). Antennular lateral flagellum distally

6 – Caridea – shrimps

biramous, or with thicker proximal section ending in short spur or swelling (Fig.  6.10f–h). Pereopods 1, 2 dissimilar, each pair symmetrical, first more robust than second. Pereopod 1 with carpo-propodal brush comprising 2

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elongate fields of serrate setae on carpus. Pereopod 2 carpus of 12–29 segments. Implicit generic attributes. Carapace smooth. Pleonites 2–5 without posteromedian spine. Arthrobranchs absent.

Key to genera of Lysmatidae 1. – 2. – 3. – 4. –

Rostrum longer than carapace, with dorsal basal crest of closely spaced teeth (Fig. 6.10a, e)���������������������������������������������2 Rostrum as long or shorter than carapace, with regularly spaced dorsal teeth, usually with ventral teeth (Fig. 6.10b–d)�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Carapace with longitudinal carinae. Pleonites 2–5 each with sharp posteromedian spine, pleura ending in 1 or 2 sharp points (Fig. 6.10e)���������������������������������������������������������������������������������������������������������������������������������Mimocaris Carapace without longitudinal carinae. Pleonites 2–5 smooth, pleura rounded�����������������������������������������Exhippolysmata Pereopods 1–4 with arthrobranchs. Pleonite 3 with posteromedian tooth�������������������������������������������������������������������� Ligur Pereopods 1–4 without arthrobranchs. Pleonites without posteromedian tooth������������������������������������������������������������������4 Pereopods 1–4 with epipods����������������������������������������������������������������������������������������������������������������������������������������������Lysmata Pereopods without epipods��������������������������������������������������������������������������������������������������������������������������������������������Lysmatella Exhippolysmata Stebbing, 1915

Diagnosis. Rostrum longer than carapace, slender, straight, with 2 well spaced teeth, 10–12 on basal crest (4 or 5 postorbital), 8 ventral teeth. Maximum cl. 17 mm. Subtidal, shelf (1–48 m), marine, occasionally in fresh water. Tropical Atlantic, Western and Central Indo-Pacific, Temperate Southern Africa. 4 species. Species of Exhippolysmata in SE Asia and in Brazil (Herrera et al. 2017) can be extremely abundant in shallow water and at least in Asia are fished for human consumption (Deshmukh 1990).

Ligur Sarato, 1885 Diagnosis. Rostrum almost as long as carapace, with 3 or 4 dorsal (1 postorbital), 4 or 5 ventral teeth. Pleonite 3 with posteromedian tooth, pleuron 5 sharp. Epipods and arthrobranchs present on pereopods 1–4. Maximum cl. 15 mm.

Slope (300–871 m). Temperate Northern and Tropical Atlantic, Western Indo-Pacific. 1 species (Lemaitre 1984).

Lysmatella Borradaile, 1915 Diagnosis. Rostrum about as long as carapace, with 6–9 regularly spaced dorsal teeth (4 postorbital), 5–7 ventral teeth. Epipods absent. Maximum cl. 5 mm. Subtidal, shelf (2–70 m). Western and Central Indo-Pacific. 1 species (Pillai et al. 2019; Xu and Li 2015).

Lysmata Risso, 1816 Diagnosis. Rostrum usually shorter than carapace, with dorsal and usually ventral teeth. Epipods present on pereopods 1–4. Maximum cl. 30 mm (Pl. 7m).

Fig. 6.10.  Lysmatidae. Anterior carapace, rostrum: a, Exhippolysmata ensirostris (Kemp, 1914); b, Ligur ensiferus (Risso, 1816); c, Lysmata seticaudata (Risso, 1816); d, Lysmatella prima Borradaile, 1915; e, Mimocaris heterocarpoides Nobili, 1903. Antennular flagella (lateral flagellum on left): f, Lysmata chiltoni Kemp, 1914; g, Lysmata hochi Baeza & Anker, 2008; h, Mimocaris heterocarpoides Nobili, 1903. Maxilliped 1: i, Lysmata. Maxilliped 3: j, Lysmata.

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Intertidal–slope (0–360 m). Cosmopolitan, except in polar seas. 50 species. Peppermint shrimps, a name applied to several species of Lysmata, are popular with aquarists who appreciate them for their bright colours. Other species have attracted a variety of other common names, such as Cleaner or Skunk shrimp, applied to both L. amboinensis (De Man, 1888) and L. grabhami (Gordon, 1935) and Blood shrimp, applied to L. debelius Bruce, 1983 and L. splendida Burukovsky, 2000. Species of Lysmata display a variety of lifestyles in a range of habitats, some living in aggregations, others solitary (Baeza 2010; Bauer 2004, 2006). The genus has been divided into species groups based on the development of the accessory branch of the lateral antennular flagellum, a division supported by molecular sequences (Anker and Cox 2011; Fiedler et al. 2010). Chace (1997) provided a key to the 25 species then known. Twice as many are now known, listed by oceanic province by Anker and Cox (2011). Keys to species are available for the south-western Atlantic Ocean (Giraldes et  al. 2018), the eastern Pacific (Wicksten 2000) and seas of China (Wang and Sha 2018). De Grave and Anker (2018) noted the importance of colour patterns in distinguishing species.

Mimocaris Nobili, 1903 Diagnosis. Carapace with longitudinal carinae. Rostrum longer than carapace, slender, upturned, with 6 or 7 teeth, 10–12 on basal crest (4 or 5 postorbital), 16 ventral teeth. Pleonites 2–5 each with sharp posteromedian spine, pleura ending in 1 or 2 sharp points. Maximum cl. 13 mm. Subtidal, estuarine (4–22 mm). Central Indo-Pacific. 1 species (Nguyen 2000). References Anker A, Cox D (2011) A new species of the shrimp genus Lysmata Risso, 1816 (Crustacea, Decapoda) from Guam. Micronesica 41, 197–214. Baeza JA (2010) Molecular systematics of peppermint and cleaner shrimps: phylogeny and taxonomy of the genera Lysmata and Exhippolysmata (Crustacea: Caridea: Hippolytidae). Zoological Journal of the Linnean Society 160, 254–265. doi:10.1111/j.1096-3642.2009.00605.x Bauer RT (2004) Remarkable shrimps. Adaptations and natural history of the caridean shrimps. University of Oklahoma Press, Norman. Bauer RT (2006) Same sexual system but variable sociobiology: evolution of protandric simultaneous hermaphroditism in Lysmata shrimps. Integrative and Comparative Biology 46, 430–438. doi:10.1093/icb/icj036 Chace FA (1997) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907–1910, Part 7: Families Atyidae, Eugonatonotidae, Rhynchocinetidae, Bathypalaemonellidae, Processidae, and Hippolytidae. Smithsonian Contributions to Zoology 587, 1–106. Christoffersen ML (1987) Phylogenetic relationships of hippolytid genera, with an assignment of new families for the Crangonoidea and Alpheoidea (Crustacea, Decapoda, Caridea). Cladistics 3, 348–362. doi:10.1111/j.1096-0031.1987.tb00898.x De Grave S, Anker A (2018) A new, distinctly coloured species of Lysmata Risso, 1816 (Malacostraca: Decapoda: Lysmatidae) from the south-central Atlantic. Zootaxa 4429, 390–400. doi:10.11646/zootaxa.4429.2.13 De Grave S, Goulding LYD (2011) Comparative morphology of the pereiopod 1 carpo-propodal (P1-CP) antennal flagellar grooming brush in caridean shrimps (Crustacea, Decapoda). Zoologischer Anzeiger 250, 280–301. doi:10.1016/j.jcz.2011.08.003

De Grave S, Li CP, Tsang LM, Chu KH, Chan T-Y (2014) Unweaving hippolytoid systematics (Crustacea, Decapoda, Hippolytidae): resurrection of several families. Zoologica Scripta 43, 496–507. doi:10.1111/ zsc.12067 Deshmukh VD (1990) Population dynamics and stock assessment of hermaphrodite prawn Exhippolysmata ensirostris Kemp, at Nawabunder, Gujarat. Journal of the Marine Biological Association of India 32, 187–192. Fiedler GC, Rhyne A, Segawa R, Aotsuka T, Schizas N (2010) The evolution of euhermaphroditism in caridean shrimps: A molecular perspective of sexual systems and systematics. BMC Evolutionary Biology 10, 297. doi:10.1186/1471-2148-10-297 Giraldes BW, Macedo TP, Brandão MC, Baeza JA, Freire AS (2018) Lysmata arvoredensis nov. sp. a new species of shrimp from the south coast of Brazil with a key to species of Lysmata (Caridea: Lysmatidae) recorded in the southwestern Atlantic. PeerJ 6, e5561. doi:10.7717/peerj.5561 Herrera DR, Pescinelli RA, Costa RCd (2017) Ecological distribution of Nematopalaemon schmitti and Exhippolysmata oplophoroides (Crustacea: Caridea) near an upwelling area off southeastern Brazil. Latin American Journal of Aquatic Research 45, 149–158. doi:10.3856/ vol45-issue1-fulltext-14 Lemaitre R (1984) Decapod crustaceans from Cay Sal Bank, Bahamas, with notes on their zoogeographic affinities. Journal of Crustacean Biology 4, 425–447. doi:10.2307/1548042 Nguyen VX (2000) Redescription of Mimocaris heterocarpoides Nobili, 1903 (Decapoda, Caridea, Hippolytidae) with remarks on Its occurrence in Vietnam. Crustaceana 73, 857–868. doi:10.1163/156854000504868 Pillai SL, Maheswarudu G, Thomas J, Ragesh L, Sreesanth M, et al. (2019) New record of Lysmatella prima Borradaile, 1915 (Crustacea, Caridea, Lysmatidae) from the Arabian Sea, southwestern India. Zootaxa 4571, 281–284. doi:10.11646/zootaxa.4571.2.10 Wang Y-R, Sha Z-L (2018) Description of two new species of Lysmata Risso, 1816 (Decapoda, Lysmatidae) from the China seas, with remarks on Lysmata vittata (Stimpson 1860). Zootaxa 4392, 28–40. doi:10.11646/zootaxa.4392.1.2 Wicksten MK (2000) The species of Lysmata (Caridea: Hippolytidae) from the eastern Pacific Ocean. Amphipacifica 2, 3–22. Xu P, Li XZ (2015) Report on the Hippolytidae Bate (sensu lato) from China seas. Zoological Systematics 40, 107–165.

Merguiidae Christoffersen, 1987 Figure 6.11, Plate 7i Merguiids are semiterrestrial shrimps from estuarine boulder and mangrove habitats. They have developed the ability to climb up mangrove prop roots out of the water. Merguia is the only genus, with two species, one on the tropical coast of the western Atlantic (Abele 1970) and the other in the Indo-West Pacific, from Africa (Gillikin et al. 2001) to Japan (Komai 2002). Diagnosis. Rostrum little longer than eyestalk, with 2–3 dorsal teeth, unarmed ventrally (Fig. 6.11a). Mandible without palp or incisor. Pereopods 1, 2 dissimilar, each pair symmetrical, first more robust than second. Pereopod 2 carpus multiarticulate, with 22–24 segments. Epipod present on maxilliped 2 only. Uropodal exopod article 1 posterolaterally produced alongside distal article (Fig. 6.11b).

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New Zealand (Yaldwyn 1971), Japan (Hayashi 1991) Australia (Poore 2004), India and the Persian Gulf (De Grave et al. 2020). Diagnosis. Rostrum small, obsolete, or absent. Eyestalks elongate, reaching nearly to distal end of antennular peduncle. Pereopods 1, 2 dissimilar, each pair symmetrical, first more robust than second. Pereopod 2 carpus of 4 or 5 segments. Ogyrides Stebbing, 1914

Fig. 6.11.  Merguiidae. a, Merguia oligodon (De Man, 1888); b, telson, uropod.

Merguia Kemp, 1914 Maximum cl. 7 mm (Pl. 7i). Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific. 2 species. References Abele LG (1970) Semi-terrestrial shrimp (Merguia rhizophorae). Nature 226, 661–662. doi:10.1038/226661a0 Gillikin DP, De Grave S, Tack JF (2001) The occurrence of the semi-terrestrial shrimp Merguia oligodon (De Man, 1888) in Neosarmatium smithii H. Milne Edwards, 1853 burrows in Kenyan mangroves. Crustaceana 74, 505–507. doi:10.1163/156854001750243081 Komai T (2002) New record of a semi-terrestrial hippolytid shrimp, Merguia oligodon (De Man) (Crustacea: Decapoda: Caridea) from Japan and Thailand. Natural History Research 7, 75–82.

Ogyrididae Holthuis, 1955 Figure 6.12, Plate 7j The elongate eyestalks immediately enable recognition of species of Ogyrides, the only genus. Ogyrides is distributed in tropical to temperate waters worldwide. Ayón-Parente and Salgado-Barragán (2013) gave a key to the three Eastern Pacific species but the genus has never been reviewed as a whole. In the Indo-West Pacific, species have been recorded in Indonesia (De Man 1902), South Africa (Barnard 1950),

Fig. 6.12.  Ogyrididae. Ogyrides delli Yaldwyn, 1971.

Maximum cl. 6 mm (Pl. 7j). Estuarine and marine, intertidal– shelf (0–55 m). Temperate Northern and Tropical Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate Southern Africa, Temperate Australasia. 13 species. References Ayón-Parente M, Salgado-Barragán J (2013) A new species of the caridean shrimp genus Ogyrides Stebbing, 1914 (Decapoda: Ogyrididae) from the eastern tropical Pacific. Zootaxa 3683, 589–594. doi:10.11646/ zootaxa.3683.5.7 Barnard KH (1950) Descriptive catalogue of South African decapod Crustacea (crabs and shrimps). Annals of the South African Museum 38, 1–837. De Grave S, Al-Kandari M, Anker A (2020) A new species of Ogyrides (Decapoda, Caridea, Ogyrididae) from Kuwait. Crustaceana 93, 225–235. doi:10.1163/15685403-bja10007 De Man JG (1902) The Decapoda of the Siboga-Expedition. Part 5. On a collection of macrurous decapod Crustacea of the Siboga Expedition, chiefly Penaeidae and Alpheidae. Siboga-Expéditie 39a4, 1–51, pls 1–4. Hayashi KI (1991) Prawns, shrimps and lobsters from Japan (62). Family Ogyrididae – Genus Ogyrides. Kaiyo To Seibutsu 13, 430–433. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Yaldwyn JC (1971) Preliminary descriptions of a new genus and twelve new species of natant decapod Crustacea from New Zealand. Records of the Dominion Museum 7, 85–94.

Thoridae Kingsley, 1879 Figure 6.13, Plate 7k The eight genera of Thoridae were until recently included in Hippolytidae. In the widely used keys to hippolytid genera (Chace 1997; Holthuis 1993) these eight are scattered between many others, suggesting that any morphological similarities were not obvious to these eminent carcinologists. The grouping of these genera in Thoridae is largely based on similarities demonstrated during analysis of nuclear and mitochondrial genes (De Grave et  al. 2014). These authors were unable to clearly define morphological synapomorphies of the family but included the club-like dorsal antennular flagellum and a protuberance or distolateral tooth on the second article of the antennular peduncle as possibilities. Genetic similarity of two thorid genera and polyphyly of

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Hippolytidae s.l. has since been confirmed (Aznar-Cormano et al. 2015). Holthuis’s (1993) key is the basis of this one. Diagnosis. Rostrum at least long twice as long as eyestalk, sometimes prominent, dentate dorsally and sometimes ventrally (Fig.  6.13a–j). Antennular lateral flagellum usually short, stout, with proximal articles annulated, bearing

many closely-spaced setae, appearing brush-like (Fig. 6.13k, l). Pereopods 1, 2 dissimilar, each pair symmetrical, first more robust than second. Pereopod 2 carpus of 6 or 7 segments (Fig. 6.13p). Implicit generic attributes. Postrostral teeth present. Mandible with palp.

Key to genera of Thoridae 1. – 2. – 3. – 4. –

Antennular peduncle article 3 with movable subtriangular distodorsal scale (Fig. 6.13l). Mandible without palp (Fig. 6.13a)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Antennular peduncle article 3 without movable distodorsal scale (may be produced distally; Fig. 6.13k). Mandible with palp (Fig. 6.13m, n)������������������������������������������������������������������������������������������������������������������������������������������3 Rostrum acute, with several (rarely 1) dorsal teeth in adults (Fig. 6.13j). Telson with 2–4 (usually 3) pairs of posterior spiniform setae (Fig. 6.13s)��������������������������������������������������������������������������������������������������������������������������������Thor Rostrum slender, acute, not exceeding antennular peduncle article, with 1 dorsal tooth (Fig. 6.13i). Telson with 3 pairs of posterior spiniform setae (Fig. 6.13r)������������������������������������������������������������������������������������������������ Thinora Carapace without supraorbital spine (Fig. 6.13c, d)��������������������������������������������������������������������������������������������������������������������4 Carapace with 1 or more supraorbital spines (Fig. 6.13a, e–h)��������������������������������������������������������������������������������������������������5 Maxilliped 3 with exopod�����������������������������������������������������������������������������������������������������������������������������������������������������Eualus Maxilliped 3 without exopod������������������������������������������������������������������������������������������������������������������������������������ Heptacarpus

Fig. 6.13.  Thoridae. Habitus: a, b, Birulia kishinouyei (Yokoya, 1930). Anterior carapace: c, Eualus; d, Heptacarpus; e, Lebbeus elegans Komai, Hayashi & Kohtsuka, 2004; f, Lebbeus clarehannah McCallum & Poore, 2010: g, Paralebbeus; h, Spirontocaris; i, Thinora; j, Thor (with antennule peduncle). Anterior carapace, eyes, antennules, scaphocerites: k, Eualus ctenomerus Komai & Fujiwara, 2012. Antennule: l, Thor. Mandible: m, n, Eualus spp.; o, Thinora. Pereopod 2: p, Thor. Pereopod 5: q, Lebbeus. Telson apex: r, Thor; s, Thinora.

6 – Caridea – shrimps

5. – 6. – 7. –

93

Carapace with 2 or more pairs of supraorbital spines (Fig. 6.13h). Maxilliped 3 with exopod���������������������� Spirontocaris Carapace with 1 pair of supraorbital spines (Fig. 6.13a, e–g). Maxilliped 3 without exopod����������������������������������������������6 Pleonites 1 and 5 with 2 dorsal carinae, pleonites 2–4 with dorsal carina. Carapace without antennal and pterygostomial spines, with large branchiostegal spine (Fig. 6.13a, b)�������������������������������������������������������������������� Birulia Pleonites dorsally rounded. Carapace with antennal and pterygostomial spines, without branchiostegal spine (Fig. 6.13e–g)���������������������������������������������������������������������������������������������������������������������������������������������������������������������7 Carapace with postrostral carapace spines (Fig. 6.13e, f). Pereopods 3–5 meri with lateral spiniform setae (Fig. 6.13q)����������������������������������������������������������������������������������������������������������������������������������������������������������������������� Lebbeus Carapace without postrostral carapace spines (Fig. 6.13g). Pereopods 3–5 meri with or without lateral spiniform setae�������������������������������������������������������������������������������������������������������������������������������������������������������� Paralebbeus Birulia Bražnikov, 1903

Diagnosis. Rostrum blade-like, directed obliquely downwards, with few small dorsal teeth. Carapace with 1 pair of supraorbital spines, without antennal spine, with prominent sharp branchiostegal spine. Pleonites 1 and 5 with 2 dorsal carinae, pleonites 2–4 with single dorsal carina. Maxilliped 3 without exopod. Pereopods 3–5 meri with lateral spines. Maximum cl. 20 mm. Subtidal, shelf (15–118 m). Temperate Northern W Pacific. 2 species. Apparently quite rare, only one species has been illustrated (Yokoya 1930).

Eualus Thallwitz, 1891 Diagnosis. Rostrum blade-like, variously dorsally and ventrally dentate. Carapace without supraorbital spines, without antennal spine, with minute branchiostegal spine. Maxilliped 3 with exopod. Pereopods 3–5 meri with movable lateral spines. Maximum cl. 10 mm (many species much smaller) (Pl. 7k). Intertidal–bathyal (0–2401 m). Arctic, Temperate Northern Atlantic, Temperate Northern Pacific, Temperate Southern Africa, Temperate Australasia, Southern Ocean. 41 species. Despite there being many species, none is tropical. The genus has not been reviewed as a whole. Komai and Fujiwara (2012) recognised a species group characterised by epipods on pereopods 1–3. The genus and Lebbeus are among the few caridean genera known from Antarctic (Nye et al. 2013) and Arctic (Greve 1963) waters.

Heptacarpus Holmes, 1900 Diagnosis. Rostrum long, slender, with dorsal and ventral teeth. Carapace without supraorbital spines, without antennal spine, without branchiostegal spine or with minute branchiostegal spine. Maxilliped 3 without exopod. Pereopods 3–5 meri with movable lateral spines. Maximum cl. 10 mm (plus long rostrum). Intertidal–slope (0–1125 m). Temperate Northern Pacific. 33 species. Some species are found in seagrass and algal habitats. The only keys to species deal only with Asian species (Hayashi 1979; Komai and Ivanov 2008) but others occur as far south as California (Wicksten 1986).

Lebbeus White, 1847 Diagnosis. Rostrum blade-like or spiniform, variously dorsally and ventrally dentate, dorsal spines extending on to carapace.

Carapace with 1 pair of supraorbital spines, with antennal spine, with prominent sharp branchiostegal spine. Maxilliped 3 without exopod. Pereopods 3–5 meri with lateral spines. Maximum cl. 16 mm. Intertidal–bathyal (0–2640 m depth). Cosmopolitan including Arctic and Southern Ocean, some species from hydrothermal vents. 72 species. Lebbeus is the largest and most widespread of the ‘hippolytoid’ genera. Approximately half of the described species are from the north-western Pacific, separated in the key by Hayashi (1992). Species are known from Antarctica (Schiaparelli et al. 2015), eight species from cold seeps and hydrothermal vents (Ahyong 2010; Chan and Komai 2017; Nye et  al. 2013). Species restricted to depths greater than 1500 m were listed by Komai (2013a). Some are spectacularly coloured with high dorsal crests (McCallum and Poore 2010). Variation in the development of epipods on pereopods 1−3 has commonly been used to diagnose and group species (Komai et al. 2004).

Paralebbeus Bruce & Chace, 1986 Diagnosis. Rostrum weak, acute, not reaching distal margin of antennular peduncle article 1, without or with 1–3 dorsal and 1 or 2 ventral spines; postrostral teeth absent. Carapace with 1 pair of supraorbital spines, with antennal spine, with minute branchiostegal spine. Maxilliped 3 without exopod. Pereopods 3–5 meri with lateral spines (one species), or without lateral spines. Maximum cl. 15.3 mm. Slope (452–1800 m). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Australasia (associated with hexactinellid sponges). 5 species (Ahyong 2019; Komai 2013b).

Spirontocaris Bate, 1888 Diagnosis. Rostrum blade-like, variously dorsally and ventrally dentate. Carapace with 2 or more pairs of supraorbital spines, with antennal spine, with minute branchiostegal spine. Maxilliped 3 with exopod. Maximum cl. 25 mm. Intertidal–slope (0–1380 m). Arctic, Temperate Northern Atlantic, Temperate Northern Pacific. 21 species. Species of Spirontocaris in shallow water are often with algae. Rathbun (1902) described 20 species from the NW Pacific but only six are accepted now (Hendrickx and Ayón Parente 2017). Three species from the Kurile Islands were reviewed by Kobjakova (1962) and three from Norway by Greve (1963).

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Thinora Bruce, 1998 Diagnosis. Rostrum slender, acute, not exceeding antennular peduncle article, with 1 dorsal spine. Carapace with 1 pair of supraorbital spines, with antennal spine, without branchiostegal spine. Telson with 3 pairs of posterior spiniform setae. Antennular peduncle article 3 with movable subtriangular dorsal scale. Mandible without palp. Maxilliped 3 with exopod. Pereopods 3–5 meri without lateral spines. Pereopod 1 sexually dimorphic, appendix masculina absent. Maximum cl. 3.3 mm. Intertidal, subtidal (0–5 m). Western and Central Indo-Pacific. 1 species (Bruce 1997 as Thorina, preoccupied name).

Thor Kingsley, 1878 Diagnosis. Rostrum acute, with several dorsal spines in adults. Carapace with or without pair of supraorbital spines, with antennal spine, without branchiostegal spine. Telson with 2–4 pairs of posterior spiniform setae. Antennular peduncle article 3 with movable subtriangular dorsal scale. Mandible without palp. Maxilliped 3 with exopod. Pereopods 3–5 meri without movable lateral spines. Pereopod 1 not sexually dimorphic. Maximum cl. 3.5 mm. Intertidal–shelf (0–58 m; some associated with sea anemones or other invertebrates). Temperate Northern and Tropical W Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific. 16 species (Bruce 1976: key to some Indo-West Pacific species; Chace 1972, 1997: keys to species). References Ahyong ST (2010) New species and new records of Caridea (Hippolytidae: Pasiphaeidae) from New Zealand. Zootaxa 2372, 341–357. doi:10.11646/​ zootaxa.2372.1.26 Ahyong ST (2019) Discovery of the sponge-associated caridean genus Paralebbeus in New Zealand and southeastern Australia: P. pegasus sp. nov. (Crustacea: Decapoda: Thoridae). Zootaxa 4652, 375–383. doi:10.11646/zootaxa.4652.2.10 Aznar-Cormano L, Brisset J, Chan T-Y, Corbari L, Puillandre N, et  al. (2015) An improved taxonomic sampling is a necessary but not sufficient condition for resolving inter-families relationships in Caridean decapods. Genetica 143, 195–205. doi:10.1007/s10709-014-9807-0 Bruce AJ (1976) A report on a small collection of shrimps from the Kenya National Marine Parks at Malindi, with notes on selected species. Zoölogische Verhandelingen 145, 3–72. Bruce AJ (1997) A new genus of hippolytid shrimp (Crustacea: Decapoda: Hippolytidae) for Thor maldivensis Borradaile. Memoirs of the Queensland Museum 42, 15–23. Chace FA (1972) The shrimps of the Smithsonian-Bredin Caribbean Expeditions with a summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology 98, 1–179. doi:10.5479/si.00810282.98 Chace FA (1997) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907–1910, Part 7: Families Atyidae, Eugonatonotidae, Rhynchocinetidae, Bathypalaemonellidae, Processidae, and Hippolytidae. Smithsonian Contributions to Zoology 587, 1–106. Chan T-Y, Komai T (2017) A new shrimp species of the genus Lebbeus White, 1847 (Crustacea: Deacpoda: Caridea: Thoridae) from a deep-sea cold seep site off southwestern Taiwan. Zootaxa 4238, 426–432. doi:10.11646/zootaxa.4238.3.9

De Grave S, Li CP, Tsang LM, Chu KH, Chan T-Y (2014) Unweaving hippolytoid systematics (Crustacea, Decapoda, Hippolytidae): resurrection of several families. Zoologica Scripta 43, 496–507. doi:10.1111/ zsc.12067 Greve L (1963) The genera Spirontocaris, Lebbeus, Eualus and Thoralus in Norwegian waters (crust. dec.). Sarsia 11, 29–42. doi:10.1080/0036482 7.1963.10410283 Hayashi KI (1979) Studies on hippolytid shrimps from Japan  – VII. The genus Heptacarpus Holmes. Suisan Daigakko Kenkyu Hokoku 28, 11–32. Hayashi KI (1992) Prawns, shrimps and lobsters from Japan (67). Family Hippolytidae  – Genera Birulia and Lebbeus. Kaiyo To Seibutsu 82, 341–345[in Japanese]. Hendrickx ME, Ayón Parente M (2017) The genus Spirontocaris Spence Bate, 1888 (Caridea, Decapoda, Thoridae) in western Mexico. Zootaxa 4320, 305–320. doi:10.11646/zootaxa.4320.2.6 Holthuis LB (1993) The Recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda) with an appendix on the Order Amphionidacea. revised edn. Nationaal Natuuristorisch Museum, Leiden. Kobjakova ZI (1962) Notes on rare and new species of decapod crustaceans (Decapoda, Malacostraca) from the region of Kuril Islands. Issledovaniya Dalnevostochnykh Morei SSSR 8, 243–247[in Russian]. Komai T (2013a) A new secies of the hippolytid shrimp genus Lebbeus (Crustacea: Decapoda: Caridea) from lower bathyal zone in the Izu Islands, Central Japan. Natural History Research 12, 81–89. Komai T (2013b) A new species of the hippolytid genus Paralebbeus Bruce & Chace, 1986 (Crustacea: Decapoda: Caridea) from the Coral Seamount, southwestern Indian Ocean. Zootaxa 3646, 171–179. doi:10.11646/zootaxa.3646.2.5 Komai T, Fujiwara Y (2012) Description of a new species of the hippolytid shrimp genus Eualus Thallwitz, 1892 from Japan, and clarification of the status of E. kikuchii Miyake & Hayashi, 1967 (Crustacea: Decapoda: Caridea). Zootaxa 3546, 68–80. doi:10.11646/zootaxa.3546.1.5 Komai T, Ivanov BG (2008) Identities of three taxa of the hippolytid shrimp genus Heptacarpus (Crustacea: Decapoda: Caridea), with description of a new species from East Asian waters. Zootaxa 1684, 1–34. doi:10.11646/zootaxa.1684.1.1 Komai T, Hayashi K-I, Kohtsuka H (2004) Two new species of the shrimp genus Lebbeus White from the Sea of Japan, with redescription of Lebbeus kuboi Hayashi (Decapoda: Caridea: Hippolytidae). Crustacean Research 33, 103–125. doi:10.18353/crustacea.33.0_103 McCallum AW, Poore GCB (2010) Two crested and colourful new species of Lebbeus (Crustacea: Caridea: Hippolytidae) from the continental margin of Western Australia. In: De Grave S, Fransen CHJM (eds) Contributions to shrimp taxonomy. Zootaxa 2372, 126–137. doi:10.11646/zootaxa.2372.1.13 Nye V, Copley JT, Linse K (2013) A new species of Eualus Thallwitz, 1892 and new record of Lebbeus antarcticus (Hale, 1941) (Crustacea: Decapoda: Caridea: Hippolytidae) from the Scotia Sea. Deep Sea Research. Part II, Topical Studies in Oceanography 92, 145–156. doi:10.1016/j.dsr2.2013.01.022 Rathbun MJ (1902) Description of new decapod crustaceans from the west coast of North America. Proceedings of the United States National Museum 24, 885–905. doi:10.5479/si.00963801.1272.885 Schiaparelli S, Ahyong ST, Bowden D (2015) Evidence of niche conservatism and host fidelity in the polar shrimp Lebbeus kiae n. sp. (Decapoda: Caridea: Thoridae) from the Ross Sea, Antarctica. Hydrobiologia 761, 45–69. doi:10.1007/s10750-015-2403-1 Wicksten MK (1986) A new species of Heptacarpus from California, with a redescription of Heptacarpus palpator (Owen) (Caridea: Hippolytidae. Bulletin of the Southern California Academy of Sciences 85, 46–55. Yokoya Y (1930) Report on the Biological Survey of Mutsu Bay. 16. Macrura of Mutsu Bay. Science Reports of the Tohoku University 5, 525–548.

6 – Caridea – shrimps

Superfamily Atyoidea De Haan, 1849 Atyoidea comprise a single family of mostly freshwater shrimps. Dense tufts of setae at the ends of the fingers of the first two pairs of pereopods are characteristic, but maxilliped 2 is also unique. Diagnosis. Mandible incisor and molar separated by sinuous, setose margin, without palp. Maxilliped 2, last article broad, crescent-shaped or subsemicircular, attached by curved margin to slender, sickle-shaped extension of penultimate article (Fig. 6.14c). Pereopods 1, 2 similar, with dense tufts of setae at ends of fingers (Fig. 6.14b).

Atyidae De Haan, 1849 Figure 6.14 Atyidae comprises 46 genera, most of which are from freshwater streams and standing water. Refer to Holthuis (1993) for a key to genera of the family. Four genera contain species tolerant of saline environments. Antecaridina lauensis (Edmondson, 1954) is the only species in its genus. It is a bright red anchialine species widely distributed in the Indo-Pacific, found in Indian Ocean islands, Red Sea, Hawaii, southern Japan, Solomon Islands, the Philippines and Fiji (Leberer and Cai 2003; Yoshihisa et al. 2015). Halocaridina rubra Holthuis, 1963 and H. palahemo Kensley & Williams, 1986 are the only two species of this genus. Both occur in anchialine environments such as lava tubes in Hawaii (Havird et al. 2015; Kensley and Williams 1986) The genus Typhlatya Creaser, 1936 comprises 17 species of small shrimps of less than 20 mm in total length, all troglobitic, inhabiting caves and cenotes in the Caribbean, Europe, and the Galapagos and Ascension Islands (Alvarez et  al. 2005). Some are restricted to freshwater systems but others tolerate saline conditions, such as anchialine caves in France (Jaume and Bréhier 2005).

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Paratya Miers, 1882 comprises 22 species from freshwater habitats in the western Pacific, with a disjunct northern range in the North Pacific (Satake and Cai 2005) and South Pacific (Page et al. 2007). Some are amphidromous (Page et al. 2005). The Paratya australiensis Kemp, 1917 species complex occurs in streams and lakes in south-eastern A ­ ustralia, with weak penetration into estuaries (Suter et al. 2022). Its biology is well understood (Yasser et al. 2018). References Alvarez F, Iliffe TM, Villalobos JL (2005) New species of the genus Typhlatya (Decapoda: Atyidae) from anchialine caves in Mexico, the Bahamas, and Honduras. Journal of Crustacean Biology 25, 81–94. doi:10.1651/C-2516 Havird JC, Vaught RC, Weese DA, Santos SR (2015) Reproduction and development in Halocaridina rubra Holthuis, 1963 (Crustacea: Atyidae) clarifies larval ecology in the Hawaiian anchialine ecosystem. The Biological Bulletin 229, 134–142. doi:10.1086/BBLv229n2p134 Holthuis LB (1993) The Recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda) with an appendix on the Order Amphionidacea. revised edn. Nationaal Natuuristorisch Museum, Leiden. Jaume D, Bréhier F (2005) A new species of Typhlatya (Crustacea: Decapoda: Atyidae) from anchialine caves on the French Mediterranean coast. Zoological Journal of the Linnean Society 144, 387–414. doi:10.1111/j.1096-3642.2005.00175.x Kensley B, Williams D (1986) New shrimps (families Procarididae and Atyidae) from submerged lava tube on Hawaii. Journal of Crustacean Biology 6, 417–437. doi:10.2307/1548182 Leberer T, Cai Y (2003) Shrimps of the family Atyidae from Guam, Mariana Island. Micronesica 35–36, 353–358. Page TJ, Baker AM, Cook BD, Hughes JD (2005) Historical transoceanic dispersal of a freshwater shrimp: the colonization of the South Pacific by the genus Paratya (Atyidae). Journal of Biogeography 32, 581–593. doi:10.1111/j.1365-2699.2004.01226.x Page TJ, von Rintelen K, Hughes JM (2007) Phylogenetic and biogeographic relationships of subterranean and surface genera of Australian Atyidae (Crustacea: Decapoda: Caridea) inferred with mitochondrial DNA. Invertebrate Systematics 21, 137–145. doi:10.1071/IS06023 Satake K, Cai Y (2005) Paratya boninensis, a new species of freshwater shrimp (Crustacea: Decapoda: Atyidae) from Ogasawara, Japan. Proceedings of the Biological Society of Washington 118, 306–311. doi:10.2988/0006-324X(2005)118[306:PBANSO]2.0.CO;2 Suter PJ, Mynott JH, Crump M (2022) New species of Paratya (Decapoda: Atyidae) from Australian inland waters – linking morphological characters with molecular lineages. Memoirs of Museum Victoria 81, 55–122. doi:10.24199/j.mmv.2022.81.04 Yasser AG, Sheldon F, Hughes JM (2018) Spatial distributions and environmental relationships of two species complexes of freshwater atyid shrimps. Ecosphere 9, e02388. doi:10.1002/ecs2.2388 Yoshihisa F, Davie PJF, Ng PKL (2015) A new stygobitic prawn of the genus Macrobrachium Spence Bate, 1864, from anchialine caves in Christmas Island, Indian Ocean; with a rediagnosis of M. miyakoense Komai & Fujita, 2005 (Crustacea: Decapoda: Caridea: Palaemonidae). Raffles Bulletin of Zoology 63, 610–625.

Superfamily Bresilioidea Calman, 1896 Fig. 6.14.  Atyidae. Paratya australiensis Kemp, 1917: a, anterior carapace; b, pereopod 1. Maxilliped 2: c, Paratya.

Separation of the five families of Bresilioidea began with confirmation of differences between Bresiliidae s.s. and

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Disciadidae, a family that had been proposed much earlier and then sunk (Forest 1977). This division was supported by Christoffersen (1986, 1990) who erected more families, partly in response to the discovery of Rimicaris and Alvinocaris at hydrothermal vents. Other families have been proposed and then synonymised with the consequence that the systematics of the superfamily has often changed (De Grave and Moosa 2004; Vereshchaka 1997). While the families are relatively easy to recognise, the superfamily is distinguished more by features that it lacks than any singular feature. Diagnosis. Mandibular molar conical, laminar or vestigial (Fig. 6.17c). Pereopods 1, 2 dissimilar, first stronger and heavier though sometimes shorter than second, fingers without distal spines. Pereopod 2 carpus entire.

Fig. 6.15.  Agostocarididae, Agostocaris: a, pereopod 1, carpus– dactylus; b, telson, uropod.

pairs of dorsolateraal spiniform setae, posterior margin with 4 pairs of spiniform setae, 2 pairs of fine setae (Fig. 6.15b).

References Christoffersen ML (1986) Phylogenetic relationships between Oplophoridae, Atyidae, Pasiphaeidae, Alvinocarididae fam. n., Bresiliidae, Psalidopodidae and Disciadidae (Crustacea Caridea Atyoidea). Boletim de Zoologia, Universidade de São Paulo 10, 273–281. Christoffersen ML (1990) A new superfamily classification of the Caridea (Crustacea: Pleocyemata) based on phylogenetic pattern. Journal of Zoological Systematics and Evolutionary Research 28, 94–106. doi:10.1111/j.1439-0469.1990.tb00369.x De Grave S, Moosa MK (2004) A new species of the enigmatic shrimp genus Pseudocheles (Decapoda: Bresiliidae) from Sulawesi (Indonesia), with the designation of a new family Pseudochelidae. Crustacean Research 33, 1–9. doi:10.18353/crustacea.33.0_1 Forest J (1977) Un groupement injustifié: la superfamilie des Bresilioida. Remarques critiques sur le statut du familles réunies sous ce nom (Crustacea Decapoda Caridea). Bulletin du Muséum National d’Histoire Naturelle, Paris, 3e série (Zoologie) 475, 869–888. Vereshchaka AL (1997) A new family for a deep-sea caridean shrimp from North Atlantic hydrothermal vents. Journal of the Marine Biological Association of the United Kingdom 77, 425–438. doi:10.1017/ S0025315400071770

Agostocarididae Hart & Manning, 1986 Figure 6.15 Hart and Manning (1986) created a new genus and family for shrimps from anchialine caves or cenotes of the Bahamas, Turks and Caicos islands. Mejía-Ortíz et  al. (2017) rediagnosed the family and Agostocaris, the only genus. They described a species from Cozumel I., Mexico, and tabulated differences between the four known species. Diagnosis. Carapace antennal spine absent. Eyes weakly pigmented. Mandible indistinctly divided into obsolete molar and incisor, with 2-articled palp. Pereopods 1, 2 ischium and merus articulated. Pereopod 1 carpus articulating with chela at middle of propodus below base of dactylus, propodus projecting (Fig. 6.15a). Exopod present on maxilliped 3, pereopods 1, 2. Telson with 4

Agostocaris Hart & Manning, 1986 Maximum cl. 7 mm. Marine anchialine caves. Tropical Atlantic (Caribbean islands only). 4 species. References Hart CW, Manning RB (1986) Two new shrimps (Procarididae and Agostocarididae, new family) from marine caves of the Western North Atlantic. Journal of Crustacean Biology 6, 408–416. doi:10.2307/1548181 Mejía-Ortíz L, Yañez G, López-Mejía M (2017) Anchialocarididae, a new family of anchialine decapods and a new species of the genus Agostocaris from Cozumel Island, Mexico. Crustaceana 90, 381–398. doi:10.1163/15685403-00003657

Alvinocarididae Christoffersen, 1986 Figure 6.16, Plate 8a Alvinocarid shrimps were discovered by the deep-diving submersible Alvin at thermal vents near the Galapagos Rift (Williams and Chace 1982). They have since been found across the globe. All inhabit reducing environments such as hydrothermal vents or brine and hydrocarbon seeps from lower shelf depths to around 5000 m depth (Ahyong 2009; Komai and Chan 2010; Komai et al. 2016; Komai and Segonzac 2003, 2008; Vereshchaka et al. 2015; Zelnio and Hourdez 2009). Many species are associated with chemoautotrophic faunas and some are seen exclusively on hydrothermal chimneys, often at extraordinary densities. The monophyly of Alvinocarididae was supported by morphological and molecular data (Christoffersen 1986; Vereshchaka et  al. 2015). Zelnio and Hourdez (2009) provided a key to seven genera and 22 species. Vereshchaka et al. (2015) also compiled a key to all genera and all species on which this one is based but their subfamilies are not adopted here.

6 – Caridea – shrimps

97

Fig. 6.16.  Alvinocarididae. Carapace (or rostrum, anterior margin): a, Alvinocaridinides; b, Alvinocaris; c, Keldyshicaris; d, Manuscaris; e, Mirocaris; f, Nautilocaris; g, Opaepele; h, Rimicaris; i, Shinkaicaris. Pleonite 3, pleuron: j, Mirocaris. Pleonites 3–5, pleura: k, Nautilocaris. Pereopod 1: l, Alvinocaris. Pereopod 3: m, Alvinocaris; n, Manuscaris. Maxilliped 3 epipod: o, Mirocaris. Telson (and right uropod): p, Manuscaris; q, Keldyshicaris; r, Opaepele; s, Rimicaris.

Diagnosis. Carapace antennal spine absent. Eyes greatly reduced, lacking faceted structure on corneal region in adults (Fig. 6.16a–i). Mandible distinctly divided into broad incisor and slender conical molar porcess, palp 2-articulated. Pereopods 1, 2 ischium and merus articulated. Pereopod 1 chela shaped like bird-head with bent beak, finger lower face excavated, cutting edges microscopically pectinate (Fig.  6.16l).

Exopods absent from pereopods 1–5. Arthrobranchs present on pereopods 1–4. Telson posterior margin rounded, with spiniform setae and/or plumose setae (Fig. 6.16p–s). Implicit generic attributes. Antennal tooth acute. Eyes broadly fused with shallow median notch, each anterior surface flat, unarmed. Telson dorsolateral spiniform setae in sinuous row, second and/or third mesial to others.

Key to genera of Alvinocarididae 1. – 2. – 3. – 4. –

Carapace with conspicuous postrostral carina extending beyond midlength (Fig. 6.16b). Pereopods 3–4 or 5 meri with spiniform setae (Fig. 6.16m)���������������������������������������������������������������������������������������������������������������Alvinocaris Carapace without conspicuous postrostral carina. Pereopods 3–5 meri without spiniform setae (Fig. 6.16n)����������������2 Epipods of maxilliped 3, pereopods 1–4 strap-like, ending in hook (Fig. 6.16o). Pleopods 2–4 appendices internae rudimentary�����������������������������������������������������������������������������������������������������������������������������������������������������������������3 Epipods of maxilliped 3, pereopods 1–4 rudimentary. Pleopods 2–4 appendices internae well developed���������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 4 Rostrum not reaching antennular peduncle article 1, dorsally smooth (Fig. 6.16e). Pleonite 3 pleuron smooth (Fig. 6.16j)��������������������������������������������������������������������������������������������������������������������������������������������������������Mirocaris Rostrum overreaching antennular peduncle article 1, with dorsal teeth (Fig. 6.16f). Pleonite 3 pleuron serrate (Fig. 6.16k)���������������������������������������������������������������������������������������������������������������������������������������������������Nautilocaris Rostrum overreaching antennular peduncle article 1, with dorsal teeth��������������������������������������������������������������������������������5 Rostrum not reaching antennular peduncle article 1, dorsally smooth����������������������������������������������������������������������������������6

98

5. – 6. – 7. – 8. –

Marine Decapod Crustacea

Rostrum and carapace bearing 11 or more dorsal teeth (Fig. 6.16d). Telson with linear row of dorsolateral spiniform setae (Fig. 6.16q). Pereopods 3, 4 ischia with spiniform setae (Fig. 6.16n)����������������������������������������������������������������������������������������������������������������������������������������������������������������� Manuscaris Rostrum and carapace 7–10 dorsal teeth (Fig. 6.16i). Telson with sinuous row of dorsolateral spiniform setae (Fig. 6.16s). Pereopods 3, 4 ischia without spiniform setae���������������������������������������������������������������������Shinkaicaris Rostrum very short, without dorsal teeth or notches (Fig. 6.16h). Uropodal exopod with 2 spiniform setae next to anterodistal tooth (Fig. 6.16s)�����������������������������������������������������������������������������������������������������������������������Rimicaris Rostrum with dorsal teeth or notches. Uropodal exopod with 1 spiniform seta next to anterodistal tooth (Fig. 6.16r)�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������7 Rostrum with acute tip, bearing > 10 dorsal teeth, ventrally unarmed (Fig. 6.16a). Pleonite 3 pleuron not serrate���������������������������������������������������������������������������������������������������������������������������������������������������������������Alvinocaridinides Rostrum with blunt tip, bearing < 7 dorsal teeth, ventrally with obsolete notches (Fig. 6.16c, g). Pleonite 3 pleuron with or without minute serrations�����������������������������������������������������������������������������������������������������������������������������8 Telson with sinuous row of dorsolateral setae, 1–3 pairs of spiniform setae on posterior margin (Fig. 6.16r). Pereopods 3, 5 ischia without spiniform setae. Pereopods 3–5 dactyli with 2 or more rows of accessory spinules�������������������������������������������������������������������������������������������������������������������������������������������������������������������������� Opaepele Telson with linear row of dorsolateral setae, 6 pairs of spiniform setae on posterior margin (Fig. 6.16q). Pereopods 3, 5 ischia with strong spiniform setae. Pereopods 3–5 dactyli with single row of accessory spinules������������������������������������������������������������������������������������������������������������������������������������������������������������������� Keldyshicaris Alvinocaridinides Komai & Chan, 2010

Manuscaris Komai & Tsuchida, 2015

Diagnosis. Rostrum dorsoventrally depressed, not overreaching distal margin of antennular peduncle article 1, blunt, dorsally toothed, ventrally smooth. Postrostral ridge low, not extending to midlength of carapace. Pleonite 3 pleuron not serrated. Maximum cl. 8.3 mm. Shelf (252–275 m). Central Indo-Pacific. 2 species (Komai and Chan 2010; Komai et al. 2016).

Diagnosis. Rostrum laterally compressed, overreaching distal margin of antennular peduncle article 1, acute, dorsally carinate, toothed, ventrally smooth. Postrostral ridge low, not extending to midlength of carapace. Pleonite 3 pleuron armed. Telson dorsolateral spiniform setae in linear row. Maximum cl. 8.4 mm. Slope (Manus Basin, 1310–1940 m). Central Indo-Pacific. 2 species (Komai and Tsuchida 2015; Wang and Sha 2016).

Alvinocaris Williams & Chace, 1982

Mirocaris Vereshchaka, 1997

Diagnosis. Rostrum laterally compressed, overreaching distal margin of antennular peduncle article 1, acute, dorsally carinate, toothed, ventrally carinate, with or without ventral tooth. Postrostral ridge high, extending beyond midlength of carapace. Eyes partly fused, with spiniform denticles on anterodorsal surface. Pereopods 3–4 or 5 meri with 1 or 2 lateral spiniform setae. Telson dorsolateral spiniform setae in linear row. Maximum cl. 20 mm (Pl. 8a). Slope, bathyal (252–3712 m). Temperate Northern Atlantic, Central Indo-Pacific, Tropical Eastern Pacific. GOODS bathyal provinces: Northern N Atlantic, N Atlantic, New Zealand/Kermadec, Nazca Plate, Indian. 16 species (Komai and Segonzac 2005).

Diagnosis. Rostrum dorsoventrally compressed, not reaching end of antennular peduncle article 1, obtuse, dorsally and ventrally not carinate, unarmed. Postrostral ridge absent, without dorsal teeth. Pleonite 3 pleuron not serrated. Eyes partly fused, anterior margin of cornea without tubercle. Maximum cl. 13 mm. Slope–bathyal (850–3480 m). GOODS bathyal provinces: N Atlantic, Indian. 2 species (Komai et al. 2006; Komai and Segonzac 2003).

Keldyshicaris Vereshchaka, Kulagin & Lunina, 2015 Diagnosis. Rostrum dorsoventrally compressed, not reaching end of antennular peduncle article 1, obtuse, dorsally not carinate, dorsally and ventrally notched. Postrostral ridge absent. Telson with 6 pairs of spiniform setae on posterior margin. Maximum cl. 11 mm. Bathyal. GOODS bathyal province: N Atlantic. 1 species (Lunina and Vereshchaka 2010; Vereshchaka et al. 2015).

Nautilocaris Komai & Segonzac, 2004 Diagnosis. Rostrum dorsoventrally compressed, overreaching end of antennular peduncle article 1, acute, dorsally and ventrally not carinate, toothed. Postrostral ridge absent, unarmed. Pleonite 3 pleuron serrated. Eyes partly fused, anterior margin of cornea without tubercle. Maximum cl. 8 mm. Bathyal (1604–2000 m). GOODS bathyal province: SE Pacific ridges. 1 species (Komai and Segonzac 2004).

Opaepele Williams & Dobbs, 1995 Diagnosis. Rostrum dorsoventrally compressed, not reaching end of antennular peduncle article 1, obtuse, dorsally and ventrally not carinate, notched or unarmed. Postrostral ridge absent. Pleonite 3

6 – Caridea – shrimps

pleuron with or without minute serrations. Telson with 1–3 pairs of spiniform setae on posterior margin Maximum cl. 13 mm. Slope–bathyal (980–2986 m). GOODS bathyal provinces: Northern N Pacific, N Atlantic. 2 species (Komai et al. 2007; Williams and Dobbs 1995).

Rimicaris Williams & Rona, 1986 Diagnosis. Rostrum dorsoventrally compressed, not reaching end of antennular peduncle article 1, obtuse, dorsally and ventrally not carinate, unarmed. Carapace with blunt antennal tooth (reduced in adults); postrostral ridge low, not extending to midlength of carapace. Maximum cl. 19 mm. Bathyal, abyssal (1640–4960 m). GOODS bathyal provinces: Northern N Atlantic, Northern N Pacific, SE Pacific ridges, Indian. 9 species. Some species have been treated as members of Chorocaris Martin & Hessler, 1990, now a junior synonym (Komai and Giguère 2019; Komai and Segonzac 2008; Komai and Tsuchida 2015). The ecology and relationship with ectosymbiotic bacteria of R. exoculata Williams & Rona, 1986, at hydrothermal vents is well researched (Zbinden and Cambon-Bonavita 2020).

Shinkaicaris Komai & Segonzac, 2005 Diagnosis. Rostrum laterally compressed, overreaching distal margin of antennular peduncle article 1, acute, dorsally carinate, toothed, ventrally smooth. Postrostral ridge extending slightly onto carapace, toothed. Pleonite 3 pleuron not serrated. Telson with sinuous row of dorsolateral spiniform setae. Maximum cl. 12.5 mm. Slope (705 m). Temperate Northern Pacific. 1 species (Komai and Segonzac 2005). References Ahyong ST (2009) New species and new records of hydrothermal vent shrimps from New Zealand (Caridea: Alvinocarididae, Hippolytidae). Crustaceana 82, 775–794. doi:10.1163/156854009X427333 Christoffersen ML (1986) Phylogenetic relationships between Oplophoridae, Atyidae, Pasiphaeidae, Alvinocarididae fam. n., Bresiliidae, Psalidopodidae and Disciadidae (Crustacea Caridea Atyoidea). Boletim de Zoologia, Universidade de São Paulo 10, 273–281. Komai T, Chan T-Y (2010) A new genus and two new species of alvinocaridid shrimps (Crustacea: Decapoda: Caridea) from a hydrothermal vent field off northeastern Taiwan. In: De Grave S, Fransen CHJM (Eds) Contributions to shrimp taxonomy. Zootaxa 2372, 15–32. doi:10.11646/zootaxa.2372.1.3 Komai T, Giguère T (2019) A new species of alvinocaridid shrimp Rimicaris Williams & Rona, 1986 (Decapoda: Caridea) from hydrothermal vents on the Mariana Back Arc Spreading Center, northwestern Pacific. Journal of Crustacean Biology 39, 640–650. doi:10.1093/jcbiol/ruz046 Komai T, Segonzac M (2003) Review of the hydrothermal vent shrimp genus Mirocaris, redescription of M. fortunata and reassessment of the taxonomic status of the family Alvinocarididae (Crustacea: Decapoda: Caridea). Cahiers de Biologie Marine 44, 199–215. Komai T, Segonzac M (2004) A new genus and species of alvinocaridid shrimp (Crustacea: Decopoda: Caridea) from hydrothermal vents on the North Fiji and Lau Basins, south-western Pacific. Journal of the Marine Biological Association of the United Kingdom 84, 1179–1188. doi:10.1017/S0025315404010628h

99

Komai T, Segonzac M (2005) A revision of the genus Alvinocaris Williams and Chace (Crustacea: Decapoda: Caridea: Alvinocarididae), with descriptions of a new genus and a new species of Alvinocaris. Journal of Natural History 39, 1111–1175. doi:10.1080/00222930400002499 Komai T, Segonzac M (2008) Taxonomic review of the hydrothermal vent shrimp genera Rimicaris Williams & Rona and Chorocaris Martin & Hessler (Crustacea: Decapoda: Caridea: Alvinocarididae). Journal of Shellfish Research 27, 21–41. doi:10.2983/0730-8000(2008)27[21:TROTHV]2.0.CO;2 Komai T, Tsuchida S (2015) New records of Alvinocarididae (Crustacea: Decapoda: Caridea) from the southwestern Pacific hydrothermal vents, with descriptions of one new genus and three new species. Journal of Natural History 49, 1789–1824. doi:10.1080/00222933.2015. 1006702 Komai T, Martin JW, Zala K, Tsuchida S, Hashimoto J (2006) A new species of Mirocaris (Crustacea: Decapoda: Caridea: Alvinocarididae) associated with hydrothermal vents on the Central Indian Ridge, Indian Ocean. Scientia Marina 70, 109–119. doi:10.3989/scimar.2006.70n1109 Komai T, Giere O, Segonzac M (2007) New record of alvinocaridid shrimps (Crustacea: Decapoda: Caridea) from hydrothermal vent fields on the southern Mid-Atlantic Ridge, including a new species of the genus Opaepele. Species Diversity 12, 237–253. doi:10.12782/ specdiv.12.237 Komai T, Menot L, Segonzac M (2016) New records of caridean shrimp (Crustacea: Decapoda) from hydrothermally influenced fields off Futuna Island, Southwest Pacific, with description of a new species assigned to the genus Alvinocaridinides Komai & Chan, 2010 (Alvinocarididae). Zootaxa 4098, 298–310. doi:10.11646/zootaxa.4098.2.5 Lunina A, Vereshchaka A (2010) A new vent shrimp (Crustacea: Decapoda: Alvinocarididae) from the Mid-Atlantic Ridge. Zootaxa 2372, 69–74. doi:10.11646/zootaxa.2372.1.8 Vereshchaka AL, Kulagin DN, Lunina AA (2015) Phylogeny and new classification of hydrothermal vent and seep shrimps of the family Alvinocarididae (Decapoda). PLoS One 10, e0129975. doi:10.1371/journal. pone.0129975 Wang Y-L, Sha Z-L (2016) A new species of the genus Manuscaris Komai & Tsuchida, 2015 (Decapoda, Caridea, Alvinocarididae) from the Manus Basin hydrothermal vents. Crustaceana 89, 1541–1550. doi:10.1163/15685403-00003591 Williams AB, Chace FA (1982) A new caridean shrimp of the family Bresiliidae from thermal vents of the Galapagos Rift. Journal of Crustacean Biology 2, 136–147. doi:10.2307/1548118 Williams A, Dobbs FC (1995) A new genus and species of caridean shrimp (Crustacea, Decapoda, Bresiliidae) from hydrothermal vents on Loihi Seamount, Hawaii. Proceedings of the Biological Society of Washington 108, 228–237. Zbinden M, Cambon-Bonavita MA (2020) Rimicaris exoculata: biology and ecology of a shrimp from deep-sea hydrothermal vents associated with ectosymbiotic bacteria. Marine Ecology Progress Series 652, 187–222. doi:10.3354/meps13467 Zelnio KA, Hourdez S (2009) A new species of Alvinocaris (Crustacea: Decapoda: Caridea: Alvinocarididae) from hydrothermal vents at the Lau Basin, southwest Pacific, and a key to the species of Alvinocarididae. Proceedings of the Biological Society of Washington 122, 52–71. doi:10.2988/07-28.1

Bresiliidae Calman, 1896 Figure 6.17, Plate 8b Bresiliidae, once a much larger family, are now restricted to two genera of small shrimps. Both are notable for the strongly

100

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Fig. 6.17.  Bresiliidae. a, Bresilia antipodarum Bruce, 1990. Bresilia: b, pereopod 1 chela; c, mandible; d, telson. Encantada spinoculata Wicksten, 1989: e, anterior carapace, eyestalks.

humped third pleonite. The presence or absence of a spine on the eyestalk distinguishes the two genera. Diagnosis. Carapace antennal spine present (Fig.  6.17e). Eyes with fully formed cornea. Mandible distinctly divided into broad incisor and slender conical molar process, palp 2-articulated (Fig. 6.17c). Pereopods 1, 2 ischium and merus articulated. Pereopods 3–5 simple. Exopods present on maxilliped 3, pereopods 1, 2. Arthrobranchs absent. Telson with posterior triangular endpiece, with 3 pairs of dorsolateral spiniform setae (Fig. 6.17d). Bresilia Calman, 1896 Diagnosis. Eyestalk without dorsal spine. Exopod on pereopods 1 and 2, arthrobranchs and pleurobranchs on pereopods 1–4. Maximum cl. 4.5 mm (including rostrum) (Pl. 8b). Shelf, slope (133–1372 m, plus shallow marine caves). Temperate Northern Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Australasia. 10 species (Komai and Kohtsuka 2017: key to species).

Encantada Wicksten, 1989 Diagnosis. Eyestalk with dorsal spine overreaching base of cornea (Fig. 6.17e). Exopod and pleurobranchs on pereopods 1–5, arthrobranchs on pereopods 1–3. Maximum cl. 3.5 mm. Shelf (55–92 m). Tropical Eastern Pacific (Galapagos Is). 1 species. The only species, Encantada spinoculata Wicksten, 1989 has never been seen since its original description (Bruce 1990; Wicksten 1989).

References Bruce AJ (1990) Two deep-sea shrimps new to the Australian fauna, Psathyrocaris hawaiiensis Rathbun (Pasiphaeidae) and Bresilia antipodarum, sp. nov. (Bresiliidae), with remarks on Encantada spinoculata Wicksten (Bresiliidae). Invertebrate Taxonomy 4, 847–866. doi:10.1071/IT9900847 Komai T, Kohtsuka H (2017) A new deep-sea species of the caridean genus Bresilia Calman, 1896 (Crustacea: Decapoda: Bresiliidae) from Sagami Bay, central Japan. Zootaxa 4299, 405–414. doi:10.11646/zootaxa.4299.3.6 Wicksten MK (1989) Encantada spinoculata, a new genus and species of shrimp from the Galapagos Islands (Caridea: Bresiliidae). Journal of Crustacean Biology 9, 667–671. doi:10.2307/1548596

Disciadidae Rathbun, 1902 Figure 6.18 Disciadidae comprise four genera, displaying a variety of ecologies. Discias is the best known and most easily recognised because of its disc-like dactylus on the first cheliped. The family was justified and separated from Bresiliidae by Forest (1977). Diagnosis. Carapace antennal spine absent. Eyes with fully formed cornea. Mandible distinctly divided into broad incisor and slender conical molar process, palp 2-articulated. Pereopods 1, 2 ischium and merus articulated. Pereopod 1 chela with dactylus semicircular, disc-like with sharp edge (Fig.  6.18f–h). Exopod present on maxilliped 3, pereopods 1–5 (rarely 1–3). Arthrobranchs absent. Telson posterior margin rounded, with spiniform setae and/or plumose setae. Implicit generic attribute. Exopod on pereopods 1–5.

Key to genera of Disciadidae 1.

Rostrum blade-like, dorsally and ventrally serrate (Fig. 6.18b). Scaphocerite narrowing distally, blade not overreaching distolateral tooth (Fig. 6.18c)��������������������������������������������������������������������������������������������������������������� Kirnasia

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Fig. 6.18.  Disciadidae. a, Discias brownae Kensley, 1983. Anterior carapace, antennule, scaphocerite: b, c, Kirnasia; d, Lucaya; e, Tridiscias. End of chela: f, Discias; g, Lucaya; h, Tridiscias.

– 2. – 3. –

Rostrum flat, unarmed (Fig. 6.18a, d, e). Scaphocerite broad distally, blade overreaching distolateral tooth (Fig. 6.18d, e)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Rostrum reaching end of antennular peduncle (Fig. 6.18d). Pleonite 3 hump-like over pleonite 4. Pereopod 1 dactylus quadrant-shaped, flat, opposing straight short acute fixed finger (Fig. 6.18g)����������������������������������� Lucaya Rostrum not reaching to end of antennular peduncle (Fig. 6.18a, e). Pleonite 3 dorsally continuous with pleonite 4 (Fig. 6.18a). Pereopod 1 dactylus disc-like or adze-shaped, opposing curved short fixed finger (Fig. 6.18f, h)���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Carapace with supraorbital and branchiostegal spines (Fig. 6.18e). Telson with 5 dorsolateral, 5 posterior spiniform setae�������������������������������������������������������������������������������������������������������������������������������������������������������������Tridiscias Carapace without supraorbital and branchiostegal spines (Fig. 6.18a). Telson with 2 or 3 dorsolateral, 3 posterior spiniform setae������������������������������������������������������������������������������������������������������������������������������������������������ Discias Discias Rathbun, 1902

Lucaya Chace, 1939

Diagnosis. Rostrum flat, triangular, not reaching end of antennular peduncle. Carapace without supraorbital and branchiostegal spines. Pleonite 3 dorsally continuous with pleonite 4. Pereopod 1 dactylus disc-like, opposing short curved, acute fixed finger with angled tip. Telson with 2 or 3 dorsolateral, 3 posterior spiniform setae. Maximum cl. 4 mm. Intertidal–slope (0–246 m). Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Temperate South America (Easter I.), Temperate Southern Africa, Temperate Australasia (associated with sponges). 8 species (Kensley 1983: key to 7 species).

Diagnosis. Rostrum flat, triangular, reaching to end of antennular peduncle. Carapace without supraorbital and branchiostegal spines. Pleonite 3 hump-like over pleonite 4. Pereopod 1 dactylus quadrant-shaped, flat, opposing straight short acute fixed finger. Telson with 2 pairs of dorsolateral spiniform setae. Maximum cl. 4.7 mm. Mesopelagic, bathypelagic. Gulf of Mexico, E Atlantic. 1 species (Chace 1940; Forest 1977). Lucaya bigelowi Chace, 1939, is the only pelagic species in the family but is rare.

Kirnasia Burukovsky, 1988 Diagnosis. Rostrum blade-like, dorsally serrate, reaching beyond antennal peduncle. Carapace with supraorbital and branchiostegal spines. Pleonite 3 hump-like over pleonite 4. Scaphocerite narrowing distally, blade not overreaching distolateral tooth. Telson with 2 pairs of dorsolateral spiniform setae. Slope (850–1200 m). Temperate Northern Atlantic. 2 species (Burukovsky 1988).

Tridiscias Kensley, 1983 Diagnosis. Rostrum flat, triangular, not reaching to end of antennular peduncle. Carapace with supraorbital and branchiostegal spines. Pleonite 3 dorsally continuous with pleonite 4. Pereopod 1 dactylus adze-shaped, flat opposing fixed finger. Telson with 5 dorsolateral, 5 posterior spiniform setae. Exopod on pereopods 1–3. Maximum cl. 2 mm. Shelf (150–200 m). Temperate Southern Africa. 1 species (Kensley 1983).

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References

Pseudocheles Chace & Brown, 1978

Burukovsky RN (1988) New representatives of the family Bresiliidae (Crustacea, Decapoda) from the north-east Atlantic [in Russian]. Zoologicheskii Zhurnal 67, 456–460. Chace FA (1940) Plankton of the Bermuda Oceanographic Expeditions. IX. The bathypelagic caridean Crustacea. Zoologica 25, 117–209. Forest J (1977) Un groupement injustifié: la superfamilie des Bresilioida. Remarques critiques sur le statut du familles réunies sous ce nom (Crustacea Decapoda Caridea). Bulletin du Muséum National d’Histoire Naturelle, Paris, 3e série (Zoologie) 475, 869–888. Kensley B (1983) New records of bresiliid shrimp from Australia, South Africa, Caribbean, and Gulf of Mexico (Decapaoda: Natantia: Caridea). Smithsonian Contributions to Zoology 394, 1–31. doi:10.5479/ si.00810282.394

Maximum cl. 1.7 mm. Subtidal, shelf (1–71 m). Tropical Atlantic, Central Indo-Pacific. 4 species (Anker 2012).

Pseudochelidae De Grave & Moosa, 2004

Superfamily Campylonotoidea Sollaud, 1913

Figure 6.19 Pseudochelid shrimps, first discovered on the Great Barrier Reef, Australia, differ from all other carideans in having all pereopods functionally chelate. The first two pairs are truly chelate, but on the last three, the dactylus opposes a long strong seta (Anker 2012). All species belong to one genus, and seem to favour coral reefs (De Grave and Moosa 2004). Diagnosis. Carapace antennal spine present. Eyes with fully formed cornea. Mandible indistinctly divided into obsolete molar and incisor, with 2-articled palp. Pereopods 1, 2 ischium and merus fused. Pereopods 3–5 pseudochelate (dactylus opposing distal spiniform seta). Exopod present on maxilliped 3, pereopods 1–5. Arthrobranchs absent. Telson with posterior triangular endpiece, with 3 pairs of dorsolateral spiniform setae.

The superfamily comprises two families of deep-water benthic shrimps distinguished most obviously on the equality or otherwise of pereopods 2. Diagnosis. Mandible not deeply bifurcate, with palp. Pereopods 1, 2 dissimilar, first more slender than second. Pereopod 2 carpus entire. Exopod absent from pereopods 1–5. Arthrobranchs present on pereopods 1–4.

References Anker A (2012) The shrimp family Pseudochelidae De Grave & Moosa, 2004 (Crustacea, Decapoda, Caridea) in Brazil, with the description of a new species. Zootaxa 3544, 79–87. doi:10.11646/zootaxa.3544.1.7 De Grave S, Moosa MK (2004) A new species of the enigmatic shrimp genus Pseudocheles (Decapoda: Bresiliidae) from Sulawesi (Indonesia), with the designation of a new family Pseudochelidae. Crustacean Research 33, 1–9. doi:10.18353/crustacea.33.0_1

Bathypalaemonellidae de Saint Laurent, 1985 Figure 6.20, Plate 8c Bathypalaemonellidae were separated from Campylonotidae by de Saint Laurent (1985). Cleva (2001) reviewed the family and provided keys to genera and all species. These shrimps are associated with deep-water corals (Watling 2010; Wicksten and Heathman 2015). Diagnosis. Maxilliped 2 last article forming oblique suture with penultimate article (Fig.  6.20e). Pereopods 2 very unequal. Epipods absent. Bathypalaemonella Balss, 1914 Diagnosis. Carapace with 3–7 postrostral spines (Fig.  6.20a). Major pereopod 2 ischium longer than merus; dactylus compressed laterally, lower margin smooth (Fig.  6.20c). Tubercle between pereopodal coxae 2 present. Maximum cl. 16 mm (Pl. 8c). Slope–bathyal (215–2750 m; from black coral, octocorals and gorgonian hosts). Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific. 8 species.

Bathypalaemonetes Cleva, 2001

Fig. 6.19.  Pseudochelidae. a, Pseudocheles chacei Kensley, 1983; b, pereopod 3, Pseudocheles.

Diagnosis. Carapace with 1 postrostral spine (Fig.  6.20b). Major pereopod 2 ischium shorter than merus; dactylus lower margin tuberculate (Fig.  6.20d). Tubercle between pereopodal coxae 2 absent. Maximum cl. 12.5 mm. Slope (221–700 m). Temperate Northern W Pacific, Central Indo-Pacific. 3 species.

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103

Fig. 6.21.  Campylonotidae. Campylonotus rathbunae Schmitt, 1926: a, habitus; b, maxilliped 2.

Campylonotus Bate, 1888

Fig. 6.20.  Bathypalaemonellidae. Anterior carapace, rostrum: a, Bathypalaemonella; b, Bathypalaemonetes. Major pereopod 2: c Bathypalaemonella; d, Bathypalaemonetes. Maxilliped 2: e, Bathypalaemonella

References Cleva R (2001) Les Bathypalaemonellidae de Saint-Laurent, 1985 (Crustacea, Decapoda, Caridea) avec description d’une espèce nouvelle et définition d’un genre nouveau. Zoosystema 23, 757–782. de Saint Laurent M (1985) Remarques sur la distribution des crustacés Décapodes. In Peuplements profonds du Golfe de Gascoyne. (Eds Laubier L, Monniot C) pp. 469–478. IFREMER, Paris. Watling L (2010) Notes on the habitat of the deep-sea caridean shrimp, Bathypalaemonella serratipalma Pequegnat. In: Fransen C, De Grave S, Ng P (Eds) Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume. Crustaceana Monographs 14, 707–714. doi:10.1163/9789047427759_052 Wicksten MK, Heathman T (2015) New host records for Bathypalaemonella serratipalma (Caridea: Bathypalaemonellidae) in the northern Gulf of Mexico. Marine Biodiversity Records 8, e96. doi:10.1017/ S1755267215000779

Campylonotidae Sollaud, 1913 Figure 6.21, Plate 8d The family comprises sublittoral to deep-sea benthic species in subantarctic and cool temperate regions of the southern hemisphere (Thatje 2003). Some species are fished commercially as Sabre Shrimps or Razor Shrimps (Arana and Ahumada 2006). Diagnosis. Maxilliped 2 last article a narrow strip on mesial margin of penultimate article (Fig. 6.0b). Pereopods 2 subequal (Fig. 6.21a). Epipods present.

Maximum cl. 35 mm. Subtidal–bathyal (18–2086 m). Temperate South America, Temperate Australasia, Southern Ocean. 5 species (Thatje 2003: key to species) (Pl. 8d). References Arana P, Ahumada M (2006) Camarón navaja (Campylonotus semistriatus), crustáceo de aguas profundas frente a la costa central de Chile (Crustacea, Decapoda, Campylonotidae). Investigaciones Marinas 34, 3–14. doi:10.4067/S0717-71782006000100001 Thatje S (2003) Campylonotus arntzianus, a new species of the Campylonotidae (Crustacea: Decapoda: Caridea) from the Scotia Sea (Antarctica. Polar Biology 26, 242–248. doi:10.1007/s00300-002-0469-5

Superfamily Crangonoidea Haworth, 1825 The two families are united on the subchelate/prehensile form of pereopod 1 described in detail for many genera by Wood and De Grave (2015). Diagnosis. Carapace subcylindrical or more-or-less dorsoventrally compressed. Pereopods 1, 2 dissimilar, each pair symmetrical, first more robust than second. Pereopod 1 subchelate, dactylus closing against subtruncate distal margin of propodus, or prehensile, carpus closing against inner surface of propodus. Pereopod 2 carpus entire or multiarticulate. Exopod present on maxilliped 3, pereopods 1–5. Reference Wood LE, De Grave S (2015) Functional morphology of the first pereiopod in crangonoid shrimps (Crustacea, Decapoda, Caridea, Crangonoidea). Zoomorphology 134, 469–486. doi:10.1007/s00435-015-0270-y

Crangonidae Haworth, 1825 Figures 6.22, 6.23, Plate 8e–i

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Fig. 6.22.  Crangonidae. a, Lissosabinea beresfordi Taylor & Collins, 2009; b, Paracrangon australis Hanamura, Wadley & Taylor, 1999; c, Philocheras victoriensis (Fulton & Grant, 1902); d, Vercoia gibbosa Baker, 1904; e, Sclerocrangon rex Komai & Matsuzaki, 2016.

Crangonids are small, robust, often somewhat flattened shrimps with characteristic subchelate first pereopods. The prehensile relationship between the dactylus and palm is seen elsewhere only in the Glyphocrangonidae, a family known only from deep water and with a more sculptured carapace. Some species of Crangon are commercially important.

Holthuis (1993) recognised 21 genera in his key, but with additions and deletions, 23 are now accepted. His key is the basis of the one below, with additional genera. Diagnosis. Pereopod 1 subchelate, dactylus closing against subtruncate distal margin of propodus.

Key to genera of Crangonidae 1. – 2. – 3. – 4. – 5. – 6. – 7. –

Pereopod 2 absent or rudimentary. Rostrum long, slender, obliquely erect, acute, with basal ventral curved tooth (Fig. 6.22b)���������������������������������������������������������������������������������������������������������������������������������������������������� Paracrangon Pereopod 2 present. Rostrum otherwise���������������������������������������������������������������������������������������������������������������������������������������2 Pereopod 2 simple����������������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Pereopod 2 chelate����������������������������������������������������������������������������������������������������������������������������������������������������������������������������7 Eyes reduced to small pointed processes; cornea absent (Fig. 6.23j)����������������������������������������������������������������Prionocrangon Eyes well developed; cornea large, well pigmented���������������������������������������������������������������������������������������������������������������������4 Pereopod 2 well developed, with broad articles, reaching beyond end of pereopod 1 merus (Fig. 6.22d). Scaphocerite without terminal tooth��������������������������������������������������������������������������������������������������������������������������� Vercoia Pereopod 2 rudimentary, thin, short, not reaching end of pereopod 1 merus (Fig. 6.23v). Scaphocerite with terminal tooth������������������������������������������������������������������������������������������������������������������������������������������������������������������������������5 Eye strongly dorsoventrally flattened, incompletely fused with cephalothorax, cornea and eyestalk not separated, dorsal surface reflective (Fig. 6.23g)������������������������������������������������������������������������������������������ Placopsicrangon Eye typically spherical���������������������������������������������������������������������������������������������������������������������������������������������������������������������6 Pleon dorsally carinate. Carapace with 3 denticulate carinae on each side (Fig. 6.23q)�������������������������������������������Sabinea Pleonites 1, 2, 4, 5 without dorsal carina. Carapace with 2 carinae on each side, each with at most 2 teeth (Fig. 6.22a)����������������������������������������������������������������������������������������������������������������������������������������������������������������Lissosabinea Pereopods 4, 5 dactyli flat, broad, natatorial (Fig. 6.23x)�������������������������������������������������������������������������������������������������� Argis Pereopods 4, 5 dactyli not broad����������������������������������������������������������������������������������������������������������������������������������������������������8

6 – Caridea – shrimps

8. – 9. – 10. – 11.

105

Pereopod 2 subequal in length to the other pereopods��������������������������������������������������������������������������������������������������������������9 Pereopod 2 shorter than other pereopods����������������������������������������������������������������������������������������������������������������������������������17 Carapace with 1 or 2 submedian spines between middorsal line and hepatic spine�����������������������������������������������������������10 Carapace without submedian spines between middorsal line and hepatic spine����������������������������������������������������������������11 Pleonite 6 posterolateral angle strongly flared (Fig. 6.23t). Pleon moderately sculptured����������������������������� Metacrangon Pleonite 6 posterolateral angle not flared (Fig. 6.23s). Pleon not or hardly sculptured���������������������������������� Mesocrangon Integument flexible. Pleon lateral surface not sculptured. Carapace branchiostegal spine not strong or flared sideways�������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 12

Fig. 6.23.  Crangonidae. Carapace, dorsal views: a, Aegaeon; b, Crangon; c, Lissocrangon; d, Mesocrangon; e, Metacrangon; f, Parapontocaris; g, Placopsicrangon; h, Pontocaris; i, Pontophilus; j, Prionocrangon; k, Rhynocrangon; l, Syncrangon. Carapace, lateral views: m, Neocrangon; n, Notocrangon; o, Parapontophilus; p, Pseudopontophilus; q, Sabinea; r, Pontocaris. Pleonite 6, dorsal views: s, Mesocrangon; t, Metacrangon. Pereopod 1: u, Pontophilus; Pereopod 2: v, Placopsicrangon; w, Vercoia. Pereopod 4: x, Argis. Male pleopod 2: y, Crangon; z, Notocrangon; A, Sclerocrangon; B, Rhynocrangon.

106

Marine Decapod Crustacea



Integument of carapace and pleon thick, strong. Pleon with sculptured surface. Carapace branchiostegal spine strong, flared sideways���������������������������������������������������������������������������������������������������������������������������������������������������15 12. Carapace middorsal line without teeth (Fig. 6.23c)�����������������������������������������������������������������������������������������������Lissocrangon – Carapace middorsal line with 1 or 2 teeth���������������������������������������������������������������������������������������������������������������������������������13 13. Carapace middorsal line with 2 teeth (Fig. 6.23m)��������������������������������������������������������������������������������������������������Neocrangon – Carapace middorsal line with 1 tooth (Fig. 6.23n)��������������������������������������������������������������������������������������������������������������������14 14. Maxilliped 3 without arthrobranch. Male pleopod 2 endopod blade vestigial, far overreached by appendix masculina (Fig. 6.23z). Pleonite 6 dorsal ridges ending in spine. Antarctic������������������������������������������������� Notocrangon – Maxilliped 3 with arthrobranch. Male pleopod 2 endopod blade longer than appendix masculina (Fig. 6.23g). Pleonite 6 dorsal ridges if present not ending in spine. N Hemisphere�������������������������������������������Crangon 15. Male pleopod 2 endopod blade small, triangular, appendix masculina adjacent to blade (Fig. 6.23B)������ Rhynocrangon – Male pleopod 2 endopod blade reduced, shorter than appendix masculina������������������������������������������������������������������������16 16. Carapace without transverse groove on cardiac region. Pleonite 6 with obtuse submedian carinae, laterally not grooved (Fig. 6.22e)��������������������������������������������������������������������������������������������������������������������������������������� Sclerocrangon – Carapace with 2 transverse grooves on cardiac region (Fig. 6.23l). Pleonite 6 with flattened middorsal carina, laterally grooved������������������������������������������������������������������������������������������������������������������������������������������ Syncrangon 17. With 1 arthrobranch on maxilliped 3, pleurobranchs above pereopods 1–5, their apices directed posteriorly ������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������18 – With 2 arthrobranchs on maxilliped 3, pleurobranchs above pereopods 1–5, their apices directed anteriorly�������������21 18. Rostrum without lateral teeth. Carapace without postorbital suture (Fig. 6.22c)������������������������������������������������Philocheras – Rostrum with 1 or 2 basal lateral teeth. Carapace with postorbital suture���������������������������������������������������������������������������19 19. Carapace with 0–24 spines on posterior half of middorsal carina (Fig. 6.23p). Pereopod 1 without exopod. Pereopod 2 reaching pereopod 1 merus distal margin��������������������������������������������������������������������������Pseudopontophilus – Carapace with 1–4 spines on anterior half of middorsal carina. Pereopod 1 with rudimentary exopod. Pereopod 2 not reaching pereopod 1 merus distal margin����������������������������������������������������������������������������������������������� 20 20. Carapace middorsal line with epigastric tooth plus 1 or 2 teeth on posterior half. Carapace lateral surface with 3 or 5 prominent spines (Fig. 6.23i)��������������������������������������������������������������������������������������������������������������Pontophilus – Carapace middorsal line with 1 or 2 epigastric teeth, sometimes with tooth on posterior half. Carapace lateral surface with 1 or 2 prominent spines (Fig. 6.23o)���������������������������������������������������������������������������Parapontophilus 21. Rostrum ending in single apex, with 2 pairs of lateral teeth. Carapace with 4 or 5 teeth on first lateral carina (Fig. 6.23f). Pleonal sternites unarmed���������������������������������������������������������������������������������������������� Parapontocaris – Rostrum ending in 1 or 2 apices, without or with 1 lateral tooth. Carapace with 6–12 teeth on first lateral carina. Pleonal sternites with median spines (Fig. 6.23r)�������������������������������������������������������������������������������������������������� 22 22. Carapace middorsal line with 4 or 5 teeth (Fig. 6.23a)��������������������������������������������������������������������������������������������������Aegaeon – Carapace middorsal line with 7 or more teeth (Fig. 6.23h)�������������������������������������������������������������������������������������� Pontocaris Aegaeon Agassiz, 1846 Diagnosis. Rostrum very short, apex simple or cleft, with 1 pair of inconspicuous lateral spines near base. Carapace middorsal line with 4 or 5 teeth, with 6–12 teeth on first lateral carina. Pereopod 2 shorter than other pereopods. Gills: 2 arthrobranchs on maxilliped 3, pleurobranchs above pereopods 1–5, their apices directed anteriorly. Maximum cl. 12 mm (Pl. 8e). Subtidal–slope (12–809 m). World-wide tropical and temperate seas except E Pacific. 5 species (Chan 1996: key to species).

Argis Krøyer, 1843 Diagnosis. Rostrum shorter than eyestalk, erect, acute. Pereopods 4, 5 dactyli flat, broad, natatorial. Maximum cl. 34 mm.

Intertidal–bathyal (0–2090 m). Temperate Northern Atlantic, Temperate Northern Pacific (A. dentata only). 10 species (Komai 1997: key to species based on mature females).

Crangon Fabricius, 1798 Diagnosis. Rostrum short, narrowly triangular. Carapace middorsal line with 1 tooth, without submedian spines between middorsal line and hepatic spine, branchiostegal spine not strong or flared sideways. Pleon lateral surface not sculptured; pleonite 6 dorsal ridges absent or not ending in spine. Pereopod 2 subequal in length to the other pereopods. Maximum cl. 18 mm (Pl. 8f). Intertidal–slope (0–630 m). Arctic, Temperate Northern Atlantic, Temperate Northern Pacific. 19 species. Crangon is a large Northern Hemisphere genus. The brown shrimp, Crangon

6 – Caridea – shrimps

crangon (Linnaeus, 1758), is the basis of a commercial and recreational fishery in the southern North Sea but occurs outside this area. For this reason, and its general abundance and availability, the species is the subject of extensive ecological and physiological research. The taxonomy of the genus as a whole has never been reviewed but Hayashi (1999) dealt with the seven East Asian species.

Lissocrangon Kuris & Carlton, 1977 Diagnosis. Rostrum as long as eyestalk, narrow, acute, curved down. Carapace middorsal line without teeth, without submedian spines between middorsal line and hepatic spine, branchiostegal spine not strong or flared sideways. Pleon lateral surface not sculptured. Pereopod 2 subequal in length to the other pereopods. Maximum cl. 12 mm. Intertidal–shelf (0–80 m). Temperate Northern Pacific (common in bays along the NW American coast). 1 species (Jarrin and Shanks 2008; Kuris and Carlton 1977).

Lissosabinea Christoffersen, 1988 Diagnosis. Rostrum wider over proximal half with pair of lateral teeth, anterior half acute. Pleonites 1, 2, 4, 5 without dorsal carina. Pereopod 2 simple, well developed, with broad articles, reaching beyond end of pereopod 1 merus. Maximum cl. 8.5 mm (Pl. 8g). Shelf, slope (110–830 m). Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate ­Australasia. 8 species (Taylor and Collins 2009: key to species).

Mesocrangon Zarenkov, 1965 Diagnosis. Rostrum short, without lateral teeth. Carapace with 1 or 2 submedian spines between middorsal line and hepatic spine. Pleonite 6 posterolateral angle not flared. Pereopod 2 subequal in length to the other pereopods. Maximum cl. 8 mm. Subtidal–slope (10–400 m). Arctic, Temperate Northern Pacific. 3 species. The three species are confined to cold waters (Zarenkov 1965) but one reaches as far south as the Gulf of ­California (Hendrickx 1993). The megalopa is better illustrated than the adult (Sedova and Grigoriev 2014a).

Metacrangon Zarenkov, 1965 Diagnosis. Rostrum short, without lateral teeth. Carapace with 1 or 2 submedian spines between middorsal line and hepatic spine. Pleonite 6 posterolateral angle strongly flared. Pereopod 2 subequal in length to the other pereopods. Maximum cl. 20 mm (Pl. 8i). Subtidal–bathyal (10–2850 m). Worldwide except polar seas, including hydrothermal vent fields (Komai et al. 2018). 46 species. The genus is divided into two informal species-groups: the M. jacqueti group of 16 species; and the M. munita group of 30 species (Komai and Ahyong 2010) (Komai 2012; Komai et  al. 2018; Komai and Taylor 2010). The only comprehensive key dealt with 21 species from the W Pacific (Komai 2012) but more have been described since (Kim and Chan 2020).

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Neocrangon Zarenkov, 1965 Diagnosis. Rostrum longer than eyestalk, laterally compressed. Carapace middorsal line with 2 teeth, without submedian spines between middorsal line and hepatic spine, branchiostegal spine not strong or flared sideways. Pleon lateral surface not sculptured. Pereopod 2 subequal in length to the other pereopods. Maximum cl. 10 mm. Subtidal–abyssal (16–4000 m). Arctic, Temperate Northern Pacific, Central Indo-Pacific. 6 species. The differences between Neocrangon and Mesocrangon have been questioned on the basis of evidence from larvae (Sedova and Grigoriev 2014b). Han and Li (2009) described the most recent species and distinguished it from three others.

Notocrangon Coutière, 1900 Diagnosis. Rostrum longer than eyestalk, laterally compressed. Carapace middorsal line with 1 tooth, without submedian spines between middorsal line and hepatic spine, branchiostegal spine not strong or flared sideways. Pleon lateral surface not sculptured; pleonite 6 dorsal ridges ending in spine. Pereopod 2 subequal in length to the other pereopods. Maximum cl. 21 mm. Subtidal–slope (5–1320 m). Southern Ocean. 1 species. (Komai et  al. 1996: illustration of only species, N. antarcticus (Pfeffer, 1887)).

Paracrangon Dana, 1852 Diagnosis. Rostrum long, slender, obliquely erect, acute, with ventral recurved tooth. Pereopod 2 absent or rudimentary. Maximum cl. 19 mm. Subtidal–slope, including hydrothermal areas (15–1422 m). Temperate Northern Pacific, Tropical Eastern Pacific, Temperate Australasia. 7 species (Komai and Kim 2004: key to species).

Parapontocaris Alcock, 1901 Diagnosis. Rostrum ending in 2 apices, without or with 1 pair of lateral teeth. Carapace middorsal line with 4 or 5 teeth, with 4 or 5 teeth on first lateral carina. Pereopod 2 shorter than other pereopods. Gills: 2 arthrobranchs on maxilliped 3, pleurobranchs above pereopods 1–5, their apices directed anteriorly. Maximum cl. 21 mm (Pl. 8h). Slope (212–885 m). Tropical Atlantic (Caribbean), Western and Central Indo-Pacific. 6 species (Chan 1996: key to species).

Parapontophilus Christoffersen, 1988 Diagnosis. Rostrum ending in single apex, with 1 or 2 pairs of lateral teeth. Carapace middorsal line with 1 or 2 epigastric teeth, with or without 1 tooth on posterior half, lateral surface with 1 or 2 prominent spines. Pereopod 2 shorter than other pereopods; chelate, not reaching pereopod 1 merus distal margin. Gills: 1 arthrobranch on maxilliped 3, pleurobranchs above pereopods 1–5, their apices directed posteriorly. Maximum cl. 18 mm.

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Subtidal–abyssal (11–5852 m). Cosmopolitan except polar seas. 19 species. Parapontophilus is divided into two species-groups: the P. gracilis group of 11 species; and the P. modumanuensis group of seven species (Kim and Chan 2020; Komai 2008: key to species).

Philocheras Stebbing, 1900 Diagnosis. Rostrum short, blunt, with or without lateral teeth. Pereopod 2 shorter than other pereopods. Gills: 1 arthrobranch on maxilliped 3, pleurobranchs above pereopods 1–5, their apices directed posteriorly. Maximum cl. 14 mm. Intertidal–slope (0–732 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Western and Central IndoPacific, Tropical Eastern Pacific, Temperate Southern Africa, Temperate Australasia, Southern Ocean. 58 species (Han and Keesing 2018: key to species, distributions; Kim and Chan 2020).

Placopsicrangon Komai & Chan, 2009 Diagnosis. Rostrum acute, with dentate dorsal margin, without lateral teeth. Carapace middorsal line serrate, with 3 pairs of sharp, serrate lateral carinae. Pleon with simple longitudinal carinae. Eye strongly dorsoventrally flattened, incompletely fused with cephalothorax, cornea and eyestalk not separated, dorsal surface reflective. Pereopod 2 simple, thin, short, overreaching end of pereopod 1 ischium. Maximum cl. 25 mm. Abyssal (4412–4824 m). GOODS abyssal province: Central Pacific (Taiwan). 1 species (Komai and Chan 2009).

Pontocaris Bate, 1888 Diagnosis. Rostrum very short, apex simple or cleft, with 1 pair of inconspicuous lateral spines near base. Carapace middorsal line with 7 or more teeth, with 6–12 teeth on first lateral carina. Pereopod 2 shorter than other pereopods. Gills: 2 arthrobranchs on maxilliped 3, pleurobranchs above pereopods 1–5, their apices directed anteriorly. Maximum cl. 21 mm. Subtidal–slope (9–1424 m). Western and Central Indo-Pacific. 10 species (Chan 1996: key to species).

Pontophilus Leach, 1817 Diagnosis. Rostrum with 1 or 2 pairs of basal lateral teeth. Carapace middorsal line with epigastric tooth, usually with 1 or 2 teeth on posterior half, lateral surface with 3 or 5 prominent spines. Pereopod 2 shorter than other pereopods; chelate, not reaching pereopod 1 merus distal margin. Gills: 1 arthrobranch on maxilliped 3, pleurobranchs above pereopods 1–5, their apices directed posteriorly. Maximum cl. 15 mm. Subtidal–bathyal (7–2261 m). Temperate Northern and Tropical Atlantic. 3 species (Alves-Júnior et al. 2018). Pontophilus norvegicus is well known in the NE Atlantic and Mediterranean (De Grave and Diaz 2001).

Prionocrangon Wood-Mason, 1891 Diagnosis. Rostrum short, spiniform, slightly ascending. Eye reduced to small pointed processes; cornea absent. Pereopod 2

simple, rudimentary, thin, short, not reaching end of pereopod 1 merus (?). Maximum cl. 11 mm. Slope–bathyal (200–2556 m). Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate Australasia. 9 species. Prionocrangon was reviewed as a whole by Kim and Chan (2005) and in the Atlantic by Anker et  al. (2014). Its range extends to the Eastern Pacific (Hendrickx and Ayón Parente 2012).

Pseudopontophilus Komai, 2004 Diagnosis. Rostrum with 1 pair of lateral teeth. Carapace middorsal line with epigastric plus 1–4 spines in anterior 0.7 length, with 1 spine superior to antennal spine, postorbital carina with 3 spines. Pereopod 2 shorter than other pereopods; chelate, reaching pereopod 1 merus distal margin. Gills: 1 arthrobranch on maxilliped 3, pleurobranchs above pereopods 1–5, their apices directed posteriorly. Maximum cl. 7 mm. Slope (235–650 m). Temperate Northern Pacific, Central IndoPacific. 1 species (Komai 2004).

Rhynocrangon Zarenkov, 1965 Diagnosis. Rostrum short, broadly triangular. Carapace without submedian spines between middorsal line and hepatic spine, branchiostegal spine strong, flared sideways. Pleon with sculptured surface. Pereopod 2 subequal in length to the other pereopods. Pereopods 4, 5 dactyli laterally compressed, with spinule close to unguis. Maximum cl. 10 mm. Shelf (10–200 m). Temperate Northern Pacific. 3 species. Komai and Komatsu (2008), who reviewed the species, likened the shrimps to the rough pebbles among which they live.

Sabinea Ross, 1835 Diagnosis. Rostrum tapering, acute. Pleon dorsally carinate. Pereopod 2 simple, well developed, with broad articles, reaching beyond end of pereopod 1 merus. Maximum cl. 9 mm. Shelf–abyssal (48–3957 m). Temperate Northern and Tropical Atlantic, Central Indo-Pacific. 3 species. The only key to species (Chace 1984) included species no longer recognised. Sabinea ­hystrix (A. Milne-Edwards, 1881) is well known in the North Atlantic (Alves-Júnior et al. 2018).

Sclerocrangon G. O. Sars, 1883 Diagnosis. Rostrum short, broadly triangular. Carapace without submedian spines between middorsal line and hepatic spine, branchiostegal spine strong, flared sideways. Pleon with sculptured surface. Pereopod 2 subequal in length to the other pereopods. Maximum cl. 41 mm. Subtidal–abyssal (10–3950 m). Arctic, Temperate Northern Pacific. GOODS bathyal province: Northern N Pacific. 9 species (Komai and Amaoka 1991: key to 8 species; Komai and Matsuzaki 2016).

6 – Caridea – shrimps

Syncrangon Kim & Hayashi, 2003 Diagnosis. Rostrum acute, with dentate dorsal margin, without lateral teeth. Carapace middorsal line with 2 teeth, cardiac region with 2 transverse grooves, branchiostegal spine strong, nearly straight. Pleonites 3–6 with flattened middorsal carina with deep groove on each side. Pereopod 2 subequal in length to the other pereopods. Maximum cl. 7.6 mm. Intertidal–shelf (0–130 m). Temperate Northern Pacific. 2 species (Kim and Hayashi 2003).

Vercoia Baker, 1904 Diagnosis. Rostrum broad, tapering, T-shaped in cross-section. Pereopod 2 simple, rudimentary, thin, short, not reaching end of pereopod 1 merus. Maximum cl. 7 mm. Subtidal, shelf (3–167 m). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Australasia. 4 species (Ďuriš 1992; Kim and Fujita 2004; Komai 1995). References Alves-Júnior FdA, Viana GFS, Araújo MdsLCd, Souza-Filho JF (2018) New records of the family Crangonidae (Decapoda: Caridea) from Southwestern Atlantic. Zootaxa 4369, 46–62. doi:10.11646/zootaxa.4369.1.2 Anker A, Pachelle PPG, Tavares M (2014) Two new species and two new records of deep-water caridean shrimps from Brazil (Decapoda: Pandalidae, Palaemonidae, Crangonidae). Zootaxa 3815, 263–278. doi:10.11646/zootaxa.3815.2.6 Chace FA (1984) The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 2: families Glyphocrangonidae and Crangonidae. Smithsonian Contributions to Zoology 397, 1–63. doi:10.5479/si.00810282.397 Chan T-Y (1996) Crustacea Decapoda Crangonidae: revision of the three closely related genera Aegaeon Agassiz, 1846, Pontocaris Bate, 1888 and Parapontocaris Alcock, 1901. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 15. Mémoires du Muséum National d’Histoire Naturelle, Paris 168, 269–336. De Grave S, Diaz D (2001) Morphometric comparison between Mediterranean and Atlantic populations of Pontophilus norvegicus (Decapoda, Crangonidae). Hydrobiologia 449, 179–186. doi:10.1023/A:1017530111635 Ďuriš Z (1992) Revision of Vercoia Baker (Crustacea: Decapoda: Crangonidae). Invertebrate Taxonomy 6, 1437–1457. doi:10.1071/IT9921437 Han Q, Keesing JK (2018) A new species of Philocheras (Decapoda: Caridea: Crangonidae) from Western Australia, with a key to all species of this genus worldwide. Zootaxa 4382, 175–184. doi:10.11646/ zootaxa.4382.1.7 Han Q, Li X (2009) Neocrangon orientalis, a new caridean shrimp species (Crustacea, Decapoda, Crangonidae) from the East China Sea. Zootaxa 2050, 65–68. doi:10.11646/zootaxa.2050.1.4 Hayashi K-I (1999) Crustacea Decapoda: Revision of Pasiphaea sivado (Risso, 1816) and related species, with descriptions of one new genus and five new species (Pasiphaeidae). In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 20. Mémoires du Muséum National d’Histoire Naturelle, Paris 180, 267–302. Hendrickx ME (1993) Crustáceos decápodos del Pacífico mexicano. In Biodiversidad Marina y Costera de Mexico. (Eds Salazar-Vallejo SI, González NE) pp. 271–318. CONABIO, Chetumal. Hendrickx ME, Ayón Parente M (2012) First record of Prionocrangon Wood Mason & Alcock, 1891 (Crustacea: Decapoda: Caridea: Crango-

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nidae) in the East Pacific and description of a new species from western Mexico. Zootaxa 3205, 63–68. doi:10.11646/zootaxa.3205.1.5 Holthuis LB (1993) The recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda) with an appendix on the Order Amphionidacea. revised edn. Nationaal Natuuristorisch Museum, Leiden. Jarrin JRM, Shanks AL (2008) Ecology of a population of Lissocrangon stylirostris (Caridea: Crangonidae), with notes on the occurrence and biology of its parasite, Argeia pugettensis (Isopoda: Bopyridae). Journal of Crustacean Biology 28, 613–621. doi:10.1651/08-2976.1 Kim JN, Chan T-Y (2005) A revision of the genus Prionocrangon (Crustacea: Decapoda: Caridea: Crangonidae). Journal of Natural History 39, 1597–1625. doi:10.1080/00222930400016788 Kim JN, Chan T-Y (2020) Crangonid shrimps (Crustacea: Decapoda: Caridea) from Papua New Guinea. In: Corbari L, Chan T-Y, Ahyong ST (Eds) Deep-sea crustaceans from Papua New Guinea. Tropical Deep-Sea Benthos vol. 31. Mémoires du Muséum National d’Histoire Naturelle, Paris 213, 207–238. Kim JN, Fujita Y (2004) A new species of the genus Vercoia from Okinawa Island, Japan Crustacea, Decapoda, Caridea, Crangonidae, with descriptions of its zoeal stages. Journal of Natural History 38, 2013–2031. doi:10.1080/0022293032000140921 Kim JN, Hayashi K-I (2003) Syncrangon, a new crangonid genus, with redescriptions of S. angusticauda (De Haan) and S. dentata (Balss) (Crustacea, Decapoda, Caridea) from East Asian waters. Zoological Science 20, 669–682. doi:10.2108/zsj.20.669 Komai T (1995) Vercoia japonica, a new species of crangonid shrimp (Crustacea: Decapoda: Caridea) from Japan. Natural History Research 3, 123–132. Komai T (1997) Revision of Argis dentata and related species (Decapoda: Caridea: Crangonidae), with description of a new species from the Okhotsk Sea. Journal of Crustacean Biology 17, 135–161. doi:10.2307/1549469 Komai T (2004) A new genus and new species of Crangonidae (Crustacea, Decapoda, Caridea) from the southwestern Pacific. Zoosystema 26, 73–85. Komai T (2008) A world-wide review of species of the deep-water crangonid genus Parapontophilus Christoffersen, 1988 (Crustacea, Decapoda, Caridea), with descriptions of ten new species. Zoosystema 30, 261–332. Komai T (2012) A review of the western Pacific species of the crangonid genus Metacrangon Zarenkov, 1965 (Decapoda: Caridea), with descriptions of seven new species. Zootaxa 3468, 1–77. doi:10.11646/ zootaxa.3468.1.1 Komai T, Ahyong ST (2010) The crangonid shrimp genus Metacrangon (Crustacea: Decapoda: Caridea) from New Zealand, with descriptions of four new species. Journal of Natural History 45, 77–111. doi:10.1080 /00222933.2010.520823 Komai T, Amaoka K (1991) A new species of the genus Sclerocrangon from Urup Island, Kurile Islands and East of Hokkaido (Crustacea, Decapoda, Crangonidae). Proceedings of the Japanese Society of ­Systematic Zoology 44, 26–37. Komai T, Chan T (2009) New genus and species of Crangonidae (Decapoda: Caridea) with a large plate-like eye from the abyssal zone off Taiwan, northwestern Pacific. Journal of Crustacean Biology 29, 254–265. doi:10.1651/08-3080R.1 Komai T, Kim JN (2004) Shrimps of the crangonid genus Paracrangon Dana (Crustacea; Decapoda: Caridea) from the northwestern Pacific: taxonomic review and description of a new species from Japan. Scientia Marina 68, 511–536. doi:10.3989/scimar.2004.68n4511 Komai T, Komatsu H (2008) A new species of the crangonid shrimp genus Rhynocrangon (Crustacea, Decapoda, Caridea) from northern Japan. Bulletin of the National Museum of Nature and Science. Series A, Zoology 34, 183–195.

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Komai T, Matsuzaki K (2016) Two deep-sea decapod crustaceans collected off eastern Hokkaido, Japan: Sclerocrangon rex n. sp. (Caridea: Crangonidae) and Munidopsis verrilli Benedict, 1902 (Anomura: Munidopsidae). Zootaxa 4162, 92–106. doi:10.11646/zootaxa.4162.1.4 Komai T, Taylor J (2010) Three new species of the crangonid genus Metacrangon Zarenzov (Crustacea: Decapoda: Caridea) from Australia. Memoirs of Museum Victoria 67, 45–59. doi:10.24199/j. mmv.2010.67.05 Komai T, Takeuchi I, Takeda M (1996) Deep-sea shrimp (Crustacea: Decapoda: Caridea) from the Antarctic Sea collected during the JARE-35 cruise. Proceedings of the National Institute of Polar Research, Tokyo, Symposium on Polar Biology 9, 179–206. Komai T, Chen C, Watanabe HK (2018) Two new species of the crangonid genus Metacrangon Zarenkov, 1965 (Crustacea: Decapoda: Caridea) from the Okinawa Trough, Japan. Zootaxa 4410, 97–112. doi:10.11646/ zootaxa.4410.1.5 Kuris AM, Carlton JT (1977) Description of a new species, Crangon handi, and new genus Lissocrangon, of crangonid shrimps (Crustacea: Caridea) from the California coast, with notes on adaptation in body shape and coloration. Biological Bulletin 153, 540–559. doi:10.2307/1540605 Sedova N, Grigoriev S (2014a) Megalopa of Mesocrangon intermedia (Decapoda: Crangonidae) from the eastern part of the Sea of Okhotsk. Zoosystematica Rossica 23, 188–196. doi:10.31610/zsr/2014.23.2.188 Sedova N, Grigoriev S (2014b) Systematic position of Neocrangon communis (Decapoda, Crangonidae) based on the features of larval morphology. Zootaxa 3827, 559–575. doi:10.11646/zootaxa.3827.4.7 Taylor J, Collins DJ (2009) New records of the shrimp genus Lissosabinea (Caridea: Crangonidae) from Australia including descriptions of three new species and a key to world species. Memoirs of Museum Victoria 66, 175–187. doi:10.24199/j.mmv.2009.66.16 Zarenkov NA (1965) Revision of the genera Crangon Fabricius and Sclerocrangon G.O. Sars (Decapoda, Crustacea). Zoologicheskii Zhurnal 44, 1761–1775 [in Russian, translation published by Fisheries Research Board of Canada, 1970].

Glyphocrangonidae Smith, 1884 goblin shrimps Figure 6.24, Plate 8j–l The only genus is immediately recognisable in deep-sea trawls by its hard, calcareous and highly sculptured carapace. They are unique among decapods in having a locking mechanism between the last three pleonites, and between

Fig. 6.24.  Glyphocrangonidae. a, Glyphocrangon kapala Komai, 2004; b, pereopod 1.

pleonite 6 and the telson, which enables these elements to act as a rigid unit (Rice 1981). Diagnosis. Rostrum strong, dorsally flattened, tapering, with 1 or 2 pairs of lateral teeth. Carapace and pleon heavily armoured, sculptured with carinae and spines, with prominent cervical and hepatic grooves. Pereopod 1 prehensile, carpus closing against inner surface of propodus (Fig. 6.24b). Glyphocrangon A. Milne-Edwards, 1881 Maximum cl. 40 mm (Pl. 8j–l). Shelf– abyssal (153–6373 m). Cosmopolitan except polar seas. 94 species. In the Indo-Pacific, two species groups are recognised: the G. spinicauda species group (of 54 species) (Komai 2004b) and the G. caeca group (of eight species) (Komai 2006). Komai (2004b) provided descriptions of several species and a key to 54 species from the Indo-West Pacific. Holthuis (1971) distinguished 12 species in the Atlantic, some of which were further described by Komai (2004a). References Holthuis LB (1971) Biological Results of the University of Miami Deep-Sea Expeditions. 75. The Atlantic shrimps of the deep-sea genus Glyphocrangon A. Milne Edwards, 1881. Bulletin of Marine Science 21, 267–373. Komai T (2004a) Deep-sea shrimps of the genus Glyphocrangon A. MilneEdwards (Crustacea, Decapoda, Caridea, Glyphocrangonidae) from off southeastern coast of Brazil collected during the REVIZEE Program. Arquivos do Museu Nacional, Rio de Janeiro 62, 31–44. Komai T (2004b) A review of the Indo-West Pacific species of the genus Glyphocrangon A. Milne-Edwards, 1881 (excluding the G. caeca species group) (Crustacea: Decapoda: Caridea: Glyphocrangonidae). In: Marshall BA, Richer de Forges B (Eds), Tropical Deep-Sea Benthos, Vol. 23. Mémoires du Muséum National d’Histoire Naturelle, Paris 191, 375–610. Komai T (2006) Revision of the Glyphocrangon caeca species group (Crustacea, Decapoda, Glyphocrangonidae). In: Richer de Forges B, Justine J-L (Eds) Tropical Deep-Sea Benthos, Vol. 24. Mémoires du Muséum National d’Histoire Naturelle 193, 243–264. Rice AL (1981) The abdominal locking mechanism in the deep-sea shrimp genus Glyphocrangon (Decapoda, Glyphocrangonidae). Crustaceana 40, 316–319. doi:10.1163/156854081X00804

Superfamily Nematocarcinoidea Smith, 1884 One of the five nematocarcinoid families is entirely freshwater. Xiphocarididae Ortmann, 1895 comprise a single genus, Xiphocaris von Martens, 1872 (Guerao et al. 2014). Two species occur in streams on Caribbean islands (Juarrero de Varona 1993) one of which is well studied (e.g. Ocasio-Torres et al. 2015). The four other families range from subtidal to bathyal depths. The superfamily was considered polyphyletic by Christoffersen (1990), a conclusion supported by molecular data (Baeza et al. 2014). Diagnosis. Mandibular molar blunt, subtruncate with ridged grinding surface (Figs 6.26c, 6.27h). Pereopods 1, 2

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dissimilar, first stronger and heavier though sometimes shorter than second. Pereopod 2 carpus entire. Exopod present on maxilliped 3, pereopods 1–5. Strap-like epipod on at least pereopods 1–3 (Fig. 6.29i). References Baeza JA, Bauer RT, Okuno J, Thiel M (2014) Molecular phylogeny of hinge-beak shrimps (Decapoda: Caridea: Rhynchocinetes and Cinetorhynchus) and allies: a formal test of familiar and generic monophyly using a multilocus phylogeny. Zoological Journal of the Linnean Society 172, 426–450. doi:10.1111/zoj.12173 Christoffersen ML (1990) A new superfamily classification of the Caridea (Crustacea: Pleocyemata) based on phylogenetic pattern. Journal of Zoological Systematics and Evolutionary Research 28, 94–106. doi:10.1111/j.1439-0469.1990.tb00369.x Guerao G, Reuschel S, Anger K, Schubart Christoph D (2014) On the presumed phylogenetic position of the Xiphocarididae (Decapoda, Caridea) based on the larval morphology of Xiphocaris elongata. In: Yeo D, Cumberlidge N, Klaus S (Eds) Advances in freshwater decapod systematics and biology. Crustaceana Monographs 19, 233–244. Juarrero de Varona A (1993) Nueva especie del género Xiphocaris (Crustacea: Atyidae) de Cuba. Poeyana 440, 1–12. Ocasio-Torres ME, Crowl TA, Sabat AM (2015) Allometric differences between two phenotypes of the amphidromous shrimp Xiphocaris elongata. Journal of Crustacean Biology 35, 747–752. doi:10.1163/​ 1937240X-00002372

Eugonatonotidae Chace, 1937 Figure 6.25, Plate 9a The two deep-water species, notable for the longitudinal carapace ridges, belong in a single genus. Diagnosis. Rostrum strongly dentate. Carapace with 3 longitudinal lateral ridges. Mandible without incisor. Eugonatonotus Schmitt, 1926 Maximum cl. 42 mm (Pl. 9a). Shelf, slope (53–914 m). Tropical W Atlantic, Central Indo-Pacific, Temperate SW Australia. 2 species (Boone 1927: figure of type species as Gonatonotus crassus A. Milne Edwards, 1881; Chan and Yu 1991). The genus Galatheacaris Vereschaka, 1997 is a junior synonym of Eugonatonotus, being based on the larva of E. chacei Chan & Yu, 1991. References Boone L (1927) Scientific results of the first oceanographic expedition of the “Pawnee”, 1925. Crustacea from the tropical east American seas. Bulletin of the Bingham Oceanographic Collection 1, 1–147. Chan T-Y, Yu H-P (1991) Eugonatonotus chacei sp. nov., second species of the genus (Crustacea, Decapoda, Eugonatonotidae). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série 13, 143–152.

Lipkiidae Burukovsky, 2012 Figure 6.26, Plate 9b The family is confined to two species from southern latitudes (D’Udekem D’Acoz and De Grave 2018).

Fig. 6.25.  Eugonatonotidae. Eugonatonotus crassus (A. Milne Edwards, 1881).

Diagnosis. Rostrum finely dentate with basal dorsal spines only. Maxilliped 2 last article a narrow strip on submesialdistal margin of broader penultimate article. Pereopods 3–5 1.5 times length of pereopod 1, carpus shorter than propodus, articulation between ischium and merus not thickened. Fresnerhynchus d’Udekem d’Acoz & De Grave, 2018 Diagnosis. Thoracic sternum area between pereopods 2–5 uneven, not forming bifid plate. Pleonite 4 posteroventrally not produced into tooth (Fig. 6.26b). Pereopods 3–5 dactyli with spiniform setae on flexor margin (Fig. 6.26e). Maximum cl. 39 mm. Slope (1889 m). Southern Ocean (seamount near Crozet Is). 1 species (D’Udekem D’Acoz and De Grave 2018).

Lipkius Yaldwyn, 1906 omega prawn Diagnosis. Thoracic sternum area between pereopods 2–5 forming bifid plate. Pleonite 4 posteroventrally produced into tooth (Fig.  6.26a). Pereopods 3–5 dactyli without spiniform setae on flexor margin (Fig. 6.26f). Maximum cl. 33 mm (Pl. 9b). Slope (350–1700 m). Temperate Australasia (366–1700 m). Maximum cl. 33 mm. 1 species (Yaldwyn 1960). The only species is taken in fish trawls in New Zealand, often with species of Nematocarcinus (Ahyong et al. 2011; Webber et al. 1990) but occurs in south-eastern Australia also (Poore 2004). References Ahyong ST, Booth J, Bruce NL, Loerz A-N, Naylor JR, et al. (2011) Phylum Arthropoda. In: Tracey DM, Anderson OF, Naylor JR (Eds), A guide to common deepsea invertebrates in New Zealand waters. Third Edition. New Zealand Aquatic Environment and Biodiversity Report 86, 159–222. D’Udekem D’Acoz C, De Grave S (2018) A new genus and species of largebodied caridean shrimp from the Crozet Islands, Southern Ocean (Crustacea, Decapoda, Lipkiidae) with a checklist of Antarctic and sub-Antarctic shrimps. Zootaxa 4392, 201–240. doi:10.11646/ zootaxa.4392.2.1

112

Marine Decapod Crustacea

Fig. 6.26.  Lipkiidae. Habitus: a, Lipkius holthuisi Yaldwyn, 1960. Pleonites 4–6: b, Fresnerhynchus. Mandible 2: c, Fresnerhynchus. Maxilliped 2: d, Fresnerhynchus. Pereopod 3, dactylus: e, Fresnerhynchus; f, Lipkius. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Webber WR, Fenaughty CM, Clark MR (1990) A guide to some common offshore shrimp and prawn species of New Zealand. New Zealand Fisheries Occasional Publication 6, 1–42. Yaldwyn JC (1960) Crustacea Decapoda Natantia from the Chatham Rise: a deep water bottom fauna from New Zealand. Bulletin – New Zealand Department of Scientific and Industrial Research 139, 13–53.

Nematocarcinidae Smith, 1884 Figure 6.27, Plate 9c Nematocarcinids are deep-water demersal shrimps notable for their extremely long slender pereopods (Burukovsky 2012; Chace 1986). Unfortunately, trawled specimens often

fare poorly, especially the legs and rostrum, so any specimens found intact should be treasured. The family occurs in all ocean basins and in Antarctic waters (Raupach et al. 2010). Most species are in one genus, Nematocarcinus; other genera contain only one or two species each. The key to genera is derived from those of Burukovsky (2012, 2013) who reviewed the systematics and distribution of the family, in Russian with English summaries. Diagnosis. Rostrum finely dentate with dorsal spines to apex, spines often movable (Fig. 6.27a–g). Mandible deeply bifurcate, with palp (Fig. 6.27h). Maxilliped 2 last article a narrow strip on submesial-distal margin of broader penultimate article (Fig.  6.27i). Pereopods 3–5 twice length of pereopod 1, carpus several times length of propodus, articulation between ischium and merus thickened (Fig. 6.27n).

Key to genera of Nematocarcinidae 1. – 2. – 3. – 4. –

Pleonite 6 with distoventral rows organ (pair of prominences posterior to longitudinal rows of setae: Fig. 6.27j). Rostrum dorsal margin and postrostral crest with only mobile teeth (Fig. 6.27d, e)�������� Nematocarcinus Pleonite 6 without distoventral organ. Rostrum dorsal margin with fixed teeth; postrostral crest with mobile or fixed teeth�������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Rostrum ventral margin with mobile teeth (Fig. 6.27c)������������������������������������������������������������������������������������� Macphersonus Rostrum ventral margin with fixed teeth (Fig. 6.27b, f, g)��������������������������������������������������������������������������������������������������������3 Rostrum dorsal margin with fixed teeth; postrostral crest with mobile teeth (Fig. 6.27g)���������������������Segonzackomaius Rostrum dorsal margin and postrostral crest with fixed teeth (Fig. 6.27a, f)������������������������������������������������������������������������4 Telson exceeding uropods, dorsal surface with long median spiniform setae (Fig. 6.27l). Carapace with curved keel connecting subcardiac and hepatic areas (Fig. 6.27f)�������������������������������������������������������������� Nigmatullinus Telson not exceeding distal margin of uropods, with 2 rows of small dorsolateral spiniform setae (Fig. 6.27k). Carapace smooth (Fig. 6.27a)������������������������������������������������������������������������������������������������������� Lenzicarcinus

6 – Caridea – shrimps

113

Fig. 6.27.  Nematocarcinidae. Carapace, rostrum: a, b, Lenzicarcinus (detail); c, Macphersonus; d, Nematocarcinus subtilis Burukovsky, 2000; e, N. productus Bate, 1888; f, Nigmatullinus; g, Segonzackomaius. Mandible: h, Nematocarcinus. Maxilliped 2: i, Nematocarcinus. Pereonite 6 sternum, longitudinal setal rows and paired dorsal organ (grey): j, Nematocarcinus. Telson, uropod: k, Lenzicarcinus; l, Nigmatullinus; m, Segonzackomaius. Pereopod 3: n, Nematocarcinus.

Lenzicarcinus Burukovsky, 2005 Diagnosis. Rostrum dorsal and ventral margins with fixed teeth; postrostral crest with fixed teeth. Telson not exceeding uropod, dorsal surface with 2 rows of dorsolateral spiniform setae. Maximum cl. 12 mm. Bathyal (2000 m). GOODS bathyal province: S Atlantic. 1 species (Burukovsky 2005).

Macphersonus Burukovsky, 2012 Diagnosis. Rostrum dorsal and ventral margins with mobile teeth; postrostral crest with mobile teeth. Telson not exceeding uropod, dorsal surface with 2 rows of dorsolateral spiniform setae. Maximum cl. 21 mm. Slope (730–1650 m). Western Indo-Pacific. 1 species (Burukovsky 2012).

Nematocarcinus A. Milne-Edwards, 1881 Diagnosis. Rostrum dorsal margin with mobile teeth; ventral margin with fixed teeth, or without teeth; postrostral crest with mobile teeth. Pleonite 6 with distoventral rows organ (pair of prominences posterior to longitudinal rows of setae). Telson not

exceeding uropod, dorsal surface with 2 rows of dorsolateral spiniform setae. Maximum cl. 30 mm (Pl. 9c). Shelf–abyssal (203–5477 m). Cosmopolitan. 43 species (­Burukovsky 2013: tabular key to species, distributions).

Nigmatullinus Burukovsky, 1991 Diagnosis. Carapace with curved keel connecting subcardiac and hepatic areas. Rostrum dorsal margin and ventral margins with fixed teeth; postrostral crest with fixed teeth. Telson exceeding uropod, dorsal surface with long median spiniform setae. Maximum cl. 40 mm. Bathyal, abyssal (2000–4160 m). GOODS bathyal provinces: Northern N Atlantic, SE Pacific ridges, Indian, W Pacific. 1 species (Burukovsky 1991; Crosnier and Forest 1973).

Segonzackomaius Burukovsky, 2011 Diagnosis. Rostrum dorsal and ventral margins with fixed teeth; postrostral crest with mobile teeth. Telson not exceeding uropod, dorsal surface with 2 rows of dorsolateral spiniform setae. Maximum cl. 29 mm. Bathyal, abyssal (2330–4446 m). GOODS bathyal province: SE Pacific ridges. 2 species (Burukovsky 2011).

114

Marine Decapod Crustacea

References Burukovsky RN (1991) Shrimps of the family Nematocarcinidae (Decapoda, Caridea) from the western part of the Indian Ocean [in Russian]. Zoologicheskij Zhurna 70, 39–46. Burukovsky RN (2005) Systematics of shrimps of the family Nematocarcinidae (Crustacea, Decapoda) from collections of the Berlin Natural History Museum and Senkenberg Museum of Natural History and Research Institute (Germany) [in Russian]. Zoologicheskij Zhurnal 84, 569–579. Burukovsky RN (2011) A new shrimp of the family Nematocarcinidae (Decapoda, Caridea). Zoologicheskij Zhurnal 90, 1–9. Burukovsky RN (2012) Deep sea shrimps of the family Nematocarcinidae (history of study, systematics, distribution, and biology) Prospekt Nauki, St Petersberg. Burukovsky RN (2013) Shrimps of the family Nematocarcinidae Smith, 1884 (Crustacea, Decapoda, Caridea) from Taiwan and the Philippines collected by the TAIWAN, PANGLAO 2005 and AURORA expeditions in the western Pacific. In: Ahyong ST, Chan T-Y, Corbari L, Ng PKL (Eds), Tropical Deep-Sea Benthos Vol. 27. Mémoires du Muséum National d’Histoire Naturelle, Paris 204, 155–189. Chace FA (1986) The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 4: families Oplophoridae and Nematocarcinidae. Smithsonian Contributions to Zoology 432, 1–82. doi:10.5479/si.00810282.432 Crosnier A, Forest J (1973) Les crevettes profondes de l’Atlantique oriental tropical. Faune Tropicale 19, 1–409. Raupach MJ, Thatje S, Dambach J, Rehm P, Misof B, et  al. (2010) Genetic homogeneity and circum-Antarctic distribution of two benthic shrimp species of the Southern Ocean, Chorismus antarcticus and Nematocarcinus lanceopes. Marine Biology 157, 1783–1797. doi:10.1007/s00227-010-1451-3

Rhynchocinetidae Ortmann, 1890 hinge-beak shrimps Figure 6.28, Plate 9d–f

Hinge-beak shrimps are colourful species from shallow water, with, as their name suggests a movable rostrum. They have been much studied because they exhibit remarkable ecological and behavioural diversity (Bauer 2004). The males of some species are larger than females, with enlarged chelipeds, which they use during intrasexual competition for receptive females (Thiel et al. 2010). The family and its two genera were found to be monophyletic by Baeza et  al. (2014). The most recent published papers do not include keys to all species. Diagnosis. Rostrum grossly dentate, hinged with carapace. Cinetorhynchus Holthuis, 1995 Diagnosis. Carapace median carina with 3 teeth; without supraorbital spine (Fig.  6.28a). Pereonite 5 with posterolateral spine (Fig. 6.28b). Maximum cl. 17 mm (Pl. 9d). Intertidal–shelf (0–67 m). Temperate Northern and Tropical Atlantic, Indo-West Pacific. 12 species (Okuno 1997: rediagnosis, key to 6 species).

Rhynchocinetes H. Milne Edwards, 1837 Diagnosis. Carapace median carina with 2 teeth; with supraorbital spine. Pereonite 5 without posterolateral spine (Fig. 6.28c). Maximum cl. 13 mm (Pl. 9e, f). Intertidal–shelf (0–220 m). Western and Central Indo-Pacific, Temperate South America, Temperate Australasia. 15 species (Okuno 1997: rediagnosis). References Baeza JA, Bauer RT, Okuno J, Thiel M (2014) Molecular phylogeny of hinge-beak shrimps (Decapoda: Caridea: Rhynchocinetes and Cinetorhynchus) and allies: a formal test of familiar and generic monophyly using a multilocus phylogeny. Zoological Journal of the Linnean Society 172, 426–450. doi:10.1111/zoj.12173 Bauer RT (2004) Remarkable shrimps. Adaptations and natural history of the caridean shrimps. University of Oklahoma Press, Norman. Okuno J (1997) Crustacea Decapoda: review on the genus Cinetorhynchus Holthuis, 1995 from the Indo-West Pacific (Caridae: Rhynchocinetidae). In Les fonds meubles des lagons de Nouvelle-Calédonie (sédimentologie, benthos). (Ed. Richer de Forges B) pp. 31–58. ORSTOM Editions, Paris. Thiel M, Chak STC, Dumont CP (2010) Male morphotypes and mating behavior of the dancing shrimp Rhynchocinetes brucei (Decapoda: Caridea). Journal of Crustacean Biology 30, 580–588. doi:10.1651/​ 09-3272.1

Superfamily Oplophoroidea Dana, 1852

Fig. 6.28.  Rhynchocinetidae. Cinetorhynchus: a, rostrum, anterior carapace; b, pleonites 4–6. c, Rhynchocinetes australis Hale, 1941.

Oplophoroids are deep-water shrimps, most being mesopelagic or bathypelagic but some suspected to be hyperbenthic. Some have been observed at almost 7000 m depth (Swan et al. 2021). Mesopelagic species are vertical migrators and all are capable of bioluminescence (Herring 1985).

6 – Caridea – shrimps

The superfamily is represented in all oceans, including subarctic and subantarctic waters (Pohle et  al. 1992; Wasmer 1986). Chace (1986) treated the two families now recognised as one. He diagnosed all the genera, and those species occurring in the Western Pacific. This foundation study has been supplemented by surveys of species from, for example, the Indian Ocean (Crosnier 1987), the tropical Atlantic (Pequegnat and Wicksten 2006) and the south-western Atlantic (Alves-Júnior et al. 2019). Separation of Oplophoroidea into two families is based on the form of the mandible, integument, the presence or absence of cuticular photophores, eye size, and molecular differences (Chan et al. 2010; Wong et al. 2015; Vereshchaka et al. 2021). Oplophoridae s.s. tend to live in shallower water than Acanthephyridae. Chan et  al. (2010) found that the two families were not sister taxa. Vereshchaka et al. (2021) provided a key to all genera of Oplophoroidea. Diagnosis. Pereopods 1, 2 similar, both chelate. Pereopod 2 carpus entire. Exopods present on maxilliped 3, pereopods 1–5. Epipod present on all pereopods, terminating in naked appendix extending vertically into branchial chamber posterior to corresponding pleurobranch (Fig. 6.29i). References Alves-Júnior FdA, Silva EdS, De Araújo MdsLC, Cardoso I, Bertrand A, et al. (2019) Taxonomy of deep-sea shrimps of the superfamily Oplophoroidea Dana 1852 (Decapoda: Caridea) from Southwestern Atlantic. Zootaxa 4613, 401–442. doi:10.11646/zootaxa.4613.3.1 Chace FA (1986) The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 4: families Oplophoridae and Nematocarcinidae. Smithsonian Contributions to Zoology 432, 1–82. doi:10.5479/si.00810282.432 Chan T-Y, Lei HC, Li CP, Chu KH (2010) Phylogenetic analysis using rDNA reveals polyphyly of Oplophoridae (Decapoda: Caridea). Invertebrate Systematics 24, 172–181. doi:10.1071/IS09049 Crosnier A (1987) Oplophoridae (Crustacea Decapoda) récoltés de 1971 à 1982 par les navires français dans l’océan Indien occidental sud. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 9, 695–726. Herring PJ (1985) Bioluminescence in the Crustacea. Journal of Crustacean Biology 5, 557–573. doi:10.2307/1548235

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Pequegnat LH, Wicksten MK (2006) Oplophorid shrimps (Decapoda: Caridea: Oplophoridae) in the Gulf of Mexico and Caribbean Sea from the collections of the research vessels Alaminos, Oregon and Oregon II. Crustacean Research 35, 92–107. doi:10.18353/ crustacea.35.0_92 Pohle G, Kenchington TJ, Halliday RG (1992) Potentially exploitable deepwater resources off Atlantic Canada. Canadian Technical Report of Fisheries and Aquatic Sciences 1843, 1–85. Swan JA, Jamieson AJ, Linley TD, Yancey PH (2021) Worldwide distribution and depth limits of decapod crustaceans (Penaeoidea, Oplophoroidea) across the abyssal-hadal transition zone of eleven subduction trenches and five additional deep-sea features. Journal of Crustacean Biology 41, doi:10.1093/jcbiol/ruaa102 Vereshchaka A, Kulagin D, Lunina A (2021) A new shrimp genus (Crustacea: Decapoda) from the deep Atlantic and an unusual cleaning mechanism of pelagic decapods. Diversity 13, 536. doi:10.3390/ d13110536 Wasmer RA (1986) Pelagic shrimps of the family Oplophoridae (Crustacea: Decapoda) from the Pacific sector of the southern ocean: USNS Eltanin cruises 10, 11, 14–16, 19–21, 24 and 25. In: Biology of the Antarctic Seas 17. Antarctic Research Series 44, 29–68. Wong JM, Pérez-Moreno JL, Chan T-Y, Frank TM, Bracken-Grissom HD (2015) Phylogenetic and transcriptomic analyses reveal the evolution of bioluminescence and light detection in marine deep-sea shrimps of the family Oplophoridae (Crustacea: Decapoda). Molecular Phylogenetics and Evolution 83, 278–292. doi:10.1016/j.ympev.2014.11.013

Acanthephyridae Bate, 1888 Figure 6.29, Plate 9g–k Acanthephyridae comprise eight genera of mesopelagic deepsea shrimps (Vereshchaka et al. 2021) previously included in Oplophoridae until Chan et al. (2010) differentiated the two families. Acanthephyrids can be dominant members of the mesopelagic community (Burdett et  al. 2017). One genus, Sclerodora Vereshchaka, Kulagin & Lunina, 2021, and one species have been added since the comprehensive review of Chace (1986) and completion of this work. It appears to be indistinguishable from Hymenodora. Diagnosis. Mandibles dissimilar, molar distinct from incisor, with distal surface triangular on right member, compressed, sublinear on left (Fig. 6.29h). Without cuticular photophores on body and appendages.

Key to genera of Acanthephyridae 1. – 2. – 3. – 4.

Pleonites not carinate (Fig. 6.29b)��������������������������������������������������������������������������������������������������������������������������������������������������2 Most pleonites carinate�������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Rostrum laterally compressed, a subtriangular vertical lamella. Carapace with blunt, sinuous carina extending posteriorly from behind orbit to near posterior margin (Fig. 6.29b)������������������������������������������������ Ephyrina Rostrum short, crest-like. Carapace without uninterrupted lateral carina (Fig. 6.29d)����������������������������������Hymenodora Carapace with hepatic spine (Fig. 6.29e)��������������������������������������������������������������������������������������������������������������������� Kemphyra Carapace without hepatic spine�����������������������������������������������������������������������������������������������������������������������������������������������������4 Carapace without uninterrupted lateral carina extending from near orbit to near posterior margin (Fig. 6.29c)������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������5

116

– 5. – 6. –

Marine Decapod Crustacea

Carapace with uninterrupted lateral carina extending from near orbit to near posterior margin (Fig. 6.29f, g)�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������6 Carapace with or without short lateral carina. Pleonite 3 tooth less than half as long as pleonite 4, pleonites 4–6 usually with posteromedian tooth (Fig. 6.29a)���������������������������������������������������������������������������������������� Acanthephyra Carapace without lateral carina. Pleonite 3 tooth more than ¾ as long as pleonite 4, pleonites 4–6 without posteromedian tooth (Fig. 6.29c)��������������������������������������������������������������������������������������������������� Heterogenys Carapace with uninterrupted lateral carina extending from near orbit to near posterior margin; dorsal margin not denticulate (Fig. 6.29f)�������������������������������������������������������������������������������������������������������������������� Meningodora Carapace with 2 or more lateral longitudinal carinae extending to posterior half; dorsal margin denticulate over nearly entire length (Fig. 6.29g)��������������������������������������������������������������������������������������������������������������������Notostomus Acanthephyra A. Milne-Edwards, 1881

Ephyrina Smith, 1885

Diagnosis. Rostrum narrow, blade-like; with at least as many dorsal as ventral teeth. Carapace without uninterrupted lateral carina extending from near orbit to near posterior margin; dorsal margin not denticulate; hepatic groove posterior margin without carina. Pleonites 3–6 carinate, 3 with posterior tooth, 4–6 usually with posteromedian tooth. Maximum cl. 55 mm (Pl. 9g, h). Mesopelagic (0–6890 m). Cosmopolitan, including subpolar oceans. 27 species (Chace 1986: key to species, distributions).

Diagnosis. Rostrum laterally compressed, subtriangular vertical lamella; unarmed. Carapace with blunt, sinuous carina extending posteriorly from behind orbit to near posterior margin; dorsal margin not denticulate; hepatic groove posterior margin with oblique carina. Pleonites not carinate. Maximum cl. 27 mm (Pl. 9i). Mesopelagic, bathypelagic (497–1682 m). Pelagic regions: Atlantic, Indian, Central and W Pacific. 6 species (Chace 1986: key to species, distributions).

Fig. 6.29.  Acanthephyridae. Habitus/carapace: a, Acanthephyra quadrispinosa Kemp, 1930; b, Ephyrina benedicti Smith, 1885; c, Heterogenys; d, Hymenodora; e, Kemphyra; f, Meningodora; g, Notostomus auriculatus Barnard, 1950. Mandibles: h, Acanthephyra. Epipod, arthrobranch: i, Notostomus.

6 – Caridea – shrimps

Heterogenys Chace, 1986 Diagnosis. Rostrum narrow, blade-like; with fewer dorsal than ventral teeth. Carapace without uninterrupted lateral carina; dorsal margin not carinate; hepatic groove posterior margin without carina. Pleonites 3–6 carinate, 3 with long posterior tooth, 4–6 without posteromedian tooth. Maximum cl. 23 mm (Pl. 9j). Bathyal, abyssal (2000–5060 m). GOODS bathyal provinces: Northern N Pacific, New Zealand/Kermadec, Subantarctic (S Australia), Cocos Plate, Indian, W Pacific. 2 species (Chace 1986: key to species, distributions).

Hymenodora G.O. Sars, 1877 Diagnosis. Rostrum short, crest-like; dentate dorsally only. Carapace without uninterrupted lateral carina; dorsal margin not denticulate; hepatic groove posterior margin without carina. Pleonites not carinate. Maximum cl. 11 mm. Mesopelagic, bathypelagic (0–5610 m). Pelagic regions: Atlantic, Indian, Central and W Pacific, E Pacific. 4 species (Chace 1986: key to species, distributions).

Kemphyra Chace, 1986 Diagnosis. Rostrum triangular in lateral view; with more dorsal than ventral teeth. Carapace with 3 carinae over posterior half; dorsal margin not denticulate; with hepatic spine; hepatic groove posterior margin without carina. Pleonites 1–6 carinate, 3–6 with median posterior tooth. Maximum cl. 26 mm. Slope, bathyal (1000–3200 m). Temperate Northern Atlantic, Eastern Indo-Pacific, Temperate Southern Africa. 1 species (Chace 1986).

Meningodora Smith, 1882 Diagnosis. Rostrum triangular in lateral view; with more dorsal than ventral teeth. Carapace with uninterrupted lateral carina extending from near orbit to near posterior margin; dorsal margin not denticulate; hepatic groove posterior margin with oblique carina. Pleonites at least 3–6 carinate, 3–6 or 4–6 with median posterior tooth. Maximum cl. 26 mm. Mesopelagic (840–2985 m). Cosmopolitan in temperate and tropical oceans. 6 species (Chace 1986: key to 5 species, distributions).

Notostomus A. Milne-Edwards, 1881 Diagnosis. Rostrum narrow, blade-like; with more dorsal than ventral teeth. Carapace with 2 or more lateral longitudinal carinae

117

extending to posterior half; dorsal margin denticulate over nearly entire length; hepatic groove posterior margin with oblique carina. Pleonites 1–6 carinate, 3–6 with median posterior tooth. Maximum cl. 60 mm (Pl. 9k). Mesopelagic (0–3932 m). Cosmopolitan in temperate and tropical oceans. 9 species (Chace 1986: key to 8 species, distributions). References Burdett EA, Fine CD, Sutton TT, Cook AB, Frank TM (2017) Geographic and depth distributions, ontogeny, and reproductive seasonality of decapod shrimps (Caridea: Oplophoridae) from the northeastern Gulf of Mexico. Bulletin of Marine Science 93, 743–767. doi:10.5343/ bms.2016.1083 Chace FA (1986) The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 4: families Oplophoridae and Nematocarcinidae. Smithsonian Contributions to Zoology 432, 1–82. doi:10.5479/si.00810282.432 Chan T-Y, Lei HC, Li CP, Chu KH (2010) Phylogenetic analysis using rDNA reveals polyphyly of Oplophoridae (Decapoda: Caridea). Invertebrate Systematics 24, 172–181. doi:10.1071/IS09049 Vereshchaka A, Kulagin D, Lunina A (2021) A new shrimp genus (Crustacea: Decapoda) from the deep Atlantic and an unusual cleaning mechanism of pelagic decapods. Diversity 13, 536. doi:10.3390/d13110536

Oplophoridae Dana, 1852 Figure 6.30, Plate 9l–n Three genera remain in Oplophoridae s.s.  – the majority formerly included now belong in Acanthephyridae. Both families are bioluminescent but only oplophorids bear superficial cuticular photophores on the surface of the body and appendages. Photophores comprise light-emitting bioluminescent cells plus reflectors, lenses and filters capable of altering the quality of the light emitted (Nowel et al. 1998; Wong et al. 2015: Fig. 1B). Oplophorid phylogeny, taxonomy and evolution have been explored by a combination of morphological and genetic methods (Chan et al. 2010; Lunina et al. 2019; Wong et al. 2015). Lunina et al. (2019) provided a key to genera and species, complementing those of Chace (1986). Only one species has been added since Chace’s review (Sha and Wang 2015). Diagnosis. Mandibles similar, molar weakly separate from incisor, consisting of channel flanked by thin walls opposing similar structure on other of pair (Fig. 6.30d), or reduced to small process (Fig.  6.30e). Superficial cuticular photophores on body and appendages.

Key to genera of Oplophoridae 1. –

Carapace moderately chitinised, suboval in cross-section. Pleonites 3, 4 without strong dorsomedial spine. Pleonite 6 about twice or more as long as pleonite 5 (Fig. 6.30c)����������������������������������������������������������������������� Systellaspis Carapace strongly chitinised, subtriangular in cross-section. Pleonites 3, 4 with strong dorsomedial spine (at least half of somite length). Pleonite 6 shorter or up to 1.5 times as long as pleonite 5 (Fig. 6.30a, b)���������������������2

118

Marine Decapod Crustacea

Fig. 6.30.  Oplophoridae. Carapace, pleon: a, Janicella; b, Oplophorus; c, Systellaspis. Mandibles: d, Janicella; e, Systellaspis. Telson: f, Janicella (apex); g, Oplophorus.

2. –

Pleonites 2–4 with strong dorsomedial spines (Fig. 6.30a). Telson with spinose endpiece, with pair of long lateral spiniform setae (Fig. 6.30f)�������������������������������������������������������������������������������������������������������������������������������Janicella Pleonites 3–5 with strong dorsomedial spines (Fig. 6.30b). Telson apex acute, without lateral spiniform setae (Fig. 6.30g)������������������������������������������������������������������������������������������������������������������������������������������������������� Oplophorus Janicella Chace, 1986

Diagnosis. Carapace strongly chitinised, subtriangular in crosssection. Pleonites 3, 4 with dorsomedial spine as least as long as somite; pleonite 6 ~1.5 times as long as pleonite 5. Telson with spinose endpiece, with pair of long lateral spiniform setae. Mandibular molar reduced to small subtriangular excavated lobe, not opposable (Fig. 6.30d). Maximum cl. 11 mm. Mesopelagic (migrating up to 30 m from 3176 m). Widespread in tropical oceans, except E Pacific. 1 species (Chace 1986).

Oplophorus H. Milne Edwards, 1837 Diagnosis. Carapace strongly chitinised, subtriangular in crosssection. Pleonites 3, 4 with dorsomedial spine as least as long as somite; pleonite 6 shorter than pleonite 5. Telson apex simple. Mandibular molar a deep channel flanked by high thin walls, opposable. Maximum cl. 21 mm (Pl. 9l, m). Mesopelagic (migrating to near surface). Cosmopolitan in temperate and tropical oceans. 4 species (Lunina et al. 2019: key to species).

Systellaspis Bate, 1888 Diagnosis. Carapace moderately chitinised, suboval in cross-section. Pleonites 3, 4 without strong dorsomedial spine; pleonite 6 about twice or more as long as pleonite 5. Telson with spinose endpiece, with pair of long lateral spiniform setae. Mandibular

molar a deep channel flanked by high thin walls, opposable (Fig. 6.30e). Maximum cl. 36 mm (Pl. 9n). Shelf–abyssal, mesopelagic-benthic (101–5025 m). Cosmopolitan in temperate and tropical oceans. 11 species (Lunina et  al. 2019: key to species, species-groups). References Chace FA (1986) The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 4: families Oplophoridae and Nematocarcinidae. Smithsonian Contributions to Zoology 432, 1–82. doi:10.5479/si.00810282.432 Chan T-Y, Lei HC, Li CP, Chu KH (2010) Phylogenetic analysis using rDNA reveals polyphyly of Oplophoridae (Decapoda: Caridea). Invertebrate Systematics 24, 172–181. doi:10.1071/IS09049 Lunina AA, Kulagin DN, Vereshchaka AL (2019) Oplophoridae (Decapoda: Crustacea): phylogeny, taxonomy and evolution studied by a combination of morphological and molecular methods. Zoological Journal of the Linnean Society 186, 213–232. doi:10.1093/zoolinnean/zly039 Nowel MS, Shelton PMJ, Herring PJ (1998) Cuticular photophores of two decapod crustaceans, Oplophorus spinosus and Systellaspis debilis. Biological Bulletin 195, 290–307. doi:10.2307/1543141 Sha Z-L, Wang Y-L (2015) A new deep-sea species of the genus Systellaspis (Decapoda, Caridea, Oplophoridae) in the western Pacific. Crustaceana 88, 1181–1192. doi:10.1163/15685403-00003484 Wong JM, Pérez-Moreno JL, Chan T-Y, Frank TM, Bracken-Grissom HD (2015) Phylogenetic and transcriptomic analyses reveal the evolution of bioluminescence and light detection in marine deep-sea shrimps of the family Oplophoridae (Crustacea: Decapoda). Molecular Phylogenetics and Evolution 83, 278–292. doi:10.1016/j.ympev.2014.11.013

6 – Caridea – shrimps

119

Superfamily Palaemonoidea Rafinesque, 1815

Palaemonidae Rafinesque, 1815

Three of the four palaemonoid families are from predominantly freshwater environments and not dealt with further here. Desmocarididae Borradaile, 1915 comprise one genus, Desmocaris Sollaud, 1911, with two species from West African forest streams (Powell 1977). Euryrhynchidae Holthuis, 1950 comprise three genera from slow-moving freshwater habitats such as streams and lakes in northern South America and West Africa (Pachelle and Tavares 2018) and in a well in India (De Grave et  al. 2018). Typhlocarididae Annandale & Kemp, 1913, with one genus, Typhlocaris Calman, 1909, includes four species distributed in subterranean fresh water in the eastern Mediterranean while one species is found in fresh- and brackish-water caves and wells in Italy (Froglia and Ungaro 2001; Tsurnamal 2008). The remaining family is predominantly marine. In fact, Palaemonidae, with 2060 accepted species is the largest family of marine decapods. Some species of some genera, such as Palaemon, are estuarine. Other genera such as Macrobrachium are dominated by freshwater species. This four-family arrangement of Palaemonoidea was found, by exploration of molecular evidence, to include paraphyletic families (Chow et al. 2020; Kou et al. 2013), an issue discussed further below. Diagnosis. Antennular lateral flagellum with accessory branch (Fig. 6.40h, j). Pereopods 1, 2 dissimilar, first more slender than second. Pereopod 2 carpus entire (Fig.  6.44). Exopods absent from pereopods 1–5. Arthrobranchs absent.

Figures 6.31–6.45, Plates 10, 11 Palaemonidae have more genera than any other family of Decapoda. Only the brachyuran family Xanthidae has a similar number. The older literature that is the foundation for identification of carideans (Chace and Bruce 1993; Holthuis 1993) recognised three families and two subfamilies that have since been included in Palaemonidae. Genera historically included in Gnathophyllidae and Hymenoceridae were long known to share some diagnostic features with some palaemonid genera (Pereira 1997), a notion reinforced by molecular data (Gan et al. 2015; De Grave et al. 2015; Kou et  al. 2013). Not all molecular analyses are in agreement, however, but Gnathophyllidae and Hymenoceridae appear to be nested within the palaemonid subfamily Pontoniinae. Likewise, Palaemoninae and Pontoniinae, the two traditionally recognised subfamilies are both polyphyletic. For practical purposes, however, these can be distinguished morphologically, an observation that enables existing keys to be the basis of this new key to genera. Another family, Anchistioididae with only one genus, was rediagnosed by Chace and Bruce (1993) and treated as a valid family by Holthuis (1993). The family was justified on larval characters (Gurney 1936), one long posterolateral pair of spiniform setae on the telson and an appendix interna on all pleopods in both sexes (Fig. 6.40y) (Bruce 1986c). However, this too has been found to be nested within Palaemonidae s. l. on molecular evidence (Chow et al. 2020). Holthuis’s (1993) keys included 101 genera, some now synonymised  – our key includes 156 genera and does not attempt to reflect phylogeny. The figures are grouped in the former family and subfamily arrangement for convenience. Palaemonids are among the most common shallow water marine shrimps; transparent shrimps found along the shore in most parts of the world are most likely to be palaemonids. Many live in deep water but more than 80% of the genera include species from coral reefs where the species form close associations with the corals themselves or with other invertebrates. These symbioses appear to have evolved at least three times (Chow et al. 2021) and predominate in the former subfamily Pontoniinae. Identification of pontoniine genera is facilitated if the symbiotic association is known. Keys are available for genera associated with echinoderms (Bruce 1982c), sponges (Bruce 2010c), crinoids (Marin 2006) and corals (Bruce 1998a). Bruce (1995) presented a synopsis of genera of the Indo-West Pacific and later Bruce (1998a) offered new keys to pontoniine genera and species. Pontoniinae were reviewed again by Li (2000), updating diagnoses and lists of genera and species. Many genera have been added since, especially as Periclimenes was subdivided (Li 2009), but this process has continued since.

References Chow LH, De Grave S, Tsang LM (2020) The family Anchistioididae Borradaile, 1915 (Decapoda: Caridea) is a synonym of Palaemonidae Rafinesque, 1815 based on molecular and morphological evidence. Journal of Crustacean Biology 40, 277–287. doi:10.1093/jcbiol/ruaa012 De Grave S, Arjun CP, Raghavan R (2018) The discovery of Euryrhynchidae (Crustacea: Decapoda) in India, with the description of a new genus and species. Zootaxa 4462, 367–378. doi:10.11646/zootaxa.4462.3.4 Froglia C, Ungaro N (2001) An unusual new record of Typhlocaris salentina (Caroli, 1923) (Decapoda: Typhlocarididae) from subterranean waters of Apulia (Southern Italy). Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano 142, 103–108. Kou Q, Li X, Chan T-Y, Chu KH, Gan Z (2013) Molecular phylogeny of the superfamily Palaemonoidea (Crustacea: Decapoda: Caridea) based on mitochondrial and nuclear DNA reveals discrepancies with the current classification. Invertebrate Systematics 27, 502–514. doi:10.1071/IS13005 Pachelle PPG, Tavares M (2018) The freshwater shrimp family Euryrhynchidae Holthuis, 1950 (Crustacea: Decapoda: Caridea) revisited, with a taxonomic revision of the genus Euryrhynchus Miers, 1878. Zootaxa 4380, 1–110. doi:10.11646/zootaxa.4380.1.1 Powell CB (1977) A revision of the African freshwater shrimp genus Desmocaris Sollaud, with ecological notes and description of a new species (Crustacea Decapoda Palaemonidae). Revue de Zoologie Africaine 91, 649–674. Tsurnamal M (2008) A new species of the stygobiotic blind prawn Typhlocaris Calman, 1909 (Decapoda, Palaemonidae, Typhlocaridinae) from Israel. Crustaceana 81, 487–501. doi:10.1163/156854008783797534

120

Marine Decapod Crustacea

Fig. 6.31.  Palaemonidae. a, Gnathophyllum taylori Ahyong, 2003; b, Palaemon dolospina dolospina Walker & Poore, 2003; c, Pinnotherotonia rumphiusi Marin & Paulay, 2010; d, Pseudopontonia minuta (Baker, 1907).

Most palaemonid genera are entirely marine but some, such as Macrobrachium and Palaemon contain freshwater, estuarine and amphidromous species. Few genera are entirely freshwater: Bruceonia, Miopontonia, Neopalaemon and Troglomexicanus. Others are estuarine: Arachnochium, Creaseria, Leptocarpus and Tenuipedium. All appear in the key.

Diagnosis. Carapace postantennal longitudinal suture complete. Antennule with accessory flagellum partially fused with dorsal flagellum (Fig.  6.40h, j). Maxilliped 1, exopod caridean lobe usually produced (Figs 6.32i, 6.34h, i, 6.41g). Telson posterior margin with 2 or 3 pairs of spiniform setae.

Key to genera of Palaemonidae * genera confined to fresh water, estuaries or anchialine caves

1. – 2. – 3. –

Maxilliped 3 operculate or suboperculate (Fig. 6.32l–o). Maxilliped 1 exopodal caridean lobe not overreaching endopodal endite (Fig. 6.32i). Mandibular incisor vestigial or absent���������������������������������������������������������������2 Maxilliped 3 usually slender, pereopod-like (Figs 6.41h, 1). Maxilliped 1 exopodal caridean lobe usually overreaching endopodal endite (Figs 6.34g–i, 6.41g). Mandibular incisor usually prominent (Figs 6.34b–d, 6.41a–e)�������������������������������������������������������������������������������������������������������������������������������������������������������������������������8 Maxilliped 3 suboperculate, last 3 articles articulating, wider and articulating with elongate basis (Fig. 6.32n, o)������������������������������������������������������������������������������������������������������������������������������������������ ‘Hymenoceridae’ … 3 Maxilliped 3 operculate, last 2 articles articulating, much narrower than fused squarish-oval proximal articles (Fig. 6.32l, m)���������������������������������������������������������������������������������������������������������������������������‘Gnathophyllidae’ … 4 ‘Hymenoceridae’ Antennular outer flagellum thread-like. Pereopod 2 chela broad, flat (Fig. 6.32q). Maxilliped 3 of 4 articles, foliaceous, second widest (Fig. 6.32o)��������������������������������������������������������������������������������������������� Phyllognathia Antennular outer flagellum broad, leaf-like (Fig. 6.32f). Pereopod 2 chela with flexor margin produced as thin flap (Fig. 6.32p). Maxilliped 3 of 4 articles, last 3 expanded, foliaceous (Fig. 6.32n)����������������������Hymenocera ‘Gnathophyllidae’

6 – Caridea – shrimps

4. – 5.

121

Pereopods 3–5 dactyli with unguis and accessory tooth (Fig. 6.32r, s)�����������������������������������������������������������������������������������5 Pereopods 3–5 dactyli broad, subtriangular, with distal spiniform seta���������������������������������������������������������������������������������7 Rostrum unarmed. Body depressed. Cornea narrower than eyestalk, without papilla (Fig. 6.32c). Pereopod 3 dactylus, distal tooth shorter than tooth on flexor margin (Fig. 6.32s)���������������������������������������Pycnocaris – Rostrum dorsally dentate (Figs 6.31a, 6.32a, b). Body not depressed. Cornea wider than eyestalk, with papilla. Pereopod 3 dactylus, distal tooth longer than tooth on flexor margin (Fig. 6.32r)��������������������������������������������6 6. Antennular peduncle article 1 and scaphocerite wide (Fig. 6.32e, h). Maxilliped 3 ischiomerus outer face convex (Fig. 6.32m). Pereopod 2 carpus much shorter than propodus, fingers toothed����������������������� Gnathophyllum – Antennular peduncle article 1 and scaphocerite slender (Fig. 6.32d, g). Maxilliped 3 ischiomerus outer face concave (Fig. 6.32l). Pereopod 2 slender, carpus almost as long as propodus, fingers without teeth����������������������������������������������������������������������������������������������������������������������������������������������������������������Gnathophylleptum 7. Maxilliped 2 strong, pediform, merus and carpus elongate (Fig. 6.32k)�������������������������������������������������������������������Levicaris – Maxilliped 2 compact, shorter than maxilliped 3 (Fig. 6.32j)���������������������������������������������������������������������� Gnathophylloides 8. Telson usually with 2 pairs of posterior spiniform setae, lateral pair shorter, 1 or more median pairs of plumose setae (Fig. 6.33x–z). Maxilliped 3 usually with pleurobranch�������������������������������������������� ‘Palaemoninae’ … 9 – Telson usually with 3 pairs of posterior spiniform setae (median pair often plumose) (Fig. 6.42). Maxilliped 3 usually without pleurobranch������������������������������������������������������������������������‘Pontoniinae’ … 29 ‘Palaemoninae’ 9. Carapace branchiostegal spine present (Fig. 6.33c, d, e, j, h, m)���������������������������������������������������������������������������������������������10 – Carapace branchiostegal spine absent (Fig. 6.33a, b, f, g, i)�����������������������������������������������������������������������������������������������������17 10. Eyes without pigment (Fig. 6.33d). Antennular peduncle article 1 distal margin concave, anterolateral spine strong. Carapace branchiostegal groove absent. Mandibular palp of 2 articles����������������������������������������������� Creaseria* – Eyes pigmented (except some freshwater genera). Antennular peduncle article 1 distal margin rounded, anterolateral spine small. Mandibular palp present or absent�������������������������������������������������������������������������������������������11 11. Male pleopod 1 with well developed appendix interna (Fig. 6.34j, k)����������������������������������������������������������������������������������� 12 – Male pleopod 1 without or with rudimentary appendix interna��������������������������������������������������������������������������������������������14 12. Rostrum with 2 strong basal teeth, crest-like. Carapace with dorsal tooth at midpoint (Fig. 6.33m). Carapace branchiostegal groove present����������������������������������������������������������������������������������������������������������������������Urocaridella – Rostrum without basal crest. Carapace branchiostegal groove absent (Fig. 6.33e)�������������������������������������������������������������13 13. Mandibular palp of 2 or 3 articles (Fig. 6.34c)���������������������������������������������������������������������������������������������������������������� Leander – Mandibular palp absent (Fig. 6.34d)���������������������������������������������������������������������������������������������������������������������������� Leandrites 14. Thoracic sternite 4 with triangular anteromedian process (Fig. 6.33n). Carapace branchiostegal groove present (Fig. 6.33j)���������������������������������������������������������������������������������������������������������������������������������������������������������������������15 – Thoracic sternite 4 smooth dorsally. Carapace branchiostegal groove absent���������������������������������������������������������������������16 15. Pleonite 5 pleura rounded (Fig. 6.33w). Pereopods 3–5 propodi expanded along flexor margin, with row of spiniform setae (Fig. 6.34q)�������������������������������������������������������������������������������������������������������������������� Rhopalaemon – Pleonite 5 pleuron generally ending in small sharp point (Figs 6.31b, 6.33v). Pereopods 3–5 propodi of even width����������������������������������������������������������������������������������������������������������������������������������������������������������������������������� Palaemon 16. Rostrum elongate, thin, with elevated basal toothed crest (Fig. 6.33h)����������������������������������������������������� Nematopalaemon – Rostrum wide, without elevated basal toothed crest, without ventral teeth (Fig. 6.33c)������������������������������ Calathaemon* 17. Carapace hepatic spine present (Fig. 6.33b, i)����������������������������������������������������������������������������������������������������������������������������18 – Carapace hepatic spine absent (Fig. 6.33a)�������������������������������������������������������������������������������������������������������������������������������� 23 18. Mandibular palp absent������������������������������������������������������������������������������������������������������������������������������������ Pseudopalaemon* – Mandibular palp present���������������������������������������������������������������������������������������������������������������������������������������������������������������19 19. Pereopods 3–5 dactyli biunguiculate (Fig. 6.34o)�����������������������������������������������������������������������������������������������Brachycarpus* – Pereopods 3–5 dactyli simple (Fig. 6.34p)��������������������������������������������������������������������������������������������������������������������������������� 20

122

Marine Decapod Crustacea

20. Pereopod 5 propodus without distal transverse rows of setae on flexor margin�������������������������������������������Neopalaemon* – Pereopod 5 propodus with distal transverse rows of setae on flexor margin�����������������������������������������������������������������������21 21. Pereopods 3–5 subequal in length, pereopod 5 shorter than pereopod 2. Pleonite 5 pleuron with curved posterolateral margin (Fig. 6.33u). Telson with more than 5 pairs of plumose setae on ventroposterior margin (Fig. 6.33x). Male thoracic sternite 8 with paired anterior lobes (Fig. 6.33q)��������������������������� Macrobrachium – Pereopods 3–5 not subequal in length, pereopod 5 longer than pereopod 2. Pleonite 5 pleuron with bilobed posterolateral margin (Fig. 6.33r). Telson with 1 or 2 pairs of plumose setae on ventroposterior margin. Male thoracic sternite 8 without anterior lobes������������������������������������������������������������������������������������������������������������������� 22 22. Branchiostegal groove running from base of hepatic spine to carapace margin (Fig. 6.33b). Thoracic sternite 5 without transverse plate (Fig. 6.33o). Plumose setae on ventroposterior margin of telson longer than inner pair of spiniform setae������������������������������������������������������������������������������������������������������������������Arachnochium* – Branchiostegal groove running below hepatic spine from carapace margin to behind hepatic spine (Fig. 6.33k). Thoracic sternite 5 with transverse plate (Fig. 6.33p). Plumose setae on ventroposterior margin of telson shorter than inner pair of spiniform setae (Fig. 6.33z)����������������������������������������������������Tenuipedium* 23. Marine. Carapace with blunt postocular tooth. Maxilliped 1 exopod lash vestigial (Fig. 6.34g). Mandibular molar flared distally (Fig. 6.34b)������������������������������������������������������������������������������������������������������������������������ Anchistioides – Freshwater. Carapace without blunt postocular tooth. Maxilliped 1 exopod lash fully formed (Fig. 6.34h, i). Mandibular molar cylindrical (Fig. 6.34c, d)���������������������������������������������������������������������������������������������������������������������� 24 24. Rostrum shorter than scaphocerite, with small teeth if any. Eyes without pigment (or single spot). Mandibular palp absent. Freshwater only������������������������������������������������������������������������������������������������������������������������������������ 25 – Rostrum usually longer than scaphocerite, toothed. Eyes pigmented. Mandibular palp present or absent������������������� 27 25. Eyes with small cornea. Pereopod 2 carpus with ring-like constriction behind and parallel with distal margin (Fig. 6.34n)����������������������������������������������������������������������������������������������������������������������������������������������� Troglindicus* – Eyes without pigment. Pereopod 2 carpus without ring-like subdistal constriction��������������������������������������������������������� 26 26. Maxilla endite bilobed (Fig. 6.34e). Maxilliped 1 epipod simple (Fig. 6.34h) (Caribbean)������������������������ Troglocubanus* – Maxilla endite simple (Fig. 6.34f). Maxilliped 1 epipod bilobed (Fig. 6.34i) (Mexico)���������������������������Troglomexicanus* 27. Rostrum with elevated basal toothed crest (Fig. 6.33f). Pleonite 4 sternite with slender median process������������������������������������������������������������������������������������������������������������������������������������������������������ Leptocarpus* – Rostrum without elevated basal toothed crest (Fig. 6.33g). Pleonite 4 sternite smooth���������������������������������������������������� 28 28. Pleonite 5 pleuron square (Fig. 6.33t). Pereopod 2 slender, smooth, carpus as long as chela (Fig. 6.34m). Mandibular palp absent or of 1 article����������������������������������������������������������������������������������������������������������Leptopalaemon* – Pleonite 5 pleuron rounded (Fig. 6.33s). Pereopod 2 robust, spinulate, carpus less than half as long as chela (Fig. 6.34l). Mandibular palp of 3 articles���������������������������������������������������������������������������������������������������Cryphiops* ‘Pontoniinae’ 29. Maxilliped 3 with 2 arthrobranchs. Thoracic sternite 5 with pair of slender submedian anterior processes (Fig. 6.40b). Pereopod 1 chela fingers longer than palm, subspatulate laterally����������������������� Exoclimenella – Maxilliped 3 without or with 1 arthrobranch. Thoracic sternite 5 without processes������������������������������������������������������ 30 30. Mandibular palp of 1 or 2 articles�����������������������������������������������������������������������������������������������������������������������������������������������31 – Mandibular palp absent����������������������������������������������������������������������������������������������������������������������������������������������������������������33 31. Carapace hepatic spine present (Fig. 6.37n)�����������������������������������������������������������������������������������������������������������Palaemonella – Carapace hepatic spine absent (Fig 6.39g)����������������������������������������������������������������������������������������������������������������������������������32 32. Carapace supraorbital carina absent (Fig. 6.39g). Pereopods 3–5 propodi flexor margin unarmed������������������������������Vir – Carapace supraorbital carina present. Pereopod 3–5 flexor margin with spinules�������������������������������������������� Eupontonia 33. Scaphocerite rudimentary (Fig. 6.40n) or absent��������������������������������������������������������������������������������������������������������������������� 34 – Scaphocerite well developed�������������������������������������������������������������������������������������������������������������������������������������������������������� 38

6 – Caridea – shrimps

123

34. Rostrum absent (Fig. 6.37r). Pereopods 3–5 dactyli simple������������������������������������������������������������������������������������� Paratypton – Rostrum present (Figs 6.39a, 6.38s, t). Pereopods 3–5 dactyli biunguiculate�����������������������������������������������������������������������35 35. Minor pereopod 2 with shearing fingers with 2 apical teeth (Fig. 6.44B). Endites of maxilla and maxilliped 1 elongate, setose, basket-like������������������������������������������������������������������������������������������������������������Typtonychus – Pereopod 2 fingers simple. Mouthparts unmodified��������������������������������������������������������������������������������������������������������������� 36 36. Labrum triangular, anteromedian margin exceeding bases of antennules (Fig. 6.40p). Maxillipedal 1 exopodal flagella strap-like, with numerous plumose distal and marginal setae (Fig. 6.41f)�����������������Onycomenes – Labrum normal. Maxillipedal exopodal flagella slender, with few plumose distal setae���������������������������������������������������37 37. Rostrum long, with 9 dorsal teeth (Fig. 6.38t)����������������������������������������������������������������������������������������������������������Typtonoides – Rostrum very short, with few or without teeth (Fig. 6.38s)�������������������������������������������������������������������������������������������� Typton 38. Maxilliped 3 (sometimes maxillipeds 1, 2) without exopod or with reduced exopod������������������������������������������������������� 39 – Maxillipeds 1–3 with exopods����������������������������������������������������������������������������������������������������������������������������������������������������� 60 39. Carapace hepatic spine present��������������������������������������������������������������������������������������������������������������������������������������������������� 40 – Carapace hepatic spine absent����������������������������������������������������������������������������������������������������������������������������������������������������� 48 40. Carapace with oblique supraorbital dentate carina (Fig. 6.35j)������������������������������������������������������������������������������������� Balssia – Carapace with or without (Figs 6.36k, 6.37c, 6.38f) supraorbital tooth beside rostrum�����������������������������������������������������41 41. Pleonites 4–6 pleura with posteroventral sharp tooth (Fig. 6.40d, f)����������������������������������������������������������������������������������� 42 – Pleonites 4–6 pleura rounded������������������������������������������������������������������������������������������������������������������������������������������������������ 43 42. Pleonites 2–6 pleura rounded, with posteroventral sharp tooth (Fig. 6.40d). Telson posterior margin with 2 large immovable teeth (Fig. 6.42g)��������������������������������������������������������������������������������������������������������������� Coutierea – Pleonites 4–6 pleura with posteroventral sharp tooth (Fig. 6.40f). Telson posterior margin convex, with 6 spiniform setae, second longest (Fig. 6.42m)�������������������������������������������������������������������������������������������������� Lipkebe 43. Pereopods 3–5 dactyli with basal protuberance (Fig. 6.45o). Pereopod 1 fingers much less than half length of palm (Fig. 6.43b)�������������������������������������������������������������������������������������������������������������������� Hamodactyloides – Pereopods 3–5 dactyli without basal protuberance (Fig. 6.45p, u). Pereopod 1 fingers at least half length of palm������������������������������������������������������������������������������������������������������������������������������������������������������������������� 44 44. Pereopod 2 dactylus much longer than fixed finger, hook-like (Fig. 6.44n)������������������������������������������������������������������������ 45 – Pereopod 2 dactylus as long fixed finger, straight�������������������������������������������������������������������������������������������������������������������� 46 45. Pereopod 2 carpus, merus and ischium with 1–3 teeth on flexor margin (Fig. 6.44n). Rostrum without teeth. Carapace pterygostomial notch present (Fig. 6.37c)������������������������������������������������������������������������������Miopontonia – Pereopod 2 carpus, merus and ischium without teeth on flexor margin. Rostrum elevated, with dorsal teeth. Carapace pterygostomial notch absent (Fig. 6.36k)���������������������������������������������������������������������������� Hamodactylus 46. Pereopods 2 similar, cutting edges without teeth (Fig. 6.44y). Pereopods 3–5 dactyli simple�������������������������Propontonia – Pereopods 2 dissimilar, cutting edges of larger with teeth (Fig. 6.44l). Pereopods 3–5 dactyli simple or biunguiculate����������������������������������������������������������������������������������������������������������������������������������������������������������������������� 47 47. Rostrum blade-like, with dorsal and sometimes few ventral teeth, with epigastric tooth (Fig. 6.36y). Maxilliped 3 without exopod (Fig. 6.42h). Western, Central Pacific�����������������������������������������������������������Mesopontonia – Rostrum dorsal margin convex, with 6 teeth, ventral margin straight, without epigastric teeth (Fig. 6.39h). Maxilliped 3 with exopod. Eastern Pacific������������������������������������������������������������������������������������������Waldola 48. Carapace with prominent dorsal horns������������������������������������������������������������������������������������������������������������������������������������� 49 – Carapace dorsally smooth������������������������������������������������������������������������������������������������������������������������������������������������������������ 50 49. Carapace with 1 dorsal conical, median horn, 2 smaller posterolateral horns (Fig. 6.38n). Pleonite 2 with pair of horns, pleonite 3 with median horn���������������������������������������������������������������������������������Sandyella – Carapace with 2 dorsal median horns in anterior quarter, small knob posterior to orbit (Fig. 6.35q). Pleonites without dorsal horns����������������������������������������������������������������������������������������������������������������������������������� Chacella

124

Marine Decapod Crustacea

50. Pleonites 3–5 pleura produced as narrow blunt projection or with posteroventral sharp tooth. Rostrum slender, dorsally and ventrally unarmed, proximal part extended laterally to form supraorbital eave (Fig. 6.38h)��������������������������������������������������������������������������������������������������������������������������������������������������������� Pseudocoutierea – Pleonites 3–5 pleura rounded�������������������������������������������������������������������������������������������������������������������������������������������������������51 51. Carapace orbital posterior margin with comb-like row of c. 20 spines (Fig. 6.35x)��������������������������������������Ctenopontonia – Carapace orbital posterior margin without comb-like row of spines������������������������������������������������������������������������������������52 52. Rostrum slender, narrow in dorsal view, orbit exposed�����������������������������������������������������������������������������������������������������������53 – Rostrum basally wide, forming roof over orbit������������������������������������������������������������������������������������������������������������������������ 54 53. Carapace postorbital spine present; branchiostegal spine absent (Fig. 6.38p). Pereopod 2 dactylus semiglobular, fixed finger a small process bearing short spine (Fig. 6.44z)��������������������������������������������������������Tectopontonia – Carapace postorbital spine absent; branchiostegal spine present (Fig. 6.37f). Pereopod 2 dactylus simple�����������������������������������������������������������������������������������������������������������������������������������������������������������������Neopericlimenes 54. Rostrum scarcely reaching beyond eyes, fully depressed, not ending in laterally compressed distal part, without erect teeth��������������������������������������������������������������������������������������������������������������������������������������������������������������������55 – Rostrum reaching beyond eyes, broad, basally depressed, ending in laterally compressed distal part, often with teeth������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������57 55. Orbits complete, closed. Antenna article 1 with prominent distolateral and lateral projections (Fig. 6.38j). Pereopod 1 fingers spatulate (Fig. 6.43f)��������������������������������������������������������������������������������� Pseudoveleronia – Orbits not closed (Fig. 6.39e, f). Antenna article 1 with slight distolateral and lateral projections. Pereopod 1 fingers simple������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 56 56. Uropodal exopod anterodistal tooth movable (Fig. 6.42z). Minor pereopod 2 sexually dimorphic, fingers of male thin, as long as palm (Fig. 6.44w)������������������������������������������������������������������������������������������������������������������ Pontonides – Uropodal exopod anterodistal tooth fixed (Fig. 6.42B). Minor pereopods 2 not sexually dimorphic, fingers stout (Fig. 6.44C)����������������������������������������������������������������������������������������������������������������������������������������������������������Veleronia 57. Carapace postorbital margin inflated, with noticeable postorbital tubercle (Fig. 6.39f). Maxilliped 2 with exopod����������������������������������������������������������������������������������������������������������������������������������������������������������������������Veleroniopsis – Carapace postorbital tubercle absent. Maxilliped 2 without exopod or without exopodal flagellum����������������������������� 58 58. Rostrum reaching beyond eyes, gradually narrowing from expanded base to compressed distal part, with dorsal teeth (Fig. 6.37g)�����������������������������������������������������������������������������������������������������������������������������Neopontonides – Rostrum basal part widely expanded, forming roof of orbit, narrowing suddenly to compressed distal part (Figs 6.36p, 6.38i)����������������������������������������������������������������������������������������������������������������������������������������������������������������������59 59. Pterygostomial sinus present, opening to branchial cavity, antennal spine small (Fig. 6.38i)��������������� Pseudopontonides – Pterygostomial sinus absent, prominent antennal spine at end of ridge below deep orbital depression (Fig. 6.36p)������������������������������������������������������������������������������������������������������������������������������������������������������������������������Izucaris 60. Carapace hepatic spine present (submarginal or set well back, sometimes very small)�����������������������������������������������������61 – Carapace hepatic spine absent��������������������������������������������������������������������������������������������������������������������������������������������������� 100 61. Rostrum with lateral carinae expanded proximally forming supraorbital eaves (Fig. 6.36s, u)����������������������������������������62 – Rostrum without or with weak basal lateral carinae��������������������������������������������������������������������������������������������������������������� 63 62. Cornea conically produced (Fig. 6.36s). Epistome with pair of acute processes (Fig. 6.40o)������������������������������� Laomenes – Cornea globular. Epistome without acute processes������������������������������������������������������������������������������������������������ Lipkemenes 63. Rostrum long, without teeth, continued posteriorly to posterior margin as high carina (Fig. 6.35p)����������������������������������������������������������������������������������������������������������������������������������������������������������Carinopontonia – Rostrum usually with dorsal teeth, not continued to posterior margin as carina�������������������������������������������������������������� 64 64. Pereopods 3–5 dactyli with large basal protuberance (Fig. 6.45k, s)������������������������������������������������������������������������������������� 65 – Pereopods 3–5 dactyli without basal protuberance����������������������������������������������������������������������������������������������������������������� 66 65. Body strongly depressed (Fig. 6.39q). Rostrum with dorsal teeth. Pereopods 3–5 dactyli with basal protuberance hoof-shaped (Fig. 6.45s)���������������������������������������������������������������������������������������������������������������������������Jocaste

6 – Caridea – shrimps

– 66. – 67. – 68. – 69. – 70. – 71. – 72. – 73. – 74. – 75. – 76. – 77. – 78. – 79. – 80. – 81.

125

Body not strongly depressed. Rostrum acute, without teeth (Fig. 6.36c). Pereopods 3–5 dactyli with basal protuberance acute or rounded (Fig. 6.45k)�����������������������������������������������������������������������������������������������������������������Dasella Pereopod 2 propodus with 2 proximal lobes fitting into cavity on distal margin of carpus (Fig. 6.44e). Carapace hepatic spine close to and lower than antennal spine (Fig. 6.35w)�������������������������������������������������Cristimenes Pereopod 2 propodus and carpus articulating simply. Carapace hepatic spine usually set well back from margin������������� 67 Pleonites 4, 5 pleura with posteroventral sharp tooth (Fig. 6.40e)���������������������������������������������������������������������������������������� 68 Pleonites 1–5 pleura rounded or blunt��������������������������������������������������������������������������������������������������������������������������������������� 69 Body not depressed. Rostrum slender, styliform, with 0–2 dorsal teeth, without ventral teeth (Fig. 6.36d)������������������������������������������������������������������������������������������������������������������������������������������������������������������� Dasycaris Body depressed. Rostrum laterally compressed, with dorsal and ventral teeth (Fig. 6.36m)�������������������������� Harpiliopsis Hepatic spine movable (Fig. 6.37e, q, s)���������������������������������������������������������������������������������������������������������������������������������������70 Hepatic spine fixed�������������������������������������������������������������������������������������������������������������������������������������������������������������������������74 Pereopod 3 propodus with spiniform setae only distally on flexor margin (Fig. 6.45x)�����������������������������������������������������71 Pereopod 3 propodus with spiniform setae along entire flexor margin (Fig. 6.45b, F)����������������������������������������������������� 73 Rostrum with few, small, dorsal and ventral teeth distally (Fig. 6.37q). Major pereopod 2 carpus much shorter than chela (Fig. 6.44t)������������������������������������������������������������������������������������������������������������������������������ Paranchistus Rostrum with teeth large over full dorsal margin. Major pereopod 2 carpus longer or shorter than chela������������������� 72 Rostrum deep, dorsal margin convex, with 10 close teeth, without ventral teeth; epigastric tooth absent (Fig. 6.37s). Pereopod 2 carpus longer than chela������������������������������������������������������������������������������������������������������ Patonia Rostrum slender, dorsal margin convex, with 8 close teeth, ventral margin with 3 teeth; epigastric tooth present, with suture (Fig. 6.37e). Pereopod 2 carpus shorter than chela��������������������������������������������������������Neoclimenes Pereopods 1 fingers spatulate (Fig. 6.43j). Pereopods 2–5 meri with small distal tooth on flexor margin (Fig. 6.45F)������������������������������������������������������������������������������������������������������������������������������������������������������������� Zenopontonia Pereopods 1 fingers simple (Fig. 6.43a). Pereopods 2–5 meri without distal tooth on flexor margin�����������Allopontonia Carapace antennal spine absent (Figs 6.37p, 6.39d)������������������������������������������������������������������������������������������������������������������75 Carapace antennal spine present�������������������������������������������������������������������������������������������������������������������������������������������������76 Rostrum ventral carina deep, with 3 or 4 prominent ventral teeth (Fig. 6.37p)����������������������������������������������� Paraclimenes Rostrum compressed, dorsal carina with numerous acute teeth, lateral carinae weak, ventral carina obsolete, weakly dentate distally (Fig. 6.39d)������������������������������������������������������������������������������������������������������������Urocaris Rostrum large, T- or cross-shaped in transverse section, without dorsal or ventral teeth. Carapace with postorbital depression (Fig. 6.38o)�����������������������������������������������������������������������������������������������������������Stegopontonia Rostrum laterally compressed, with at least dorsal teeth�������������������������������������������������������������������������������������������������������� 77 Rostrum basal part with narrow lateral wings, narrowing to compressed distal part, with small dorsal and usually ventral teeth. Carapace postorbital groove distinct, bordered by postorbital carina and carina from antennal to hepatic spine (Fig. 6.38r)��������������������������������������������������������������������������������������������Tuleariocaris Rostrum without lateral wings, with conspicuous dorsal teeth, ventral teeth present or absent (Figs 6.36a, m, v, w, 6.38k, l). Carapace postorbital groove narrow or indistinct��������������������������������������������������������������������������������78 Thoracic sternite 4 with slender median spine. Pereopods 3–5 dactyli simple������������������������������������������������������������������� 79 Thoracic sternite 4 without process, or at most with short angular lobe����������������������������������������������������������������������������� 82 Pereopod 1 fingers stout, deeply subspatulate, with mesial subspatulate concavity, without distal lamina (Fig. 6.43d). Pereopod 2 merus with distoventral tooth, major chela with fossae on opposable cutting margins of both fingers (Fig. 6.44u)���������������������������������������������������������������������������������Periclimenella Pereopod 1 fingers not subspatulate. Pereopod 2 without fossae on fingers����������������������������������������������������������������������� 80 Pereopods 3–5 propodi distally setose, without spiniform setae, dactyli short, stout, hook-like (Fig. 6.45q)���Harpilius Pereopods 3–5 propodi with spiniform setae on flexor margin, dactylus slender��������������������������������������������������������������81 Pereopods 2 feeble, symmetrical, barely exceeding pereopods 1; fingers simple, longer than palm (Fig. 6.44k)�����������������������������������������������������������������������������������������������������������������������������������������������������������������Madangella

126

Marine Decapod Crustacea

– Pereopods 2 well developed, symmetrical or not, well exceeding pereopods 1; fingers shorter than palm��������Cuapetes 82. Pereopods 3–5 propodi prehensile, with 2–5 pairs of long straight spiniform setae; dactyli slender, curved, simple (Fig. 6.45C). Carapace epigastric spine with basal suture (Fig. 6.38k, l)���������������������������������������������� 83 – Pereopods 3–5 propodi not prehensile, without pairs of long straight spiniform setae. Carapace epigastric spine without basal suture������������������������������������������������������������������������������������������������������������������������������������ 84 83. Pereopod 1 fingers narrow, simple (Fig. 6.43g). Rostrum dorsal margin convex (Fig. 6.38k)��������������������������� Rapimenes – Pereopod 1 fingers expanded, spatulate (Fig. 6.43h). Rostrum high at base (Fig. 6.38l)���������������������������������Rapipontonia 84. Rostrum slender, far over-reaching antennal scale. Carapace epigastric tooth and 2 posterior rostral teeth large, similar, curved anteriorly, without sutures (Fig. 6.36w). Maxilliped 3 coxa with 1 or more long plumose setae and 1 arthrobranch������������������������������������������������������������������������������������������������������������������� Manipontonia – Rostrum not over-reaching antennal scale. Carapace epigastric tooth smaller than posterior rostral teeth, if present. Maxilliped 3 coxa without long plumose seta������������������������������������������������������������������������������������������������������ 85 85. Pereopod 2 fingers 3 times as long as palm. Rostrum unarmed ventrally. Carapace supraorbital spine absent (Fig. 6.35m). Pereopods 3–5 propodi with row of long distal setae extending nearly to base of dactylus unguis, dactyli simple (Fig. 6.45g)����������������������������������������������������������������������������������������������������������� Brucecaris – Pereopod 2 fingers usually subequal or shorter than palm. Rostrum with or without ventral teeth. Carapace supraorbital spine present or absent. Pereopods 3–5 propodi without distal setae extending to near end of dactylus; dactyli simple, biunguiculate or more ornate��������������������������������������������������������������������������� 86 86. Pereopods 3–5 dactyli corpus upper margin with distal accessory spinules (Fig. 6.45E). Mandibular incisor wider distally (Fig. 6.41e)����������������������������������������������������������������������������������������������������������������������������������������� Unguicaris – Pereopods 3–5 dactyli corpus without accessory spinules. Mandibular incisor not wider distally�������������������������������� 87 87. Thoracic sternite 3 ~4 times as long as thoracic sternite 4. Antennular peduncle article 2 elongate, flat (Fig. 6.40k). Pereopod 2 slender, fingers with recurved teeth on cutting edges (Fig. 6.44j). Rostrum not over-reaching end of antennular peduncle article 1 (Fig. 6.36t)������������������������������������������������������������������ Leptomenaeus – Thoracic sternite 3 not elongate. Antennular peduncle article 2 not elongate (Fig. 6.40i)������������������������������������������������ 88 88. Rostrum well developed, toothless. Carapace orbit deep, with posterior notch; supraorbital, epigastric spines absent (Fig. 6.36x). Pereopods 3–5 dactyli biunguiculate, unguis elongate, corpus with crenulate distoventral accessory tooth, with transverse row of simple setae distolaterally (Fig. 6.45t)������������������������Margitonia – Rostrum usually with numerous dorsal and sometimes fewer ventral teeth (Figs 6.35v, 6.37w). Carapace orbits without posterior notch; supraorbital, epigastric spines present or absent. Pereopods 3–5 dactyli simple or biunguiculate, sometimes ornate������������������������������������������������������������������������������������������������������������������������� 89 89. Antennular peduncle article 1 with more than 1 distolateral teeth, mesial ventral tooth robust (Fig. 6.40i). Pereopods 2 highly dissimilar, unequal, asymmetrical, major fixed finger with 2 proximal teeth fitting into 2 sockets on dactylus (Fig. 6.44d). Pereopod 3 propodus with serrated flexor border, without spiniform setae; dactylus biunguiculate, with slender, curved unguis (Fig. 6.45i)����������������������������������������� Crinotonia – Antennular peduncle article 1 with 1 distolateral tooth, mesial ventral tooth small (Fig. 6.40h). Pereopods 2 equal or unequal. Pereopod 3 dactylus simple, biunguiculate or with small distal spines on flexor margin��������������� 90 90. Thoracic sternite 4 with short triangular anterior lobe (Fig. 6.40a). Pereopods 2 similar to others. Pereopods 3–5 dactyli biunguiculate (Fig. 6.45z)�������������������������������������������������������������������������������������������������������� Phycomenes – Thoracic sternite 4 without triangular anterior process. Pereopods 2 distinctly longer than others. ­Pereopods 3–5 dactyli biunguiculate or not�������������������������������������������������������������������������������������������������������������������������91 91. Carapace orbital rim round, lower half laminar, raised laterally, separated by distinct notch from leaf-like inferior orbital angle; hepatic spine submarginal below antennal spine (Fig. 6.35l). Pereopod 3 dactylus simple��������Blepharocaris – Carapace orbital inferior angle not leaf-like; hepatic spine rarely submarginal����������������������������������������������������������������� 92 92. Pereopods 3–5 propodi with bunches of long setae along flexor margin; dactyli simple (Fig. 6.45D). Body and appendages covered by dense pubescence����������������������������������������������������������������������������������������������������Sandimenes – Pereopods 3–5 propodi without bunches of long setae along flexor surface; dactyli usually biunguiculate, or with small distal spines on flexor margin. Body and appendages not covered by dense pubescence������������������� 93

6 – Caridea – shrimps

93. – 94.

– 95.

– 96. – 97. – 98. – 99. – 100. – 101. – 102. – 103. – 104. – 105. – 106.

127

Carapace prominently humped in adult female (Fig. 6.35f). Telson with 2 pairs of minute dorsal spines on posterior third (Fig. 6.42d)������������������������������������������������������������������������������������������������������������������������������������� Ancylocaris Carapace not prominently humped in adult females. Telson otherwise ornamented������������������������������������������������������� 94 Pereopods 3–5 dactyli with stout unarmed corpus, unguis elongate, curved (Fig. 6.45a). Rostrum with 7–10 dorsal teeth (posterior tooth behind orbits), 0–2 ventral teeth. Carapace hepatic spine almost level with antennal spine (Fig. 6.35a). Telson with 2 pairs of large dorsal marginal spines on anterior and posterior thirds (Fig. 6.42a)����������������������������������������������������������������������������������������������������������������������������������������������������Actinimenes Pereopods 3–5 dactyli, rostrum, hepatic spine, telson together not as above���������������������������������������������������������������������� 95 Rostrum horizontal, ventral margin straight, dentate, dorsal margin dentate (Fig. 6.37b). Pereopod 1 fingers with cutting edges unarmed; dactylus not subspatulate. Major pereopod 2 fixed finger with proximal excavation on cutting surface receiving and overlapping proximal part of dactylar cutting edge; dactylus with flange on upper margin (Fig. 6.44m)��������������������������������������������������������������������������� Michaelimenes Rostrum, pereopod 1 not as above. Pereopod 2 fingers cutting edges meeting, not overlapping������������������������������������ 96 Major pereopod 2 palm granulate, fingers with lateral flange; dactylus cutting edge with basal tooth-like process���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 97 Major pereopod 2 palm rarely granulate, fingers without lateral flanges; dactylus cutting edge without basal tooth-like process����������������������������������������������������������������������������������������������������������������������������������������������������������� 98 Carapace hepatic spine behind anterior margin. Rostrum with most posterior tooth clearly separate from others, epigastric (Fig. 6.35k). Telson with 3–7 pairs of dorsolateral spiniform setae. Pereopods 3–5 dactyli biunguiculate (Fig. 6.45f)�������������������������������������������������������������������������������������������������������������������������� Bathymenes Carapace hepatic spine overlapping anterior margin. Rostrum teeth evenly spaced (Fig. 6.36f). Telson with 2 pairs of dorsolateral spiniform setae (Fig. 6.42h). Pereopods 3–5 dactyli truncate subdistally, with denticulate lobe on flexor margin (Fig. 6.45l)����������������������������������������������������������������������������������������Echinopericlimenes Pleon ~4 times as long as carapace. Inferior orbital margin strongly produced, with reflected inner ventral flange (Fig. 6.40c). Antennular peduncle article 1 with large anterolateral lobe (Fig. 6.40h)��������������������Ancylomenes Pleon much less than 4 times as long as carapace. Inferior orbital margin not strongly produced, without reflected inner ventral flange (Figs 6.37t, 6.39j). Antennular peduncle article 1 without large anterolateral lobe��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 99 Pereopods 3–5 dactyli with small distal spines on flexor margin (Fig. 6.45G)��������������������������������������������������������Zoukaris Pereopods 3–5 dactyli biungiculate or rarely simple����������������������������������������������������������������������������������������������Periclimenes Carapace, pleon and limbs depressed, completely densely covered in short setae (Fig. 6.31c)����������������� Pinnotherotonia Carapace, pleon and limbs smooth, not pubescent all over���������������������������������������������������������������������������������������������������101 Pereopods 3–5 dactyli with basal protuberance (Fig. 6.45h, n)������������������������������������������������������������������������������������������� 102 Pereopods 3–5 dactyli without basal protuberance��������������������������������������������������������������������������������������������������������������� 104 Body strongly depressed (Fig. 6.35u). Pereopods 3–5 dactyli with basal protuberance hoof-shaped (Fig. 6.45h)��������������������������������������������������������������������������������������������������������������������������������������������������������������� Coralliocaris Body rounded or depressed. Pereopods 3–5 dactyli with basal protuberance rounded (Fig. 6.45n)������������������������������103 Carapace postorbital ridge with 3–5 spines in oblique row behind antennal spine (Fig. 6.36i)����������������������������� Fennera Carapace postorbital ridge without spines (Fig. 6.35t)������������������������������������������������������������������������������������������ Conchodytes Rostrum vestigial or short, triangular. Scaphocerite tapering, acute or bifid������������������������������������������������������������������� 105 Rostrum well developed. Scaphocerite blade-like������������������������������������������������������������������������������������������������������������������ 106 Rostrum triangular, with 1 dorsal tooth. Scaphocerite bifid (Fig. 6.36r). Telson with 2 pairs of strong dorsal spiniform setae in anterior quarter����������������������������������������������������������������������������������������������������������������������� Kaviengella Rostrum short, triangular (Fig. 6.37h). Scaphocerite tapering. Telson with second pair of dorsal spiniform setae at midlength (Fig. 6.42o)������������������������������������������������������������������������������������������������������������������������������Nippontonia Rostrum laterally compressed or triangular, surface not concave, usually with teeth (e.g. Figs 6.35d, r, 6.36n, 6.37k, o, 6.38q, u)������������������������������������������������������������������������������������������������������������������������ 107

128

– 107. – 108. – 109. – 110. – 111. – 112. – 113. – 114. – 115. – 116. – 117. – 118. – 119.

– 120. – 121. –

Marine Decapod Crustacea

Rostrum depressed (surface concave) or cylindrical-conical, usually toothless (Figs 6.35c, 6.37a, y, 6.38c, d)����������� 148 Pereopods 1 unequal, carpus segmented (Fig. 6.43i)��������������������������������������������������������������������������������������� Thaumastocaris Pereopods 1 equal, carpus not segmented������������������������������������������������������������������������������������������������������������������������������� 108 Pereopods 2 unequal, major dactylus or finger with molar-shaped tooth fitting into socket of opposite finger, or dactylus entire, fitting into longitudinal groove on finger����������������������������������������������������������������������������� 109 Pereopods 2 equal or unequal, fingers without opposing molar-fossa��������������������������������������������������������������������������������115 Major pereopod 2, fixed finger with 1 or 2 teeth, dactylus with matching sockets (Fig. 6.44s)��������������������������������������110 Major pereopod 2 dactylus entire or with tooth, fixed finger with groove�������������������������������������������������������������������������111 Carapace supraorbital tooth absent (Fig. 6.37o). Uropodal exopod with numerous spiniform setae on suture and anterior margin (Fig. 6.42y)���������������������������������������������������������������������������������������������������������������������Paraclimenaeus Carapace supraorbital tooth present (Fig. 6.39b, b’). Uropodal exopod suture with 5 spiniform setae laterally, middle one longest (Fig. 6.42A)��������������������������������������������������������������������������������������������������������������� Unesconia Pereopod 2 chela spinose, flattened (Fig. 6.44b)�������������������������������������������������������������������������������������������������� Anisomenaeus Pereopod 2 chela smooth, oval���������������������������������������������������������������������������������������������������������������������������������������������������112 Major pereopod 2 dactylus entire, fitting into longitudinal depression on finger (Fig. 6.44v)��������������������Plesiomenaeus Major pereopod 2 dactylus with molar-shaped tooth fitting into socket of fixed finger�������������������������������������������������113 Scaphocerite shorter than antenna peduncle. Minor pereopod 2 similar to major, dactylus with molar, fixed finger with fossa (Fig. 6.44A)��������������������������������������������������������������������������������������������������������������Typtonomenaeus Scaphocerite longer than antenna peduncle. Minor pereopod 2 dissimilar to major, dactylus, fixed finger without molar-fossa����������������������������������������������������������������������������������������������������������������������������������������������������������������114 Telson with 3 pairs of dorsal and marginal spiniform setae (Fig. 6.42k). Mandibular incisor vestigial (Fig. 6.41b)���������������������������������������������������������������������������������������������������������������������������������������������������������������Holthuisaeus Telson with 2 pairs of dorsal spiniform setae (Fig. 6.42q). Mandibular incisor prominent (Fig. 6.41c)����Periclimenaeus Major pereopod 2 fingers with opposing fossae (Fig. 6.44c). Orbit with supraorbital tooth (Fig. 6.35r)�������� Climeniperaeus Major pereopod 2 fingers without opposing fossae (Fig. 6.44q). Orbit without supraorbital tooth (Figs 6.35d, 6.36o)�������������������������������������������������������������������������������������������������������������������������������������������������������������������116 Pereopod 3 dactylus more than 6 times as long as wide, with long unguis about as long as propodus (Fig. 6.45v)������������������������������������������������������������������������������������������������������������������������������������������������������������Onycocaridites Pereopods 3–5 dactyli similar, not more than 4 times as long as wide, less than half as long as propodus������������������117 Uropodal exopod anterior margin spinose or suture lobed, with spiniform setae������������������������������������������������������������118 Uropodal exopod anterior margin straight, ending in tooth and spiniform seta��������������������������������������������������������������119 Uropodal exopod suture lobed, with spiniform setae (Fig. 6.42w). Rostrum with supraocular tooth, triangular, with 5 dorsal teeth, depressed tip (Fig. 6.35g)�������������������������������������������������������������������������������Apopontonia Uropodal exopod anterior margin spinose (Fig. 6.42v). Rostrum steep, with ~15 small dorsal teeth from tip to middle carapace (Fig. 6.35d)����������������������������������������������������������������������������������������������������������������������Anapontonia Carapace with 3 or more teeth behind orbit, extending to mid-carapace (Fig. 6.36o). Uropodal exopod spiniform seta almost reaching end of exopod, curved (Fig. 6.42x). Telson with all spiniform setae on distal third of posterior margin. Body laterally compressed�����������������������������������������������������������������������Ischnopontonia Carapace with 0–3 teeth behind orbit, on anterior quarter of carapace (Figs 6.35h, 6.36e, j, l, 6.38a, 6.39i). Uropodal exopod spiniform seta small, not curved��������������������������������������������������������������������������������������������������������� 120 Telson far exceeding uropods, distally expanded, with 3 pairs of dorsal spiniform setae, without posterior spiniform setae (Fig. 6.42C)����������������������������������������������������������������������������������������������������������������������������������� Yemenicaris Telson about as long as uropods, not distally expanded, with 2 or 4 pairs of dorsal spiniform setae, with posterior spiniform setae�������������������������������������������������������������������������������������������������������������������������������������������������������121 Telson with 4 pairs of dorsal spiniform setae (Fig. 6.42s)����������������������������������������������������������������������������������Plesiopontonia Telson with 2 pairs of dorsal spiniform setae������������������������������������������������������������������������������������������������������������������������� 122

6 – Caridea – shrimps

129

122. Telson posterior margin without spiniform setae, ending in 2 large downcurved teeth (Fig. 6.42j)��������� Hamopontonia – Telson posterior margin with 6 setae (Fig. 6.42e) or without spiniform setae, rounded������������������������������������������������� 123 123. Mandibular incisor wider distally (Fig. 6.41a). Pereopod 3 dactylus tip comprising tooth, unguis, spiniform seta (Fig. 6.45e)���������������������������������������������������������������������������������������������������������������������������������� Araiopontonia – Mandibular incisor not wider distally. Pereopod 3 dactylus simple or biunguiculate����������������������������������������������������� 124 124. Pereopods 3–5 dactyli largely hidden by dense tuft of propodal setae (Fig. 6.45y)�������������������������������������������������Philarius – Pereopods 3–5 dactyli not hidden by setae (Fig. 6.45j, r, m)������������������������������������������������������������������������������������������������� 125 125. Carapace with 1–3 epigastric teeth (Figs 6.36e, j, 6.38a, g)��������������������������������������������������������������������������������������������������� 126 – Carapace without epigastric teeth, all teeth on rostrum (Figs 6.36g, 6.38e)���������������������������������������������������������������������� 129 126. Pereopods 2 unequal, major fixed finger cutting edge with bifid lobe (Fig. 6.44h). Rostrum with 2 ventral teeth (Fig. 6.36j)������������������������������������������������������������������������������������������������������������������������������������������������������������� Hamiger – Pereopods 2 equal, fixed finger cutting edge with up to 2 small proximal teeth. Rostrum with 0–2 ventral teeth����������� 127 127. Rostrum high, not reaching beyond antennular peduncle article 1, with 4 dorsal teeth, without ventral teeth (Fig. 6.38a). Pereopods 3–5 dactyli hook-like (Fig. 6.45A)�������������������������������������������������������������������� Pliopontonia – Rostrum slender, reaching beyond antennular peduncle article 2; with dorsal and ventral teeth. Pereopods 3–5 dactyli biunguiculate (Fig. 6.45B)�������������������������������������������������������������������������������������������������������������������������������� 128 128. Rostrum slender, with lateral carina, with 7–9 dorsal and 1 ventral teeth (Fig. 6.36e)������������������������������������ Diapontonia – Rostrum without lateral carina, with 7 or 8 dorsal teeth (2 postorbital) and 2 distal ventral teeth (Fig. 6.38g)���������������������������������������������������������������������������������������������������������������������������������������������������������� Pseudoclimenes 129. Rostrum shorter than carapace, unarmed or with evenly spaced sharp dorsal teeth (Figs 6.37v, y, 6.38m)������������������ 130 – Rostrum as long as carapace, with 3 unevenly spaced dorsal teeth, 1 distal ventral tooth (Fig. 6.38e)���������� Poripontonia 130. Pleonite 6 posterolateral angle ending in slender spine ��������������������������������������������������������������������������������������������������������131 – Pleonite 6 posterolateral angle rounded or triangular���������������������������������������������������������������������������������������������������������� 132 131. Rostrum with 5 prominent dorsal teeth (Fig. 6.37m)���������������������������������������������������������������������������������������� Orthopontonia – Rostrum dorsally unarmed or with minute subdistal denticle (Fig. 6.38)��������������������������������������������������������������� Pontonia 132. Telson dorsal spiniform setae with one pair each in anterior and posterior halves (Fig. 6.42p)�������������������������������������133 – Telson with both pairs of dorsal spiniform setae in either anterior half or posterior half (Fig. 6.42f, i, n)������������������� 136 133. Pereopod 1 fingers less than half length of palm. Pereopod 2 palm laterally compressed, fixed finger usually with accessory tooth near tip (Fig. 6.44r)������������������������������������������������������������������������������������������������ Onycocaris – Pereopod 1 fingers as long as palm (Fig. 6.43c). Pereopod 2 palm subcylindrical, fixed finger with simple tip (Fig. 6.44p)������������������������������������������������������������������������������������������������������������������������������������������������������������ 134 134. Carapace antennal spine absent (Fig. 6.37j). Rostrum not reaching end of eyes. Pereopod 1 finger tips spatulate (Fig. 6.43c)������������������������������������������������������������������������������������������������������������������������������������������ Onycocaridella – Carapace antennal spine present (Fig. 6.37i). Rostrum not reaching to or beyond eyes. Pereopod 1 finger tips simple, pointed���������������������������������������������������������������������������������������������������������������������������������������������������������������� 135 135. Rostrum stout, thick, without dorsal, ventral or lateral carinae, suboval in section, reaching to end of eyes. Body and appendages setose (Fig. 6.35n). Pereopods 3–5 dactyli biunguiculate����������������������������������������������Bruceonia – Rostrum broadly triangular, reaching well beyond eyes. Body and appendages with few setae. Pereopods 3–5 dactyli simple or biunguiculate������������������������������������������������������������������������������������������������������������������������Odontonia 136. Telson with both pairs of dorsal spiniform setae in anterior half (Fig. 6.42f, i)�����������������������������������������������������������������137 – Telson with both pairs of dorsal spiniform setae in posterior half (Fig. 6.42n)�����������������������������������������������������������������145 137. Pereopod 2 unequal, one dactylus blunt, much longer than fixed finger (Fig. 6.44g). Pereopods 3–5 dactyli biunguiculate (Fig. 6.45m)����������������������������������������������������������������������������������������������������������������������� Exopontonia – Pereopod 2 equal or unequal, dactyli acute, as long as fixed finger. Pereopods 3–5 dactyli simple ­biunguiculate or multidentate���������������������������������������������������������������������������������������������������������������������������������������������� 138 138. Rostrum with 6 or more dorsal teeth (Fig. 6.36g) ����������������������������������������������������������������������������������������������������������������� 139 – Rostrum dorsally unarmed or with 2–3 teeth (Figs 6.35i, o, 6.36b, 6.38m)����������������������������������������������������������������������� 140

130

139. – 140. – 141. – 142. – 143. – 144. – 145. – 146. – 147. – 148. – 149. – 150. – 151. – 152. – 153. – 154. – 155. –

Marine Decapod Crustacea

Pereopod 3–5 dactylus less than 2 times as long as high��������������������������������������������������������������������������������� Periclimenoides Pereopod 3–5 dactylus more than 3 times as long as high������������������������������������������������������������������������������������ Epipontonia Rostrum reduced, not reaching beyond eyestalks, with ventral carina (Fig. 6.35o)����������������������������������������������� Cainonia Rostrum reaching beyond eyestalks������������������������������������������������������������������������������������������������������������������������������������������141 Rostrum with 2 or 3 dorsal teeth, lateral carinae not developed (Fig. 6.38m)���������������������������������������������������������Rostronia Rostrum without or with small subdistal dorsal tooth, lateral carinae developed����������������������������������������������������������� 142 Antennular article 1 medioventral tooth strongly developed (Fig. 6.40l)��������������������������������������������������������������Colemonia Antennular article 1 medioventral tooth small (Fig. 6.40j)���������������������������������������������������������������������������������������������������143 Pereopods 3–5 dactyli flexor margin with row of tubercles (Fig. 6.45j). (Rostrum with dorsal carina; Fig. 6.36b)������������������������������������������������������������������������������������������������������������������������������������������������������������������� Dactylonia Pereopods 3–5 dactyli flexor margin smooth������������������������������������������������������������������������������������������������������������������������� 144 Antennal scaphocerite without distolateral spine. Rostrum with dorsal carina (Fig. 6.37l)������������������������������� Opaepupu Antennal scaphocerite with distolateral spine. Rostrum without dorsal carina (Fig. 6.35i)�������������������������������� Ascidonia Pereopods 3–5 dactyli simple, hook-shaped. Rostrum linguiform to broadly triangular����������������������������������������������� 146 Pereopods 3–5 dactyli biunguiculate (Fig. 6.45c). Rostrum slender, sharply triangular��������������������������������������������������147 Rostrum linguiform, flat, distally rounded, without teeth. Carapace antennal spine absent (Fig. 6.37d)������������������������������������������������������������������������������������������������������������������������������������������������������������� Neoanchistus Rostrum distally laterally compressed, acute, usually with few denticles. Carapace antennal spine usually present��������������������������������������������������������������������������������������������������������������������������������������������������� Anchistus Rostrum not reaching beyond eyes, without dorsal teeth (Fig. 6.38c)������������������������������������������������������������ Pontoniopsides Rostrum reaching well beyond eyes, with obscure dorsolateral teeth in male only (Fig. 6.35b)��������������������Altopontonia Rostrum short, unarmed, with large blunt median tooth behind. Pterygostomial margin roundly produced forward beyond rostrum (Fig. 6.37a)����������������������������������������������������������������������������������������������Metapontonia Rostrum without large blunt tooth behind. Pterygostomial angle not produced forward����������������������������������������������149 Carapace antennal spine present���������������������������������������������������������������������������������������������������������������������������������������������� 150 Carapace antennal spine absent������������������������������������������������������������������������������������������������������������������������������������������������ 154 Rostrum broadly triangular, not reaching beyond eyes (Fig. 6.37y). Pereopods 2 subequal. Pereopods 3–5 dactyli simple�����������������������������������������������������������������������������������������������������������������������������Platypontonia Rostrum reaching eyes or beyond (Figs 6.35c, e, 6.38c)���������������������������������������������������������������������������������������������������������151 Pereopods 2 very unequal, major carpus cup-shaped, minor carpus slender��������������������������������������������������������������������152 Pereopods 2 sometimes unequal in size, symmetrical, both carpi cup-shaped�����������������������������������������������������������������153 Rostrum spiniform, cylindrical (Fig. 6.35c). Telson with 3 pairs of lateral setae equally spaced (Fig. 6.42b). Major pereopod 2 dactylus without flange, chela twice as long as carapace (Fig. 6.44a). Pereopod 1 fingers simple���������������������������������������������������������������������������������������������������������� Amphipontonia Rostrum triangular, with orbital shelf (Fig. 6.38d). Telson with both pairs of dorsal spiniform setae small, in posterior half. Major pereopod 2 dactylus with flange on upper half of lateral surface (Fig. 6.44x), chela not longer than carapace. Pereopod 1 fingers spatulate (Fig. 6.43e)���������������������������������������������������� Pontoniopsis Major pereopod 2 chela strongly compressed, twice as long as carapace (rostrum included) (Fig. 6.44i). Telson with pairs of spiniform setae submarginal, evenly spaced (Fig. 6.42l)�����������������������������������������������Isopontonia Major pereopod 2 chela not compressed, less than 1.5 times length of postorbital carapace. Telson with both pairs of dorsal spiniform setae in anterior third (Fig. 6.42c) or half���������������������������������������������� Anchiopontonia Body with deep postorbital depression formed by rostrum lateral margin and oblique ridge from lower orbital angle (Fig. 6.37x)��������������������������������������������������������������������������������������������������������������������������������������������� Platycaris Body without postorbital depression�����������������������������������������������������������������������������������������������������������������������������������������155 Pereopod 2 palm huge, palm compressed, wide, flexor margin convex, carinate, fixed finger 3 times as wide as dactylus (Fig. 6.44o)�������������������������������������������������������������������������������������������������������������������������� Notopontonia Pereopod 2 palm short, subcylindrical, fixed finger narrower than dactylus����������������������������������������������Pseudopontonia

6 – Caridea – shrimps

131

Gnathophyllid genera

Levicaris Bruce, 1973

Diagnosis of gnathophyllid group. Maxilliped 1 exopod caridean lobe not overreaching endite. Maxilliped 3 operculate, last 2 articles articulating, much narrower than fused squarish-oval proximal articles.

Diagnosis. Maxilliped 2 strong, pediform, merus and carpus elongate. Pereopods 3–5 dactyli simple. Telson posterior margin incised. Maximum cl. 4 mm. Intertidal, subtidal (0–5 m). Eastern Indo-Pacific (Hawaii). 1 species (Bruce 1973a).

Gnathophylloides Schmitt, 1933 Diagnosis. Maxilliped 2 shorter than maxilliped 3. Pereopods 3–5 dactyli simple. Maximum cl. 6 mm. Intertidal–shelf (0–94 m; associated with echinoids). Tropical Atlantic, Indo-West Pacific, Temperate Australasia. 2 species (Bruce 1973a).

Gnathophyllum Latreille, 1819

Pycnocaris Bruce, 1972 Diagnosis. Body depressed. Rostrum prominent, unarmed. Cornea narrower than eyestalk. Pereopods 3–5 dactyli broad, subtriangular, distal spiniform seta shorter than tooth on flexor margin. Maximum cl. 4.6 mm. Intertidal (associated with holothurians). Western IndoPacific (Chagos Archipelago). 1 species (Bruce 1972c).

Diagnosis. Rostrum with dorsal teeth only. Cornea wider than eyestalk, with papilla. Maxilliped 3 ischiomerus outer face convex. Pereopods 3–5 dactyli broad, subtriangular, distal tooth longer than tooth on flexor margin. Maximum cl. 5 mm (Pl. 10a). Intertidal–shelf (0–50 m). Cosmopolitan except in polar seas. 10 species (Ahyong 2003, 2015).

Diagnosis of hymenocerid group. Maxilliped 1 exopod caridean lobe not overreaching endite. Maxilliped 3 suboperculate, last 3 articles articulating, wider and articulating with elongate basis.

Gnathophylleptum d’Udekem d’Acoz, 2001

Hymenocera Latreille, 1819

Diagnosis. Cornea wider than eyestalk, with papilla. Scaphocerite 3 times as long as wide. Maxilliped 3 ischiomerus outer face concave. Maximum cl. 2.8 mm. Subtidal (15 m). Temperate Northern Atlantic (Canary Is). 1 species (d’Udekem d’Acoz 2001).

Hymenocerid genera

Harlequin shrimps Diagnosis. Antennular outer flagellum broad, leaf-like. Maxilliped 3 of 4 articles, last 3 expanded, foliaceous. Pereopod 2 chelae with flexor border produced as thin flap. Maximum cl. 10 mm (Pl. 10b).

Fig. 6.32.  Palaemonidae, gnathophyllid and hymenocerid genera. Anterior carapace, rostrum: a, Gnathophylloides; b, Gnathophyllum; c, Pycnocaris. Right antennular peduncle: d, Gnathophylleptum; e, Gnathophyllum; f, Hymenocera (with flagellum). Scaphocerite: g, Gnathophylleptum; h, Gnathophyllum. Maxilliped 1: i, Gnathophyllum. Maxilliped 2: j, Gnathophylloides; k, Levicaris. Maxilliped 3: l, Gnathophylleptum; m, Gnathophyllum (caridean lobe arrowed); n, Hymenocera. o, Phyllognathia. Pereopod 2: p, Hymenocera; q, Phyllognathia. Pereopod 3, distal articles: r, Gnathophyllum; s, Pycnocaris.

132

Marine Decapod Crustacea

Subtidal (1–20 m). Indo-West Pacific, Tropical Eastern Pacific. 1 species (Mitsuhashi et al. 2005: figures).

Phyllognathia Borradaile, 1915 Diagnosis. Antennular outer flagellum thread-like. Maxilliped 3 of 4 articles, foliaceous, second widest. Pereopod 2 chelae broad, flat. Maximum cl. 11 mm. Subtidal, shelf (2–50 m). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Australasia. 2 species (Fujino 1973b).

Palaemonine genera Diagnosis of palaemonine group. Maxilliped 1 exopod caridean lobe overreaching endite. Telson usually with 2 pairs of posterior spiniform setae, lateral pair shorter, 1 or more median pairs of plumose setae. Pleurobranch usually present at base of maxilliped 3.

Implicit generic characters. Carapace branchiostegal spine absent; hepatic spine absent. Pleonite 4 smooth dorsally; pleonite 5 pleuron posteroventrally rounded. Cornea well developed, pigmented. Mandibular molar cylindrical; palp of 3 articles. Pereopod 5 propodus without distal transverse rows of setae on flexor margin. Male pleopod 1 without or with rudimentary appendix interna. Anchistioides Paul’son, 1875 Diagnosis. Rostrum laterally compressed, with dorsal and ventral teeth, with pair of blunt postocular teeth. Mandibular molar distally flared; palp absent. Male pleopod 1 with well developed appendix interna. Maximum cl. 13 mm. Subtidal–shelf (9–68 m). Tropical W Atlantic, Western and Central Indo-Pacific. 3 species (Bruce 1991b; Chace and Bruce 1993; Fransen 2020: redescriptions, key to Indo-West Pacific species). The species are associated with sponges (Soledade et al. 2017).

Fig. 6.33.  Palaemonidae, palaemonine genera. Carapace, rostrum: a, Anchistioides; b, Arachnochium; c, Calathaemon; d, Creaseria; e, Leandrites; f, Leptocarpus; g, Leptopalaemon; h, Nematopalaemon; i, Pseudopalaemon; j, Rhopalaemon; k, Tenuipedium; l, Troglindicus; m, Urocaridella. Thoracic sternite 4, anterior margin: n, Rhopalaemon. Thoracic sternites 4, 5: o, Arachnochium; p, Tenuipedium. Male thoracic sternite 8, coxae 5: q, Macrobrachium. Pleon, posterior pleonites: r, Arachnochium; s, Cryphiops; t, Leptopalaemon; u, Macrobrachium; v, Palaemon; w, Rhopalaemon. Telson: x, Macrobrachium; y, Palaemon; z, Tenuipedium.

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Fig. 6.34.  Palaemonidae, palaemonine genera. Right antennular peduncle: a, Palaemon. Mandible: b, Anchistioides; c, Leander; d, Leandrites. Maxilla: e, Troglocubanus; f, Troglomexicanus. Maxilliped 1: g, Anchistioides; h, Troglocubanus; i, Troglomexicanus (caridean lobe arrowed). Pleopod 1, with appendices interna, masculina: j, Anchistioides; k, Leander. Pereopod 2, distal articles: l Cryphiops; m, Leptopalaemon; n, Troglindicus. Pereopod 3, distal articles: o, Brachycarpus; p, Neopalaemon; q, Rhopalaemon. I, incisor; M, molar.

Arachnochium Wowor & Ng, 2010

Creaseria Holthuis, 1950

Diagnosis. Rostrum without elevated basal crest. Carapace branchiostegal groove running from base of hepatic spine to carapace margin; hepatic spine present. Male thoracic sternite 8 without anterior lobes. Pereopods 3–5 not subequal in length, pereopod longer than pereopod 2. Pereopod 5 propodus with distal transverse rows of setae on flexor margin. Maximum cl. 12 mm. Estuarine, freshwater. Ganges Delta to S Vietnam. 2 species (Wowor and Ng 2010).

Diagnosis. Rostrum without elevated basal crest. Carapace branchiostegal spine present. Cornea without pigment. Antennular peduncle article 1 distal margin concave, anterolateral spine strong. Mandibular palp of 2 articles. Pereopod 5 propodus with distal transverse rows of setae on flexor margin. Maximum cl. 15 mm. Marine anchialine caves. Yucatan, Mexico. 1 species (ChávezSolís et al. 2018; Holthuis 1952).

Brachycarpus Bate, 1888

Cryphiops Dana, 1852

Diagnosis. Rostrum without elevated basal crest. Carapace branchiostegal groove present; hepatic spine present. Pereopods 3–5 dactyli biunguiculate; pereopod 5 propodus with 1 distal transverse row of setae on flexor margin. Male pleopod 1 with well developed appendix interna. Maximum cl. 15 mm (Pl. 10c). Marine, intertidal–shelf (0–56 m). Temperate Northern and Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific. 3 species (Bruce 1998b: key to species).

Diagnosis. Rostrum without elevated basal crest. Pereopod 5 propodus with distal transverse rows of setae on flexor margin. Maximum cl. 40 mm. Freshwater (rivers, caves). Chiapas, Mexico; Chile; Brazil. 6 species (Baldari et al. 2010: four species from Chiapas; Villalobos Hiriart et al. 1989: key to five species).

Calathaemon Bruce & Short, 1993 Diagnosis. Rostrum longer than scaphocerite, with dorsal teeth only. Carapace branchiostegal spine present. Pleonite 5 pleuron posteroventrally square. Cornea without pigment. Mandibular palp absent. Pereopod 5 propodus with distal transverse rows of setae on flexor margin. Maximum cl. 5 mm. Freshwater (caves). TX, USA. 1 species (Short et  al. 2013: rediagnosis).

Leander Desmarest, 1849 Diagnosis. Rostrum without elevated basal crest. Carapace branchiostegal spine present. Mandibular palp of 2 articles. Male pleopod 1 with well developed appendix interna. Maximum cl. 8 mm (Pl. 10d, f). Freshwater, estuarine, marine intertidal–shelf (0–55 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Australasia (L. tenuicornis also on floating algae in oceanic environments). 5 species (Bruce 1994: key to four species).

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Leandrites Holthuis, 1950

Neopalaemon Hobbs, 1973

Diagnosis. Rostrum without elevated basal crest. Carapace branchiostegal spine present. Pleonite 4 with slender median process. Mandibular palp absent. Male pleopod 1 with well developed appendix interna. Maximum cl. 10 mm. Estuarine, intertidal–shelf (0–56 m). Temperate Northern Pacific, Western and Central Indo-Pacific. 4 species (Bruce 2000a: distribution; Chace and Bruce 1993: key to species).

Diagnosis. Rostrum without elevated basal crest. Carapace branchiostegal groove present; hepatic spine present. Cornea without pigment. Mandibular palp of 2 articles. Maximum cl. 11 mm. Marine anchialine caves. Caribbean. 1 species (Hobbs 1973).

Leptocarpus Holthuis, 1950 Diagnosis. Rostrum with elevated basal toothed crest. Carapace branchiostegal groove present. Pleonite 4 with slender median process. Maximum cl. 10 mm. Estuarine, freshwater. Western Indo-Pacific (India). 3 species (Jayachandran 1992: key to species).

Leptopalaemon Bruce & Short, 1993 Diagnosis. Rostrum without elevated basal crest. Pleonite 5 pleuron posteroventrally square. Mandibular palp of 1 article, or absent. Pereopod 5 propodus with distal transverse rows of setae on flexor margin. Maximum cl. 8.5 mm. Freshwater. NT, Australia. 5 species (Short et al. 2013: rediagnosis, key to species).

Macrobrachium Bate, 1868 Diagnosis. Rostrum without elevated basal crest. Carapace branchiostegal groove present; hepatic spine present. Male thoracic sternite 8 with anterior lobes. Pereopod 5 propodus with distal transverse rows of setae on flexor margin. Telson, posterior margin with more than 5 pairs of plumose setae. Maximum cl. 100 mm. Freshwater, estuarine, marine, anchialine caves. Cosmopolitan except polar seas. 262 species (~16 marine or estuarine). Macrobrachium is largely a freshwater genus that includes important edible species such as the giant freshwater prawn, M. rosenbergi (see, e.g. Banu and Christianus 2016; Wowor and Choy 2001). Some species rely on a period as juveniles in estuaries to complete their life history (Fukuda et al. 2017). Neotropical and West African clades of species, of which some spend time in estuaries, appear genetically isolated from the rest of the genus (Anger 2013). Twelve species are stygobitic, at least one in an anchialine environment (Fujita et al. 2015; Makombu et al. 2019).

Nematopalaemon Holthuis, 1950 Diagnosis. Rostrum elongate, thin, with elevated basal toothed crest. Carapace branchiostegal spine present. Pereopod 5 propodus with distal transverse rows of setae on flexor margin. Maximum cl. 14 mm. Intertidal–subtidal (0–20 m). Tropical Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate Southern Africa. 5 species (Ashelby et al. 2018: figures; Chace and Bruce 1993: key to 5 species).

Palaemon Weber, 1795 Diagnosis. Rostrum with elevated basal toothed crest, or without elevated basal crest. Carapace branchiostegal spine present; branchiostegal groove present. Pleonite 4 with slender median process; 5 pleuron posteroventrally square. Mandibular palp of 2 or 3 articles, or absent. Pereopod 5 propodus with distal transverse rows of setae on flexor margin. Maximum cl. 13 mm (Pl. 10c). Freshwater, estuarine, marine, intertidal, subtidal (0–26 m). Cosmopolitan except in polar seas. 88 species (c. 33 marine, another 11 brackish, rest freshwater). Half of the species are confined to fresh water, the other half to estuarine or marine environments, some being amphidromous (Carvalho et al. 2017; De Grave and Ashelby 2013: list of species with distributions and ecology). The genus may not be monophyletic (Ashelby et al. 2012; De Grave and Ashelby 2013), However, the phylogeny of the genus suggests three lineages based on molecular relationships or four monophyletic groups based on morphology (Carvalho et al. 2017).

Pseudopalaemon Sollaud, 1911 Diagnosis. Rostrum without elevated basal crest. Carapace branchiostegal groove present; hepatic spine present. Mandibular palp absent. Pereopod 5 propodus with distal transverse rows of setae on flexor margin (usually). Maximum cl. 7 mm. Freshwater, estuarine. W South American rivers and lakes. 7 species (García-Dávila and Magalhães 2003: 7 species compared; Kensley and Walker 1982: key to 5 species).

Rhopalaemon Ashelby & De Grave, 2010 Diagnosis. Rostrum without elevated basal crest. Carapace branchiostegal spine present; branchiostegal groove present. Pleonite 4 with slender median process. Pereopods 3–5 propodi expanded along flexor margin, with row of spiniform setae. Maximum cl. 10 mm. Intertidal. Western Indo-Pacific (India). 1 species (Ashelby and De Grave 2010).

Tenuipedium Wowor & Ng, 2010 Diagnosis. Rostrum without elevated basal crest. Carapace branchiostegal groove running below hepatic spine from margin to behind hepatic spine; hepatic spine present. Male thoracic sternite 8 without anterior lobes. Pereopod 5 propodus with distal transverse rows of setae on flexor margin. Maximum cl. 5.7 mm. Freshwater. Sumatra, Indonesia. 1 species (Wowor and Ng 2010).

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Fig. 6.35.  Palaemonidae, pontoniine genera. Carapace, rostrum: a, Actinimenes; b, Altopontonia; c, Amphipontonia; d, Anapontonia; e, Anchiopontonia; f, f’, Ancylocaris (female, male); g, Apopontonia; h, Araiopontonia; i, i’, Ascidonia; j, Balssia; k, Bathymenes; l, Blepharocaris; m, Brucecaris; n, Bruceonia; o, Cainonia; p, Carinopontonia; q, Chacella; r, Climeniperaeus; s, Colemonia; t, Conchodytes; u, Coralliocaris; v, Crinotonia; w, Cristimenes; x, Ctenopontonia

Troglindicus Sankolli & Shenoy, 1979 Diagnosis. Rostrum short, without dorsal teeth. Cornea with small pigment spot. Mandibular palp absent. Maximum cl. 8 mm. Freshwater, groundwater (wells). India. 1 species (Sankolli and Shenoy 1979).

Troglocubanus Holthuis, 1949 Diagnosis. Rostrum longer than scaphocerite, with dorsal teeth only, or short, with or without dorsal teeth. Carapace branchiostegal groove present. Cornea without pigment. Maxilla endite bilobed, maxilliped 1 epipod rounded. Mandibular palp absent. Maximum cl. 12 mm. Freshwater (pools, caves). Caribbean islands. 5 species (Holthuis 1952: key to 4 species).

Troglomexicanus Villalobos, Alvarez & Iliffe, 1999 Diagnosis. Rostrum short, with or without teeth. Cornea without pigment. Maxilla endite simple, maxilliped 1 epipod bilobed. Mandibular palp absent. Maximum cl. 8 mm.

Freshwater (caves). Mexico. 3 species (Villalobos et  al. 1999: key to species).

Urocaridella Borradaile, 1915 Diagnosis. Rostrum with 2 strong basal teeth, crest-like. Carapace with dorsal tooth at midpoint; branchiostegal spine present; branchiostegal groove present. Mandibular palp of 2 articles, or absent. Male pleopod 1 with well developed appendix interna. Maximum cl. 6.4 mm. Intertidal–shelf (0–130 m). Western and Central Indo-Pacific. 8 species (Prakash and Baeza 2018). One species of Urocaridella has invaded the Mediterranean from the Red Sea. Some species are known fish cleaners (Bos and Fransen 2018).

Pontoniine genera Diagnosis of pontoniine group. Maxilliped 1 exopod caridean lobe overreaching endite. Telson usually with 3 pairs of posterior spiniform setae (median pair often plumose). Pleurobranch usually absent from base of maxilliped 3.

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Marine Decapod Crustacea

Fig. 6.36.  Palaemonidae, pontoniine genera. Carapace, rostrum continued: a, Cuapetes; b, Dactylonia; c, Dasella; d, Dasycaris; e, Diapontonia; f, Echinopericlimenes; g, Epipontonia; h, Exopontonia; i, Fennera; j, Hamiger; k, Hamodactylus; l, Hamopontonia; m, Harpiliopsis; n, Holthuisaeus; o, Ischnopontonia; p, Izucaris; q, Jocaste; r, Kaviengella; s, Laomenes; t, Leptomenaeus; u, u’, Lipkemenes; v, Madangella; w, Manipontonia; x, Margitonia; y, Mesopontonia.

Fig. 6.37.  Palaemonidae, pontoniine genera. Carapace, rostrum continued: a, Metapontonia; b, Michaelimenes; c, Miopontonia; d, Neoanchistus; e, Neoclimenes; f, Neopericlimenes; g, Neopontonides; h, Nippontonia; i, i’, Odontonia; j, Onycocaridella; k, Onycocaridites; l, Opaepupu; m, Orthopontonia; n, Palaemonella; o, o’, Paraclimenaeus; p, Paraclimenes; q, Paranchistus; r, Paratypton; s, Patonia; t, Periclimenes; u, Plesiopontonia; v, Philarius; w, Phycomenes; x, x’, Platycaris; y, y’, Platypontonia.

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Fig. 6.38.  Palaemonidae, pontoniine genera. Carapace, rostrum continued: a, Pliopontonia; b, Pontonia; c, Pontoniopsides; d, Pontoniopsis; e, Poripontonia; f, Propontonia; g, Pseudoclimenes; h, h’, Pseudocoutierea; i, i’, Pseudopontonides; j, Pseudoveleronia; k, Rapimenes; l, Rapipontonia; m, Rostronia; n, Sandyella; o, Stegopontonia; p, Tectopontonia; q, Thaumastocaris; r, Tuleariocaris; s, Typton; t, Typtonoides; u, u’, Typtonomenaeus.

Fig. 6.39.  Palaemonidae, pontoniine genera. Carapace, rostrum continued: a, Typtonychus; b, b’, Unesconia; c, Unguicaris; d, Urocaris; e, Veleronia; f, Veleroniopsis; g, Vir; h, Waldola; i, Yemenicaris; j, Zoukaris.

Implicit generic attributes. Rostrum blade-like, with dorsal and ventral teeth. Carapace antennal spine present; branchiostegal spine absent; hepatic spine absent. Pleonites 1–5 rounded. Mandibular palp absent. Maxillipeds 1–3 with exopods. Pereopods 1 equal. Pereopods 3–5 dactyli simple, without basal protuberance. Telson with 2 pairs of dorsal spiniform setae.

Actinimenes Ďuriš & Horka, 2017 Diagnosis. Rostrum with 7–10 dorsal teeth (posterior tooth behind orbits), 0–2 ventral teeth. Carapace hepatic spine almost level with antennal spine. Thoracic sternite 4 with broad transverse ridge, with median incision. Pereopods 3–5 dactyli with stout unarmed corpus, unguis elongate, curved. Maximum cl. 3.1 mm.

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Marine Decapod Crustacea

Fig. 6.40.  Palaemonidae, pontoniine genera. Thoracic sternite 4, anterior margin: a, Phycomenes. Thoracic sternite 5, anterior lobes: b, Exoclimenella. Carapace, pleon: c, Ancylomenes. Pleon, posterior pleonites: d, Coutierea; e, Harpiliopsis; f, Lipkebe. Pleonite 6, dorsal: g, Orthopontonia. Right antennular peduncle: h, Ancylomenes; i, Crinotonia; j, Dactylonia; k, Leptomenaeus (pair, rostrum, eyes); l, Odontonia; m, Veleronia. Antenna, scaphocerite: n, Paratypton. Epistome, labrum: o, Laomenes; p, Onycomenes.

Fig. 6.41.  Palaemonidae, pontoniine genera. Mandible: a, Araiopontonia; b, Holthuisaeus; c, Periclimenaeus; d, Periclimenoides; e, Unguicaris. Maxilliped 1: f, Onycomenes; g, Typtonychus (caridean lobe arrowed). Maxilliped 3: h, Mesopontonia; i, Propontonia. I, incisor; M, molar.

Subtidal (1–18 m; associated with sea anemones). Western and Central Indo-Pacific. 3 species (Ďuriš and Horká 2017: key to species).

Allopontonia Bruce, 1972 Diagnosis. Rostrum with 7–10 dorsal teeth, 1 ventral tooth. Carapace hepatic spine movable. Pereopod 2 carpus about third length of chela. Pereopods 3–5 propodi with spiniform setae on flexor margin. Maximum cl. 2.8 mm. Subtidal, shelf (10–105 m; associated with echinoids). Central Indo-Pacific. 2 species (Bruce 1972a, 2010b).

Altopontonia Bruce, 1990 Diagnosis. Rostrum slender, with obscure dorsal teeth. Pereopods 3–5 dactyli biunguiculate. Telson with both pairs of dorsal spiniform setae in posterior half. Maximum cl. 3.3 mm. Slope (350–503 m; possibly associated with echinoids). Central Indo-Pacific (New Caledonia). 1 species (Bruce 1990).

Amphipontonia Bruce, 1991 Diagnosis. Rostrum cylindrical, curving downwards. Pereopods 2 very unequal, major carpus cup-shaped, minor carpus slender.

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Fig. 6.42.  Palaemonidae, pontoniine genera. Telson: a, Actinimenes; b, Amphipontonia; c, Anchiopontonia; d, Ancylocaris; e, Araiopontonia; f, Ascidonia; g, Coutierea; h, Echinopericlimenes; i, Exopontonia; j, Hamopontonia; k, Holthuisaeus; l, Isopontonia; m, Lipkebe; n, Neoanchistus; o, Nippontonia; p, Onycocaris; q, Periclimenaeus; r, Periclimenes; s, Plesiopontonia; t, Pontonia; u, Propontonia. Telson, right uropod: v, Anapontonia; w, Apopontonia; x, Ischnopontonia; y, Paraclimenaeus; z, Pontonides (exopod suture only); A, Unesconia; B, Veleronia; C, Yemenicaris.

Telson with 3 pairs of lateral setae equally spaced. Maximum cl. 8 mm. Slope (300–365 m; probably associated with antipatharian corals). Central Indo-Pacific. 2 species (Bruce 1991a; Ďuriš and Šobáňová 2020).

Anapontonia Bruce, 1966 Diagnosis. Body rather stout. Rostrum steep, with ~15 small dorsal teeth from tip to middle carapace. Uropodal exopod anterior margin spinose. Maximum cl. 3.2 mm.

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Marine Decapod Crustacea

Fig. 6.43.  Palaemonidae, pontoniine genera. Pereopod 1, distal articles: a, Allopontonia; b, Hamodactyloides; c, Onycocaridella; d, Periclimenella; e, Pontoniopsis; f, Pseudoveleronia; g, Rapimenes; h, Rapipontonia; i, Thaumastocaris; j, Zenopontonia.

Intertidal, subtidal (0–1 m; associated with corals). Western and Central Indo-Pacific. 1 species (Bruce 1966b, 1967).

Anchiopontonia Bruce, 1992 Diagnosis. Rostrum cylindrical, with minute distal teeth. Pereopods 2 unequal. Pereopods 3–5 dactyli with tip widened, truncate. Telson with 3 pairs of lateral setae equally spaced. Maximum cl. 5 mm (Pl. 10g). Subtidal (8–30 m; associated with bivalves, Spondylus). Central and Eastern Indo-Pacific. 1 species (Bruce 1992).

Anchistus Borradaile, 1898 Diagnosis. Rostrum distally laterally compressed, usually with few small teeth. Telson with both pairs of dorsal spiniform setae in posterior half. Maximum cl. 8 mm (Pl. 10h). Intertidal–shelf (0–110 m; associated with bivalves). Indo-West Pacific, Temperate Australasia. 7 species (Bruce 1977; Chace and Bruce 1993: key to species).

Ancylocaris Schenkel, 1902 Diagnosis. Rostrum blade-like, with dorsal teeth, 1 or 2 distoventral teeth, carapace prominently humped in adult female. Carapace hepatic spine present. Pereopods 3–5 dactyli with small distal tooth on stout corpus, unguis elongate, curved. Telson with 2 pairs of minute dorsal spines on posterior third. Maximum cl. 8.5 mm (Pl. 10i). Intertidal, subtidal (0–30 m; associated with sea anemones). Indo-West Pacific. 1 species (Ďuriš and Horká 2017: diagnosis; Kemp 1922: description, figures of only A. brevicarpalis).

Ancylomenes Okuno & Bruce, 2010 Diagnosis. Rostrum usually arched, dorsal margin with 7–13 teeth, with 0 or few ventral teeth. Carapace inferior orbital margin strongly produced, with reflected inner ventral flange; hepatic spine present. Antennular peduncle article 1 with large anterolateral lobe. Maximum cl. 7 mm (Pl. 10j).

Intertidal–shelf (0–181 m; associated with cnidarians, fish-cleaners). Temperate Northern Pacific, Tropical Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate Southern Africa. 22 species (Okuno and Bruce 2010: key to 18 species).

Anisomenaeus Bruce, 2010 Diagnosis. Rostrum downcurved, with 9 dorsal teeth, 1 distoventral tooth. Pereopods 2 unequal, similar, dactylus entire, fitting into longitudinal depression on finger; major palm densely spinose, fixed finger longitudinally grooved, dactylus spinose, cutting edge entire. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 3.1 mm. Shelf (70 m; associated with sponges). Western Indo-Pacific (Somalia). 1 species (Bruce 2010c).

Apopontonia Bruce, 1976 Diagnosis. Rostrum with supraocular tooth, triangular, with 5 dorsal teeth, depressed tip. Uropodal exopod anterior part of suture lobed, with spiniform setae. Maximum cl. 2 mm. Subtidal–shelf (10–73 m; associated with sponges). Western and Central Indo-Pacific. 1 species (Bruce 1976b).

Araiopontonia Fujino & Miyake, 1970 Diagnosis. Rostrum laterally compressed, with dorsal and ventral teeth. Mandibular incisor wider distally. Pereopod 3 dactylus tip comprising tooth, unguis, spiniform seta. Telson posterior margin, without spiniform seta. Maximum cl. 2.6 mm. Subtidal to at least 5 m (associated with crinoids). Central Indo-Pacific. 1 species (Bruce 1983: figures; Fujino and Miyake 1970: diagnosis, figures).

Ascidonia Fransen, 2002 Diagnosis. Rostrum dorsally unarmed or with 1 subdistal tooth, dorsal carina absent, lateral carinae well developed, ventral margin unarmed. Carapace antennal spine merged with infraorbital

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Fig. 6.44.  Palaemonidae, pontoniine genera. Pereopod 2, distal articles: a, Amphipontonia; b, Anisomenaeus; c, Climeniperaeus; d, Crinotonia; e, Cristimenes; f, Eupontonia; g, Exopontonia; h, Hamiger (major); i, Isopontonia; j, Leptomenaeus; k, Madangella; l, Mesopontonia; m, Michaelimenes; n, Miopontonia; o, Notopontonia; p, Onycocaridella; q, Onycocaridites; r, Onycocaris; s, Paraclimenaeus; t, t’, Paranchistus (major); u, Periclimenella; v, Plesiomenaeus; w, Pontonides (male); x, Pontoniopsis; y, Propontonia; z, Tectopontonia; A, Typtonomenaeus; B, Typtonychus (minor); C, Veleronia; D, Vir.

angle, acute. Thoracic sternite 4 anteriorly produced as triangular plate. Antennular peduncle medioventral tooth small. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 7.3 mm. Intertidal–shelf (0–100 m; associated with ascidians). Temperate Northern Atlantic, Tropical Eastern Pacific. 5 species (Fransen 2002: key to species).

Balssia Kemp, 1922 Diagnosis. Rostrum with dorsal teeth only. Carapace with oblique supraorbital dentate carina. Pleonites 1–3 pleura rounded, 4, 5 with posteroventral sharp tooth. Maxillipeds 1–3 without exopods. Maximum cl. 4 mm.

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Marine Decapod Crustacea

Fig. 6.45.  Palaemonidae, pontoniine genera. Pereopod 3, distal articles: a, Actinimenes; b, Allopontonia; c, Altopontonia; d, Anchiopontonia; e, Araiopontonia; f, Bathymenes; g, Brucecaris; h, Coralliocaris; i, Crinotonia; j, Dactylonia; k, Dasella; l, Echinopericlimenes; m, Exopontonia; n, Fennera; o, Hamodactyloides; p, Hamodactylus; q, Harpilius; r, Isopontonia; s, Jocaste; t, Margitonia; u, Miopontonia; v, Onycocaridites; w, Onycocaris; x, Paranchistus; y, Philarius; z, Phycomenes; A, Pliopontonia; B, Pseudoclimenes; C, Rapimenes; D, Sandimenes; E, Unguicaris; F, Zenopontonia; G, Zoukaris.

Subtidal, shelf (12–120 m; associated with gorgonians and alcyonarians). Temperate Northern Atlantic, Temperate Australasia. 3 species (Bruce 2004a: key to species).

Bathymenes Kou, Li & Bruce, 2016 Diagnosis. Rostrum with most posterior tooth clearly separate from others, epigastric. Carapace hepatic spine present. Cornea

reduced, without accessory pigment spot. Pereopod 2 granulate, fingers with lateral flange; cutting edge with basal tooth-like process. Pereopods 3–5 dactyli biunguiculate. Telson with 3–7 pairs of dorsolateral spiniform setae. Maximum cl. 10.9 mm. Slope (187–824 m; some associated with sea anemones, echinoids, pennatulaceans). Indo-West Pacific, Tropical Eastern Pacific. 15 species (Kou et al. 2016a: distribution, key to species; Li and Bruce 2006: descriptions of 5 species).

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Blepharocaris Mitsuhashi & Chan, 2007

Climeniperaeus Bruce, 1995

Diagnosis. Rostrum with dorsal teeth only, extending on to carapace. Carapace orbital rim round, lower half laminar, raised laterally, separated by notch from leaf-like infraorbital angle; hepatic spine submarginal below antennal spine. Maximum cl. 2.7 mm. Subtidal (20 m; associated with scleractinian corals). Central Indo-Pacific (Philippines). 1 species (Mitsuhashi and Chan 2007).

Diagnosis. Rostrum with 6 dorsal teeth. Pereopods 2 unequal; pereopod 2 major chela fingers with opposing fossae. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 3.5 mm. Subtidal, shelf (2–97 m; associated with sponges). Indo-West Pacific. 3 species (Bruce 1996: figures; Odijk and Fransen 2017: key to species).

Brucecaris Marin & Chan, 2006

Colemonia Bruce, 2005

Diagnosis. Rostrum with 6 similar prominent dorsal teeth, without ventral teeth. Carapace hepatic spine present. Pereopod 2 fingers 3 times as long as palm. Pereopods 3–5 propodi with row of long distal setae extending nearly to base of dactylus unguis. Maximum cl. 2.5 mm. Subtidal (8–17 m; associated with crinoids). Western and Central Indo-Pacific. 1 species (Marin and Chan 2006).

Diagnosis. Rostrum short, acute, unarmed, dorsal carina distinct, lateral carinae narrow. Carapace pterygostomial region broadly rounded. Maximum cl. 2.2 mm. Subtidal (12 m; associated with ascidians). Central IndoPacific (Papua New Guinea). 1 species (Bruce 2005b).

Bruceonia Fransen, 2002

Diagnosis. Body strongly depressed. Rostrum short, stout, triangular, thickened, toothless. Pereopods 3–5 dactyli with rounded basal protuberance. Maximum cl. 9 mm (Pl. 10k). Intertidal–shelf (0–40 m; association with large bivalves, including giant clams). Temperate Northern Pacific, Western and Central Indo-Pacific. 11 species (Fransen 1994: figures of 4 species).

Diagnosis. Rostrum short, blunt, depressed, dorsally unarmed, dorsal carina absent, lateral carinae well developed, ventral margin unarmed. Antennular peduncle medioventral tooth small. Pereopods 3–5 dactyli biunguiculate, unguis with patch of distal scales. Maximum cl. 5.2 mm. Subtidal (18–21 m; associated with molluscs). Central IndoPacific (Great Barrier Reef, Australia). 1 species (Fransen 2002).

Cainonia Bruce, 2005 Diagnosis. Rostrum very short, acute, unarmed, dorsal carina feebly developed, lateral carinae broad, not reaching beyond eyestalks. Pereopods 3–5 dactyli biunguiculate, with denticles on flexor margin. Maximum cl. 6.1 mm. Subtidal (2–5 m; associated with molluscs, Spondylus). Central and Eastern Indo-Pacific. 1 species (Bruce 2005b).

Carinopontonia Bruce, 1988

Conchodytes Peters, 1852

Coralliocaris Stimpson, 1860 Diagnosis. Body strongly depressed. Rostrum with lateral carinae proximally expanded as supraocular eave, compressed distally, with 1–5 dorsal teeth, 0–2 ventral teeth. Pereopods 3–5 dactyli with hoof-shaped basal protuberance. Maximum cl. 4.1 mm (Pl. 10l). Intertidal, shelf (0–54 m; associated with corals). Indo-West Pacific. 11 species (Mitsuhashi and Takeda 2008: key to 10 species).

Coutierea Nobili, 1901

Diagnosis. Rostrum long, without teeth, continued posteriorly to posterior margin as high carina. Carapace hepatic spine present. Maximum cl. 2.2 mm. Shelf (83 m). Central Indo-Pacific (NW Australia). 1 species (Bruce 1988b).

Diagnosis. Rostrum prominent, unarmed. Carapace with supraorbital tooth beside rostrum; hepatic spine present. Pleonites 2–6 pleura with posteroventral sharp tooth. Maxillipeds 2, 3 without exopod. Telson posterior margin with 2 large immovable teeth. Maximum cl. 7.5 mm. Shelf (148–172 m). Tropical W Atlantic. 1 species (Holthuis 1978).

Chacella Bruce, 1986

Crinotonia Marin, 2006

Diagnosis. Carapace with 2 dorsal median horns in anterior quarter, small knob posterior to orbit. Pleonites without dorsal horns. Rostrum short, broad, truncate, not reaching end of eyes, overhanging orbits, without lateral teeth, with high, blunt dorsal tooth, another tooth behind. Pleonites 3–5 pleura produced as narrow blunt projection or with posteroventral sharp tooth. Maxillipeds 2, 3 without exopod. Subtidal–shelf (13–30 m; associated with antipatharian). Tropical Eastern Pacific. 1 species (Bruce 1986a).

Diagnosis. Rostrum with 2–4 dorsal teeth, without ventral teeth. Carapace hepatic spine very small. Antennular peduncle article 1 with more than 1 distolateral tooth, mesial ventral tooth robust. Pereopods 2 unequal, major fixed finger with 2 proximal teeth fitting into 2 sockets on dactylus. Pereopods 3–5 dactyli biunguiculate; pereopod 3 dactylus with slender, curved unguis. Maximum cl. 5.5 mm. Subtidal–shelf (10–40 m; associated with crinoids). Central Indo-Pacific. 2 species (Marin 2008b).

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Marine Decapod Crustacea

Cristimenes Ďuriš & Horka, 2017 Diagnosis. Rostrum with 2–4 dorsal teeth, without ventral teeth. Carapace hepatic spine very small. Antennular peduncle article 1 with more than 1 distolateral tooth, mesial ventral tooth robust. Pereopods 2 unequal, major fixed finger with 2 proximal teeth fitting into 2 sockets on dactylus. Pereopods 3–5 dactyli biunguiculate; pereopod 3 dactylus with slender, curved unguis. Maximum cl. 5.5 mm. Intertidal–shelf (0–40 m; associated with crinoids or echinoids). Temperate Northern Pacific, Western and Central IndoPacific. 4 species (Ďuriš and Horká 2017: key to 3 species; Park et al. 2019: fourth species).

Subtidal, shelf (4–130 m; associated with antipatharians). Western and Central Indo-Pacific. 4 species (Bruce 1973c: key to species).

Diapontonia Bruce, 1986 Diagnosis. Rostrum with narrow lateral carinae, reaching beyond antennular peduncle article 2; with 8 dorsal and 1 ventral teeth, with 2 or 3 middorsal teeth behind orbit. Pereopod 2 fingers more than half as long as palm. Maximum cl. 5.1 mm. Slope (204–309 m; associated with echinoids). Tropical Atlantic (Bahamas). 1 species (Bruce 1986b).

Ctenopontonia Bruce, 1979

Echinopericlimenes Marin & Chan, 2014

Diagnosis. Rostrum triangular, with lateral carinae, acute, unarmed. Orbit posterior margin with comb-like row of c. 20 spines. Maxillipeds 1–3 without exopods. Maximum cl. 2.5 mm. Subtidal (5–15 m; associated with corals). Central Indo-Pacific (Eniwetok Atoll). 1 species (Bruce 1979a).

Diagnosis. Rostrum long, slender, with dorsal teeth, ventral lamina weak. Carapace hepatic spine long, overlapping anterior carapace margin. Cornea large, oval. Pereopod 2 cutting edge with basal tooth-like process. Pereopods 3–5 dactyli truncate subdistally, with denticulate lobe on flexor margin. Maximum cl. 9.8 mm. Shelf, slope (45–1280 m; associated with echinoids). Temperate Northern Pacific, Central Indo-Pacific. 4 species (Ďuriš and Šobáňová 2020: phylogeny of species based on COI; Marin and Chan 2014: key to species).

Cuapetes Clark, 1919 Diagnosis. Carapace hepatic spine present. Thoracic sternite 4 with slender median spine. Pereopods 3–5 propodi with spiniform setae on flexor margin. Maximum cl. 4 mm. Intertidal–shelf (0–191 m; most free-living, some associated with corals, hydroids or algae, including floating Sargassum). Tropical Atlantic, Indo-West Pacific, Temperate ­ Australasia. Migrant to Mediterranean. 29 species (Bruce 2004c: key to 24 species of Kemponia, junior synonym).

Epipontonia Bruce, 1977 Diagnosis. Rostrum with evenly spaced sharp dorsal teeth, 1 distal ventral tooth. Carapace infraorbital spine usually present. Telson with 2 pairs of dorsal robust setae in anterior half. Maximum cl. 3.4 mm. Subtidal (6–18 m; associated with sponges). Indo-West Pacific. 4 species (Bruce 2004b: key to species).

Dactylonia Fransen, 2002 Diagnosis. Rostrum dorsally unarmed; dorsal carina bordered by lateral depressions, lateral carinae well developed. Antennular peduncle medioventral tooth small. Pereopods 3–5 dactyli biunguiculate, with denticles on flexor margin (unguis with patch of distal scales). Maximum cl. 3.3 mm. Subtidal, shelf (2–185 m; associated with ascidians). Western and Central Indo-Pacific. 8 species (Fransen 2002: key to 6 species).

Dasella Lebour, 1945 Diagnosis. Rostrum prominent, unarmed. Carapace hepatic spine movable. Pereopods 3–5 dactyli with rounded basal protuberance. Maximum cl. 3.1 mm. Subtidal (3–27 m; inside ascidians). Western and Central IndoPacific, Temperate Australasia. 3 species (Berggren 1990: 3 species compared).

Dasycaris Kemp, 1922 Diagnosis. Rostrum slender, styliform, with 0–2 dorsal teeth, without ventral teeth. Carapace hepatic spine present. Maximum cl. 4.3 mm.

Eupontonia Bruce, 1971 Diagnosis. Carapace supraorbital ridges present. Pleonite 4 with slender median spine. Cornea narrower than eyestalk. Mandibular palp of 1 or 2 articles. Pereopod 2 carpus slender, longer than the palm. Pereopods 3–5 propodi with spiniform setae on flexor margin. Maximum cl. 3.2 mm. Intertidal, shallow subtidal (0–5 m; associated with corals). Indo-West Pacific. 4 species (Marin 2014b: key to species).

Exoclimenella Bruce, 1995 Diagnosis. Carapace hepatic spine present. Thoracic sternite 5 with pair of submedian processes. Pereopod 1 chela with fingers longer than palm, subspatulate laterally, with transparent distolateral lamina. Maximum cl. 3.4 mm. Subtidal–shelf (9–60 m). Indo-West Pacific. 4 species (Ďuriš and Bruce 1995: key to species as petitthouarsii-species group).

Exopontonia Bruce, 1988 Diagnosis. Rostrum short, tapering, with 2 small dorsal teeth. Pereopods 2 unequal, one dactylus blunt, much longer than fixed

6 – Caridea – shrimps

finger. Pereopods 3–5 dactyli biunguiculate. Telson with 2 pairs of lateral setae in anterior half. Maximum cl. 2 mm. Intertidal–shelf (0–55 m). Central Indo-Pacific. 1 species (Bruce 1988a).

Fennera Holthuis, 1951 Diagnosis. Body strongly depressed. Rostrum with lateral carinae proximally expanded as supraocular eave, compressed distally, with 3 dorsal teeth, without ventral teeth. Carapace postorbital ridge with 3–5 spines in oblique row behind antennal spine. Mandibular incisor wider distally. Pereopods 3–5 dactyli with rounded basal protuberance. Maximum cl. 2.5 mm. Intertidal, subtidal (0–15 m; associated with corals). Eastern Indo-Pacific, Tropical Eastern Pacific. 2 species (Marin 2011: key to species).

Hamiger Borradaile, 1916

145

1–3 pleura rounded, 4, 5 with posteroventral sharp tooth. Maximum cl. 8 mm. Intertidal–shelf (0–54 m; associated with scleractinian corals). Indo-West Pacific, Tropical Eastern Pacific. 3 species (Bruce 1976d; Chace and Bruce 1993: key to species).

Harpilius Dana, 1852 Diagnosis. Hepatic spine present. Thoracic sternite 4 with slender median spine. Pereopods 3–5 propodi distally setose, without spiniform setae; dactyli short, stout, hook-like. Maximum cl. 4.2 mm. Subtidal (2–6 m; associated with scleractinian corals). Western and Central Indo-Pacific. 4 species (Bruce 2004c: rediagnosis, key to 3 species; Bruce 2006: 4 species compared).

Holthuisaeus Anker & De Grave, 2010

Diagnosis. Rostrum blade-like, with 6–9 dorsal teeth, with 2 ventral teeth, with 2 or 3 middorsal teeth behind orbit. Pereopods 2 unequal, major fixed finger cutting edge with bilobed lobe. Maximum cl. 8.6 mm. Shelf (128 m; in hexactinellid sponges). Temperate Australasia (N of New Zealand). 1 species (Bruce 1986d).

Diagnosis. Rostrum with dorsal teeth only. Mandibular incisor vestigial. Pereopods 2 unequal, dissimilar, major dactylus with molar-shaped tooth fitting into socket of fixed finger. Pereopods 3–5 dactyli biunguiculate. Telson with 3 pairs of dorsal spiniform setae. Maximum cl. 10.7 mm. Intertidal–shelf (0–88 m; associated with sponges). Tropical W Atlantic. 1 species (Anker and De Grave 2010).

Hamodactyloides Fujino, 1973

Ischnopontonia Bruce, 1966

Diagnosis. Rostrum with 4 dorsal teeth, ventrally unarmed. Carapace hepatic spine present. Maxillipeds 2, 3 without exopod. Pereopod 1 fingers much less than half length of palm. Pereopods 3–5 dactyli with rounded basal protuberance. Maximum cl. 1.5 mm. Subtidal (1–24 m; associated with hydroids). Western and Central Indo-Pacific. 2 species (Bruce 1976c; Fujino 1973a).

Diagnosis. Body laterally compressed. Rostrum dentate with 3 or more middorsal teeth behind orbit, extending to mid-carapace. Uropodal exopod spiniform seta almost reaching end of exopod, curved. Telson with all spiniform setae on distal third of posterior margin. Maximum cl. 2.5 mm. Subtidal (0–5 m; associated with scleractinian corals). Western and Central Indo-Pacific. 1 species (Bruce 1966b).

Hamodactylus Holthuis, 1952 Diagnosis. Rostrum dorsal margin elevated, dentate, ventral margin obsolete. Carapace hepatic spine present. Maxillipeds 2, 3 without exopod. Pereopod 1 fingers at least half length of palm. Pereopod 2 dactylus much longer than fixed finger, hook-like. Maximum cl. 2 mm. Subtidal–shelf (2–40 m; associated with gorgonians, alcyonaceans and scleractinian corals). Western and Central IndoPacific. 6 species (Horká et al. 2016: key to species).

Isopontonia Bruce, 1982 Diagnosis. Rostrum cylindrical, acute. Pereopods 2 unequal; pereopod 2 major chela twice as long as carapace. Maximum cl. 1.5 mm. Subtidal (1–15 m). Central Indo-Pacific (Chesterfield Is). 1 species (Bruce 1982b).

Izucaris Okuno, 1999

Diagnosis. Rostrum with dorsal teeth only. Telson without spiniform setae, ending in 2 large downcurved teeth. Subtidal (0–10 m; associated with corals). Central Indo-Pacific. 5 species (Marin 2014a: key to species).

Diagnosis. Rostrum reaching beyond eyes, basal part widely expanded, forming roof of orbit, narrowing to compressed distal part. Carapace antennal spine prominent, at end of sharp ridge below deep orbital depression; pterygostomial lobe small. Maxillipeds 1–3 without exopods. Maximum cl. 3.4 mm. Shelf (20–110 m; associated with actinarians). Temperate Northern W Pacific, Eastern Indo-Pacific. 2 species (Okuno 1999).

Harpiliopsis Borradaile, 1917

Jocaste Holthuis, 1952

Diagnosis. Body depressed. Rostrum laterally compressed, with dorsal and ventral teeth. Carapace hepatic spine present. Pleonites

Diagnosis. Body strongly depressed. Rostrum with broad base forming supraorbital eaves, sharply tapering to compressed apex,

Hamopontonia Bruce, 1970

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Marine Decapod Crustacea

with dorsal and sometimes ventral spines. Carapace hepatic spine present. Pereopods 3–5 dactyli with hoof-shaped basal protuberance. Maximum cl. 3.2 mm (Pl. 10m). Subtidal–shelf (1–55 m; associated with corals). Western and Central Indo-Pacific. 3 species (Fransen 1994: species compared).

Kaviengella Šobáňová & Ďuriš, 2018 Diagnosis. Body vermiform. Rostrum very short, triangular, with 1 dorsal tooth. Scaphocerite bifid. Pereopod 1 fixed finger tridentate, with smaller distal teeth; dactylus spatulate, with denticulate margin. Pereopods 3–5 propodi with spiniform setae on flexor margin. Maximum cl. 1.45 mm. Subtidal (20 m; associated with sponges). Central Indo-Pacific (Papua New Guinea). 1 species (Šobáňová and Ďuriš 2018).

Laomenes Clark, 1919 Diagnosis. Rostrum with lateral carinae expanded proximally forming supraorbital eaves. Carapace hepatic spine present. Epistome with pair of acute processes. Cornea conically produced. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 4 mm (Pl. 10n). Subtidal, shelf (1.5–95 m; associated with crinoids). Temperate Northern Pacific, Central Indo-Pacific. 12 species (Marin 2009a: key to 8 species; Marin et al. 2012).

Leptomenaeus Bruce, 2007 Diagnosis. Body slender. Rostrum with 7–9 dorsal teeth, plus 1 distant tooth on carapace midline, without or with 1 small ventral tooth. Carapace hepatic spine present. Thoracic sternite 3 ~4 times as long as thoracic sternite 4. Cornea wider than eyestalk. Antennular peduncle article 2 elongate, flat. Pereopod 2 fingers 1.5 times as long as palm, with recurved teeth on cutting edges. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 2.3 mm. Subtidal (1–2 m). Central Indo-Pacific. 2 species (Ďuriš and Horká 2008).

Lipkebe Chace, 1969 Diagnosis. Rostrum prominent, unarmed. Carapace with orbit with supraorbital tooth beside rostrum; hepatic spine present. Pleonites 4–6 pleura with posteroventral sharp tooth. Maxillipeds 2, 3 without exopod. Maximum cl. 4.6 mm. Shelf (64–150 m; associated with crinoids). Tropical W Atlantic. 1 species (Bruce 1976a; Shaw et al. 1977).

Lipkemenes Bruce & Okuno, 2010 Diagnosis. Body depressed. Rostrum long, with 9 dorsal teeth, 1 distoventral tooth. Carapace hepatic spine present. Pleonites 1–3 pleura rounded, 4, 5 with posteroventral sharp tooth. Pereopods 3–5 propodi with long spiniform setae along flexor margin; dactyli biunguiculate. Maximum cl. 4.3 mm. Subtidal, shelf (15–120 m; associated with gorgonocephalid ophiuroids). Western and Central Indo-Pacific. 1 species (Bruce and Okuno 2010).

Madangella Frolová & Ďuriš, 2018 Diagnosis. Carapace hepatic spine present. Thoracic sternite 4 with slender median spine. Pereopods 2 feeble, symmetrical, barely exceeding pereopods 1; fingers simple, longer than palm. Maximum cl. 1.6 mm. Subtidal (4–8 m). Central Indo-Pacific (Papua New Guinea). 1 species (Frolová and Ďuriš 2018).

Manipontonia Bruce, Okuno & Li, 2005 Diagnosis. Rostrum twice as long as carapace, slender, curved, with 2–4 proximal dorsal teeth, without ventral teeth, epigastric tooth and 2 posterior rostral teeth large, similar, curved anteriorly, without sutures. Carapace hepatic spine present. Maximum cl. 4.1 mm. Intertidal–shelf (0–105 m). Western and Central Indo-Pacific. 3 species (Bruce 2012).

Margitonia Bruce, 2007 Diagnosis. Rostrum prominent, unarmed. Orbit deep, with posterior notch; hepatic spine present. Pereopods 3–5 propodi with distal tufts of setae; dactyli biunguiculate or elongate, corpus with crenulate distoventral accessory tooth, with transverse row of simple setae distolaterally. Maximum cl. 1.7 mm. Intertidal (associated with echinoids). Eastern Indo-Pacific (Hawaii). 1 species (Bruce 1974: description of only species as Periclimenes insolitus; Bruce 2007a).

Mesopontonia Bruce, 1967 Diagnosis. Rostrum blade-like, with dorsal and sometimes few ventral teeth, with epigastric tooth separate from rostrum. Carapace antennal spine absent; hepatic spine present. Maxilliped 3 without exopod. Pereopod 1 fingers at least half length of palm. Pereopods 2 dissimilar, cutting edges of major fingers with teeth; pereopod 2 dactylus as long fixed finger, straight. Maximum cl. 3.6 mm. Shelf–slope (35–460 m). Western and Central Indo-Pacific. 6 species (Bruce 1996: key to species; Li and Bruce 2006: refined key).

Metapontonia Bruce, 1967 Diagnosis. Rostrum short, unarmed, with large median blunt tooth behind. Pterygostomial margin roundly produced forward beyond rostrum. Maximum cl. 2.6 mm. Subtidal (2–19 m; associated with corals). Western and Central Indo-Pacific. 2 species (Bruce 1967; Ďuriš and Lin 2016).

Michaelimenes Okuno, 2017 Diagnosis. Rostrum horizontal, ventral margin straight, dentate, dorsa margin dentate. Carapace with orbit weakly developed; hepatic spine present. Pereopod 2 major chela fixed finger with proximal excavation on cutting surface receiving proximal part of

6 – Caridea – shrimps

dactylar cutting edge, overlapping; pereopod 2 short, with flange on upper margin, cutting edge proximally dentate. Pereopods 3–5 propodi with spiniform setae on flexor margin; dactyli biunguiculate. Maximum cl. 2.7 mm. Subtidal, shelf (20–304 m; associated with soft corals). Temperate Northern W Pacific, Indo-West Pacific. 3 species (Okuno 2017: key to species).

Miopontonia Bruce, 1985 Diagnosis. Rostrum prominent, unarmed. Carapace hepatic spine present; pterygostomial sinus opening to branchial cavity present. Maxillipeds 2, 3 without exopod. Pereopod 1 fingers at least half length of palm. Pereopod 2 carpus, merus and ischium with 1–3 teeth on flexor margin; dactylus much longer than fixed finger, hook-like. Maximum cl. 7 mm. Shelf (40–80 m; associated with gorgonians and antipatharians). Temperate Northern W Pacific, Central Indo-Pacific. 1 species (Bruce 1985b).

Neoanchistus Bruce, 1975 Diagnosis. Rostrum flat, depressed, truncate, without teeth. Carapace antennal spine absent. Telson with both pairs of dorsal spiniform setae in posterior half. Maximum cl. 10 mm. Subtidal (10 m; associated with bivalves). Western and Central Indo-Pacific. 2 species (Bruce 1975; Holthuis 1986).

147

Subtidal (7–15 m; associates of sponges). Central Indo-Pacific. 3 species (Bruce 2014a; Fransen 2013).

Notopontonia Bruce, 1991 Diagnosis. Body compressed. Rostrum triangular, without teeth. Pereopod 2 palm wide, compressed, flexor margin convex, carinate; fixed finger 3 times as wide as dactylus. Maximum cl. 14 mm. Subtidal–shelf (7–64 m; inside ascidians). Temperate Australasia. 1 species (Berggren 1999).

Odontonia Fransen, 2002 Diagnosis. Rostrum dorsally unarmed; dorsal carina bordered by lateral depressions, lateral carinae well developed. Carapace pterygostomial angle square. Antennular peduncle medioventral tooth strongly developed. Pereopods 3–5 dactyli often with denticles on flexor margin or unguis with patch of distal scales. Maximum cl. 6 mm. Subtidal, shelf (2–71 m; associated with ascidians). Western and Central Indo-Pacific. 10 species (Fransen 2002: key to 6 species).

Onycocaridella Bruce, 1981

Diagnosis. Rostrum slender, dorsal margin convex, with 8 close teeth, ventral margin with 3 teeth. Carapace hepatic spine movable. Pereopod 2 carpus longer than chela. Maximum cl. 4.7 mm. Slope (500 m). Central Indo-Pacific. 1 species (Mitsuhashi et al. 2010).

Diagnosis. Rostrum reduced, compressed, unarmed or with dorsal apical tooth. Pereopod 1 fingers as long as palm, spatulate. Pereopod 2 palm subcylindrical, fixed finger short, unarmed. Telson with anterior pair of dorsal spiniform setae in anterior half. Maximum cl. 2 mm. Intertidal, subtidal (0–15 m; associated with sponges). Western and Central Indo-Pacific. 7 species (Bruce 1981; Šobáňová and Ďuriš 2021: rediagnosis, key to species).

Neopericlimenes Heard, Spotte & Bubucis, 1993

Onycocaridites Bruce, 1987

Diagnosis. Rostrum well developed, deep, with dorsal teeth, lacking lateral eave. Carapace branchiostegal spine present. Maxillipeds 1–3 with rudimentary exopods. Maximum cl. 3.2 mm. Shelf (40 m; associated with antipatharians). Tropical W Atlantic. 1 species (Heard et al. 1993).

Diagnosis. Body rather stout. Rostrum short, tapering, with 3 small dorsal teeth. Pereopod 3 dactylus more than 6 times as long as wide, with long unguis about as long as propodus. Maximum cl. 3 mm. Shelf (60 m; associated with sponges). Central Indo-Pacific. 1 species (Bruce 1987).

Neoclimenes Mitsuhashi, Li & Chan, 2010

Neopontonides Holthuis, 1951 Diagnosis. Rostrum reaching beyond eyes, gradually narrowing from base to compressed distal part, with dorsal teeth. Maxillipeds 2, 3 without exopod. Maximum cl. 3 mm. Intertidal–shelf (0–32 m; associated with gorgonians). Tropical W Atlantic, Tropical Eastern Pacific. 5 species (Fransen and de Almeida 2009; Heard 1986: key to 3 species).

Nippontonia Bruce & Bauer, 1997 Diagnosis. Body vermiform. Rostrum absent. Scaphocerite tapering, acute. Uropodal exopod anterior margin distally dentate. Maximum cl. 4.8 mm.

Onycocaris Nobili, 1904 Diagnosis. Rostrum from compressed, dorsally dentate to reduced, unarmed. Pereopod 1 fingers less than half length of palm, nonspatulate. Pereopod 2 palm laterally compressed, fixed finger long, usually with lateral flange. Telson with anterior pair of dorsal spiniform setae in anterior half. Maximum cl. 3.5 mm. Subtidal, shelf (0–84 m; associated with sponges). Temperate Northern W Pacific, Indo-West Pacific. 22 species. Onycocaris has been divided into species-groups, for some of which keys to species exist (Bruce 2011b; Marin 2015). Another key deals with 16 species (Bruce 2011c).

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Marine Decapod Crustacea

Onycomenes Bruce, 2009 Diagnosis. Rostrum short, depressed, with small preterminal dorsal tooth. Scaphocerite rudimentary. Maxillipedal exopodal flagella strap-like, with numerous plumose distal and marginal setae. Pereopods 3–5 dactyli simple (fifth only) or biunguiculate. Maximum cl. 2.7 mm. Subtidal (10–15 m; inside sponges). Central Indo-Pacific (Vietnam). 1 species (Marin 2007b: redescription of type species as Onycocaridella antokha Marin, 2007).

Opaepupu Anker & De Grave, 2021 Diagnosis. Body strongly depressed. Rostrum broadly dorsally triangular, carinate. Carapace antennal spine prominent. Pereopods 2 unequal. Scaphocerite unarmed. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 4.3 mm. Subtidal (14 m; inside mantle cavity of bivalve, Trapezium oblongum). Eastern Indo-Pacific (Hawaii). 1 species (Anker and De Grave 2021).

absent; hepatic spine present. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 5.3 mm. Shelf (109–520 m; associated with gorgonians). Central IndoPacific. 3 species (Bruce 1995).

Paranchistus Holthuis, 1952 Diagnosis. Rostrum with few, small, dorsal and ventral teeth distally. Hepatic spine movable. Pereopod 2 carpus about third length of chela. Maximum cl. 4.2 mm. Intertidal, subtidal (0–10 m; associated with bivalves). Temperate Northern W Pacific, Central Indo-Pacific. 6 species (Li et al. 2004; Suzuki 1971: key to 4 species).

Paratypton Balss, 1914 Diagnosis. Rostrum absent. Scaphocerite rudimentary. Maximum cl. 4.1 mm. Subtidal (1–14 m; forming cysts in acroporid corals). Western and Central Indo-Pacific. 1 species (Bruce 1969b).

Orthopontonia Bruce, 1982

Patonia Mitsuhashi & Chan, 2006

Diagnosis. Rostrum with dorsal teeth only. Pleonite 6 posterolateral angle ending in slender spine. Pereopod 2 minor fingers with numerous flat ridged tubercles on cutting faces. Maximum cl. 4.6 mm. Subtidal–shelf (0.5–46 m; associated with sponges). Western and Central Indo-Pacific. 1 species (Bruce 1982a).

Diagnosis. Body laterally compressed. Rostrum deep, dorsal margin convex, with 10 close teeth, without ventral teeth. Carapace hepatic spine movable. Pereopod 2 carpus longer than chela. Maximum cl. 4 mm. Slope (209–280 m). Central Indo-Pacific. 1 species (Mitsuhashi and Chan 2006).

Palaemonella Dana, 1852 Diagnosis. Carapace hepatic spine present. Thoracic sternite 4 with slender median spine. Mandibular palp of 1 or 2 articles. Pereopod 1 fingers stout, deeply subspatulate, with mesial subspatulate concavity, without distal lamina. Maximum cl. 3.7 mm. Intertidal–slope (0–544 m; most free-living, some associated with fishes or invertebrates). Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Tropical Eastern Pacific. 23 species (Komai and Yamada 2015: key to 20 Indo-West Pacific species).

Paraclimenaeus Bruce, 1988 Diagnosis. Rostrum dorsally dentate, ventrally unarmed, lateral carinae moderately expanded. Pereopods 2 unequal; pereopod 2 major chela fixed finger with 1 or 2 teeth, dactylus with matching sockets. Pereopods 3–5 dactyli biunguiculate. Uropodal exopod with numerous spiniform setae on suture and anterior margin. Maximum cl. 3.2 mm. Subtidal, shelf (3–90 m; associated with sponges). Western and Central Indo-Pacific. 5 species (Bruce 2009a; Odijk and Fransen 2017: key to species).

Periclimenaeus Borradaile, 1915 Diagnosis. Rostrum with dorsal teeth only. Pereopods 2 unequal, dissimilar, major dactylus with molar-shaped tooth fitting into socket of fixed finger. Maximum cl. 4 mm (Pl. 10o). Subtidal–slope (0–450 m depth). Cosmopolitan in tropical and subtropical seas (on coral reefs, associated with sponges, gorgonians and ascidians). 87 species. Periclimenaeus is the second largest pontoniine genus despite smaller genera being hived off. The only keys to species deal with limited groups defined on the shape of the dactylus of pereopod 2 (Fransen 2006), or the dactylus of the ambulatory pereopods (Bruce 2014b). The dactylus of pereopod 3 can be used to differentiate Indo-West Pacific species (Bruce 2013).

Periclimenella Bruce, 1995 Diagnosis. Carapace hepatic spine present. Thoracic sternite 4 with slender median spine. Pereopod 1 fingers stout, deeply subspatulate, with mesial subspatulate concavity, without distal lamina. Pereopod 2 major chela fingers with opposing fossae. Maximum cl. 5.8 mm. Intertidal–shelf (0–34 m). Indo-West Pacific. 2 species (Ďuriš and Bruce 1995: key to species).

Paraclimenes Bruce, 1995

Periclimenes Costa, 1844

Diagnosis. Rostrum compressed, with numerous acute dorsal teeth, 3–4 prominent ventral teeth. Carapace antennal spine

Diagnosis. Carapace hepatic spine present. Pereopods 3–5 dactyli simple (rarely) or biunguiculate (Pl. 11g).

6 – Caridea – shrimps

Subtidal–slope (0–1290 m). Cosmopolitan in tropical and subtropical seas. 119 species. Periclimenes remains the largest pontoniine genus although many genera have recently been created or resurrected for single species or small groups of species (see review by Li 2009 of Periclimenes and other genera created later). Partly for this reason the genus is difficult to diagnose – it is diagnosed more by absence of the features of the smaller genera than by any synapomorphy. Several species-groups persist within the remaining species  – keys to species exist for some (e.g. Li et  al. 2008; Martínez-Mayén et al. 2006). Mediterranean species can be resolved using the key of Grippa and d’Udekem d’Acoz (1996) and those of the Indo-West Pacific using Chace and Bruce (1993) but some of these species may now be in other genera. Holthuis’s (1993) figure of the type species, Periclimenes amethysteus (Risso, 1827), is reproduced here (Fig. 6.31b).

Periclimenoides Bruce, 1990 Diagnosis. Rostrum blade-like, dorsally serrate, with or without 1 ventral tooth. Mandibular incisor bifid or trifid. Telson with 2 pairs of lateral setae in anterior half. Maximum cl. 3.2 mm. Subtidal (5–20 m; associated with sponges). Central IndoPacific. 3 species (Šobáňová et al. 2017: key to species).

Philarius Holthuis, 1952 Diagnosis. Rostrum deep, with 4–10 dorsal teeth, 1 or 2 ventral teeth. Pereopods 3–5 dactyli simple, hook-shaped, hidden by tuft of setae. Telson with both pairs of dorsal spiniform setae in posterior half. Maximum cl. 3.2 mm. Intertidal–shelf (0–28 m; associates of branching Acropora). Indo-West Pacific. 10 species (Kou and Li 2016: tabulated differences between 8 species; Marin and Anker 2011: key to 7 species).

Phycomenes Bruce, 2008 Diagnosis. Carapace hepatic spine present. Thoracic sternite 4 with short triangular anterior lobe. Pereopods 1 and 2 similar. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 2.4 mm. Intertidal, subtidal (0–4 m; from seagrass). Tropical Atlantic, Western and Central Indo-Pacific, Temperate Australasia. 5 species (Bruce 2008b; 2010a: key to species).

Pinnotherotonia Marin & Paulay, 2010 Diagnosis. Body depressed, completely covered by dense pubescence. Rostrum broad, truncate. Cornea reduced, without accessory pigment spot. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 6.2 mm. Intertidal, subtidal (0–10 m; in mantle cavity of bivalve, Periglypta crispata). Central Indo-Pacific (Palau). 1 species (Marin and Paulay 2010).

Platycaris Holthuis, 1952 Diagnosis. Body strongly depressed, with deep postorbital depression formed by rostrum lateral margin and oblique ridge from

149

lower orbital angle. Rostrum broadly triangular, with wide supraocular eaves. Pereopods 2 unequal. Maximum cl. 5 mm. Intertidal, subtidal (0–26 m; associated with corals, Galaxea). Indo-West Pacific. 1 species (Bruce 1966a).

Platypontonia Bruce, 1968 Diagnosis. Body strongly depressed. Rostrum triangular, not reaching beyond eyes. Pereopods 2 subequal. Maximum cl. 5.3 mm. Subtidal (2–22 m; associated with bivalves). Western IndoPacific. 2 species (Hipeau-Jacquotte 1971: key to species).

Plesiomenaeus Bruce, 2009 Diagnosis. Rostrum shorter than eyestalk, compressed, with 1 tooth. Pereopods 2 unequal, similar, dactylus entire, fitting into longitudinal depression on finger. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 6 mm. Subtidal (0.1 m). Western Indo-Pacific. 1 species (Bruce 2009b).

Plesiopontonia Bruce, 1985 Diagnosis. Rostrum laterally compressed, with dorsal and ventral teeth. Telson with 4 pairs of dorsal spiniform setae. Maximum cl. 5.3 mm. Slope (242–320 m). Central Indo-Pacific (Philippines). 1 species (Bruce 1985a; Li et al. 2008).

Pliopontonia Bruce, 1973 Diagnosis. Rostrum high, not reaching beyond antennular peduncle article 1, with 4 dorsal, without ventral teeth, with 1–3 middorsal teeth behind orbit. Pereopod 2 fingers half as long as palm. Pereopods 3–5 dactyli short, stout, hook-like. Maximum cl. 6.1 mm (Pl. 11a). Subtidal–shelf (1–40 m; associated with corals). Western and Central Indo-Pacific. 2 species (Okuno 2009: compared 2 species).

Pontonia Latreille, 1829 Diagnosis. Rostrum with at most 1 subdistal dorsal tooth, dorsal carina not developed. Thoracic sternite 4 anteriorly produced as triangular plate. Pleonite 6 posterolateral angle ending in slender spine. Antennular peduncle medioventral tooth small. Pereopods 3–5 dactyli biunguiculate. Telson with 2 pairs of lateral setae equally spaced. Maximum cl. 13 mm (Pl. 11b, c). Intertidal–shelf (1–290 m; associated with molluscs). Temperate Northern and Tropical Atlantic, Tropical Eastern Pacific. 11 species (Fransen 2002: key to 10 species).

Pontonides Borradaile, 1917 Diagnosis. Body depressed. Rostrum basally wide, not reaching beyond eyes, fully depressed, without teeth. Carapace with orbit complete or incomplete, open posteriorly. Maxillipeds 1–3 with rudimentary exopods. Pereopods 2 (minor) sexually dimorphic,

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Marine Decapod Crustacea

fingers of male thin, as long as palm. Uropodal exopod anterodistal tooth movable. Maximum cl. 4.8 mm. Subtidal, shelf (5–112 m; associated with black and dendrophylliid corals). Indo-West Pacific. 7 species (Marin 2007a: key to 8 species; 2008c: rediagnosis).

Pleonites 3–5 pleura produced as narrow blunt projection or with posteroventral sharp tooth. Maxillipeds 2, 3 without exopod. Maximum cl. 2.5 mm. Subtidal–shelf (3–138 m; on antipatharians). Tropical Atlantic, Tropical Eastern Pacific. 8 species (Olthof et al. 2018).

Pontoniopsides Bruce, 2005

Pseudopontonia Bruce, 1992

Diagnosis. Rostrum acute, without orbital shelf, without teeth. Pereopods 2 dissimilar, unequal, major chela massive. Telson with both pairs of dorsal spiniform setae in posterior half (small). Maximum cl. 3.2 mm. Shelf (63 m; associated with echinoids). Tropical Atlantic (Florida, USA). 1 species (Bruce 2005a).

Diagnosis. Body compressed. Rostrum triangular, without teeth, distally compressed. Telson with 6 pairs of posterior spiniform setae. Maximum cl. 4.6 mm. Subtidal (inside ascidians). Western Indo-Pacific, Temperate Australasia. 1 species (Bruce 1972b, 1992).

Pontoniopsis Borradaile, 1915 Diagnosis. Rostrum triangular, centrally depressed with orbital shelf, without teeth. Pereopod 1 fingers spatulate. Pereopods 2 very unequal, major carpus cup-shaped, minor carpus slender; pereopod 2 major dactylus with flange on upper half of lateral surface. Pereopods 3–5 dactyli simple (with small accessory tooth). Telson with both pairs of dorsal spiniform setae in posterior half (small). Maximum cl. 1.05 mm. Intertidal, subtidal (0–22 m; associated with crinoids). IndoWest Pacific. 1 species (Bruce 1980).

Poripontonia Fransen, 2003 Diagnosis. Rostrum with 3 unevenly spaced dorsal teeth, 1 distal ventral tooth. Carapace with sharp antennal spine. Maximum cl. 3.4 mm. Subtidal (to 25 m; associated with sponges). Central IndoPacific (Bali, Indonesia). 2 species (Fransen 2003).

Pseudopontonides Heard, 1986 Diagnosis. Rostrum reaching beyond eyes, basal part widely expanded, forming roof of orbit, narrowing to compressed distal part. Antennal spine small; pterygostomial sinus opening to branchial cavity present. Maxillipeds 2, 3 without exopod. Maximum cl. 3.2 mm. Subtidal, shelf (1–65 m; associated with antipatharians). Tropical W Atlantic. 2 species (Heard 1986; Snijders and Fransen 2010).

Pseudoveleronia Marin, 2008 Diagnosis. Body depressed. Rostrum basally wide, forming roof over orbit. Carapace with orbit complete, closed. Maxillipeds 2, 3 without exopod. Pereopod 1 fingers spatulate. Maximum cl. 2 mm. Subtidal (2–20 m; associated with gorgonians). Tropical Eastern Pacific. 1 species (Marin 2008c).

Rapimenes Ďuriš & Horká, 2017

Diagnosis. Rostrum dorsal margin convex, with 6 teeth, ventral margin with 1 subdistal tooth. Carapace hepatic spine present. Maxilliped 3 without exopod. Pereopod 1 fingers at least half length of palm. Pereopod 2 dactylus as long fixed finger, straight. Maximum cl. 2 mm. Intertidal, subtidal (0–3 m; associated with alcyonarians). Western and Central Indo-Pacific. 1 species (Bruce 1969a).

Diagnosis. Rostrum subequal to antennular peduncle, with 5–7 low dorsal and 0–3 ventral teeth, lateral carinae with depressed supraorbital tooth, epigastric tooth smaller than posterior rostral teeth, with basal suture. Carapace hepatic spine present. Pereopods 3–5 propodi prehensile, with 2–5 pairs of long straight spiniform setae along flexor margin. Maximum cl. 2.5 mm. Subtidal, shelf (14–52 m; associated with antipatharians, hydroids, pennatularians and scyphozoans). Western and Central Indo-Pacific. 3 species (Ďuriš 2010: descriptions of species as Periclimenes granulimanus species-group; Ďuriš and Horká 2017: key to species).

Pseudoclimenes Bruce, 2008

Rapipontonia Marin, 2007

Diagnosis. Rostrum blade-like, with dorsal and ventral teeth, with 2 or 3 middorsal teeth behind orbit. Maximum cl. 1.85 mm. Shelf (114 m). Central Indo-Pacific (NW Australia). 1 species (Bruce 2008a).

Diagnosis. Rostrum with 6 dorsal teeth, prominent basally, without ventral teeth, epigastric tooth smaller than posterior rostral teeth, with basal suture. Carapace hepatic spine present. Pereopod 1 fingers expanded, forming scoops, extremity bearing 3 small recurved teeth. Pereopods 3–5 propodi prehensile, with long distal paired and unpaired spiniform setae along flexor margin. Maximum cl. 3.4 mm. Subtidal (1–26 m; associated with hydroids). Tropical Atlantic, Indo-West Pacific. 4 species (Marin 2009b: key to species).

Propontonia Bruce, 1969

Pseudocoutierea Holthuis, 1951 Diagnosis. Rostrum slender, reaching beyond scaphocerite, basally extended as supraorbital eave. Antennal spine apparently bifid.

6 – Caridea – shrimps

Rostronia Fransen, 2002 Diagnosis. Rostrum with 2 or 3 dorsal distal teeth, lateral carinae not developed. Antennular peduncle medioventral tooth small. Pereopods 3–5 dactyli biunguiculate, with denticles on flexor margin. Maximum cl. 7 mm. Subtidal, shelf (1–42 m; associated with ascidians). Western and Central Indo-Pacific. 1 species (Fransen 2002).

Sandimenes Li, 2009 Diagnosis. Body covered by dense pubescence on ventral surface, and on appendages. Rostrum slender, straight, dentate dorsally, compressed, lateral carinae distinct. Carapace hepatic spine present. Pereopods 3–5 propodi with bunches of long setae along flexor margin. Maximum cl. 5.2 mm. Intertidal–shelf (0–119 m; associated with echinoids). Western and Central Indo-Pacific. 1 species (Kou et al. 2016b: erroneously included 2 species; Li 2009).

Sandyella Marin, 2009 Diagnosis. Rostrum absent. Carapace with 1 dorsal conical, median horn, 2 smaller posterolateral horns. Pleonite 2 with pair of horns, pleonite 3 with median horn. Pleonites 4–6 pleura with posteroventral sharp tooth. Maxillipeds 2, 3 without exopod. Maximum cl. 2.6 mm. Shelf (30–55 m; associated with antipatharians and gorgonians). Tropical Eastern Pacific. 5 species (Li and Poupin 2009: key to species as Chacella; Marin 2009c).

Stegopontonia Nobili, 1906 Diagnosis. Rostrum large, T- or +-shaped in transverse section, without dorsal or ventral teeth. Carapace hepatic spine present. Intertidal, subtidal (0–1 m; associated with echinoids). IndoWest Pacific. 1 species (Bruce 1982c).

Tectopontonia Bruce, 1973 Diagnosis. Rostrum with 6 dorsal teeth. Carapace postorbital spine present. Maxilliped 3 without exopod. Pereopod 2 globular. Maximum cl. 1.85 mm. Subtidal (4 m; on scleractinian corals). Western Indo-Pacific. 1 species (Bruce 1973b).

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Carapace postorbital groove distinct, bordered by postorbital carina and carina from antennal to hepatic spine. Maximum cl. 1.7 mm. Intertidal, subtidal (0–15 m; associated with echinoids). Tropical Atlantic, Western and Central Indo-Pacific. 4 species (Berggren 1994: key to species).

Typton Costa, 1844 Diagnosis. Rostrum short. Scaphocerite rudimentary. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 3.5 mm (Pl. 11e). Intertidal, subtidal (0–73 m; some associated with echinoderms, sponges). Temperate Northern and Tropical Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific. 21 species (Anker et  al. 2021; Ayón-Parente et  al. 2015: key to E Pacific species; Bruce 2000b: key to 6 Indo-West Pacific species).

Typtonoides Bruce, 2010 Diagnosis. Rostrum long, with 9 dorsal teeth, 1 distoventral tooth. Scaphocerite rudimentary. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 1.25 mm. Subtidal (15 m). Central Indo-Pacific (Chesterfield Is). 1 species (Bruce 2010d).

Typtonomenaeus Marin & Chan, 2013 Diagnosis. Rostrum with dorsal teeth only. Scaphocerite shorter than antennal peduncle. Pereopods 2 unequal, similar, both major and minor dactyli with molar-shaped tooth fitting into socket of fixed finger. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 3.2 mm. Subtidal (5–6 m; associated with sponges). Central IndoPacific (Taiwan). 1 species (Marin and Chan 2013).

Typtonychus Bruce, 1996 Diagnosis. Rostrum short. Scaphocerite rudimentary. Basal endites of maxilla and maxilliped 1 elongate, setose, forming basket. Pereopod 2 fingers with 2 sharp apical teeth. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 4.2 mm. Subtidal (3–38 m; inside sponges). Western and Central IndoPacific. 6 species (Bruce 2011a: key to species).

Unesconia Anker, 2020

Diagnosis. Rostrum laterally compressed, with dorsal and ventral teeth. Pereopods 1 unequal; carpus segmented. Maximum cl. 8 mm (Pl. 11d). Subtidal–shelf (6–141 m; associated with sponges). Western and Central Indo-Pacific. 1 species (Bruce 1979b).

Diagnosis. Rostrum dorsally carinate, dentate, ventrally unarmed, lateral carina proximally expanded. Pereopods 2 unequal; pereopod 2 major chela swollen, with 2 opposing deep fossae, one on fixed finger, one on dactylus. Pereopods 3–5 dactyli biunguiculate. Uropodal exopod suture with 5 spiniform setae laterally, middle one longest. Maximum cl. 3.1 mm. Subtidal (1–2 m; associated with sponges). Tropical Eastern Pacific. 1 species (Anker 2020).

Tuleariocaris Hipeau-Jacquotte, 1965

Unguicaris Marin & Chan, 2006

Diagnosis. Rostrum basal part with narrow lateral wings, narrowing to compressed distal part, with small dorsal and ventral teeth.

Diagnosis. Rostrum with dorsal teeth only. Carapace hepatic spine present. Mandibular incisor wider distally. Pereopods 3–5 dactyli

Thaumastocaris Kemp, 1922

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Marine Decapod Crustacea

corpus upper margin with distal accessory spinules. Maximum cl. 5.1 mm. Subtidal–shelf (6–37 m; associated with crinoids). Central Indo-Pacific. 4 species (Marin and Chan 2006: key to species).

Urocaris Stimpson, 1860 Diagnosis. Rostrum compressed, dorsal carina with numerous acute teeth, lateral carinae weak, ventral carina obsolete, weakly dentate distally. Carapace antennal spine absent; hepatic spine present. Epistome with acute median process. Pereopods 3–5 dactyli biunguiculate. Maximum cl. 2.5 mm. Subtidal, shelf (1–27 m). Temperate Northern and Tropical W Atlantic. 1 species (Bruce 2007b).

Subtidal to 9 m. Western Indo-Pacific (Yemen). 1 species (Bruce 1997).

Zenopontonia Bruce, 1975 Diagnosis. Rostrum with dorsal teeth only. Carapace hepatic spine movable. Thoracic sternite 4 with broad transverse ridge, with median incision. Pereopod 1 fingers spatulate. Pereopod 2 carpus about third length of chela. Pereopods 2–5 meri with distal tooth on flexor margin; 3–5 propodi with spiniform setae on flexor margin; 3–5 dactyli obscurely biunguiculate. Maximum cl. 7 mm (Pl. 11f). Intertidal–shelf (0–25 m; on asteroids, nudibranchs, holothurians). Western and Central Indo-Pacific. 3 species (Kou et  al. 2016b: suggested polyphyly; Marin 2012: key to 3 species).

Veleronia Holthuis, 1951 Diagnosis. Body depressed. Rostrum basally wide, forming roof over orbit. Carapace with orbit complete or incomplete, open posteriorly. Maxillipeds 2, 3 without exopod. Maximum cl. 2 mm. Subtidal, shelf (10–23 m; associated with black corals, gorgonians). Eastern Indo-Pacific, Tropical Eastern Pacific. 2 species (Marin 2008c: key to species).

Veleroniopsis Gore, 1981 Diagnosis. Body depressed. Rostrum reaching beyond eyes, broad, basally wide and depressed, ending in laterally compressed distal part. Carapace postorbital margin inflated. with postorbital tubercle. Maxillipeds 2, 3 without exopod. Maximum cl. 3.2 mm. Subtidal (18 m). Tropical Atlantic (FL, USA). 1 species (Gore 1981).

Vir Holthuis, 1952 Diagnosis. Mandibular palp of 1 or 2 articles. Pereopod 2 carpus subequal to or shorter than palm. Pereopods 3–5 propodi unarmed. Maximum cl. 5.6 mm. Subtidal–shelf (3–30 m; most symbionts of caryophyllid corals). Central Indo-Pacific. 6 species (Marin 2008a: key to species; Marin 2007c).

Waldola Holthuis, 1951 Diagnosis. Rostrum dorsal margin convex, with 6 teeth, ventral margin straight. Carapace antennal spine absent; hepatic spine present. Maxillipeds 2, 3 without exopod. Pereopod 1 fingers at least half length of palm. Pereopods 2 dissimilar, cutting edges of major fingers with teeth; dactylus as long fixed finger, straight. Maximum cl. 7 mm. Subtidal, shelf (18–46 m). Tropical Eastern Pacific. 1 species (Holthuis 1951).

Yemenicaris Bruce, 1997 Diagnosis. Rostrum with dorsal teeth only. Telson far exceeding uropods, distally expanded, with 3 pairs of dorsal spiniform setae, without posterior spiniform setae. Maximum cl. 1.7 mm.

Zoukaris Anker & Corbari, 2020 Diagnosis. Rostrum with 7 dorsal teeth, 2 distoventral teeth. Carapace hepatic spine present. Pereopods 3–5 dactyli with small distal spines on flexor margin. Maximum cl. 11.6 mm. Shelf (208–385 m; possibly associated with echinoids). Tropical W Atlantic. 1 species (Anker and Corbari 2020). References Ahyong ST (2003) Gnathophyllum taylori, a new species of caridean shrimp from southeastern Australia (Crustacea: Decapoda: Gnathophyllidae). Memoirs of Museum Victoria 60, 237–242. doi:10.24199/j. mmv.2003.60.24 Ahyong ST (2015) Decapod crustacea of the Kermadec Biodiscovery Expedition 2011. Bulletin of the Auckland Museum 20, 405–442. Anger K (2013) Neotropical Macrobrachium (Caridea: Palaemonidae): on the biology, origin, and radiation of freshwater-invading shrimp. Journal of Crustacean Biology 33, 151–183. doi:10.1163/1937240X-00002124 Anker A (2020) Unesconia coibensis, gen. et sp. nov., a miniature spongeassociated shrimp from a biodiversity hotspot in the tropical eastern Pacific (Decapoda: Palaemonidae). Zootaxa 4731, 115–133. doi:10.11646/​ zootaxa.4731.1.8 Anker A, Corbari L (2020) A new deep-water palaemonid shrimp genus and species from the French Antilles, with a new record of Periclimenes milleri Bruce, 1986 (Decapoda: Caridea). Zootaxa 4834, 407–424. doi:10.11646/zootaxa.4834.3.4 Anker A, De Grave S (2010) Holthuisaeus, a new genus for Periclimenes (Periclimenaeus) bermudensis Armstrong, 1940 (Decapoda, Palaemonidae, Pontoniinae). In: Fransen C, De Grave S, Ng P (Eds) Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume. Crustaceana Monographs 14, 115–131. doi:10.1163/9789047427759_005 Anker A, De Grave S (2021) Opaepupu, a new genus and species of bivalveassociated shrimp (Decapoda: Caridea: Palaemonidae) from Hawai’i. Zootaxa 4903, 55–70. doi:10.11646/zootaxa.4903.1.3 Anker A, Pachelle PPG, Leray M (2021) Two new species of Typton Costa, 1844 from tropical American waters, with taxonomic notes on T. tortugae McClendon, 1911 and a new record of T. granulosus AyónParente, Hendrickx & Galvan-Villa, 2015 (Decapoda: Caridea: Palaemonidae). Zootaxa 4950, 267–295. doi:10.11646/zootaxa.4950.2.3 Ashelby CW, De Grave S (2010) A new genus of palaemonid shrimps (Crustacea: Decapoda: Palaemonidae) to accomodate Leander belindae Kemp, 1925, with a redescripion of the species. Zootaxa 2372, 369–378. doi:10.11646/zootaxa.2372.1.29

6 – Caridea – shrimps

Ashelby CW, Page TJ, De Grave S, Hughes JM, Johnson ML (2012) Regional scale speciation reveals multiple invasions of freshwater in Palaemoninae (Decapoda). Zoologica Scripta 41, 293–306. doi:10.1111/j.1463-6409.2012.00535.x Ashelby CW, Lin WC, Grave S, Chan T-Y (2018) Notes on the shrimp genus Palaemon Weber, 1795 (Crustacea, Decapoda, Palaemonidae) and related genera from Taiwan. Zootaxa 4446, 575–595. doi:10.11646/ zootaxa.4446.4.10 Ayón-Parente M, Hendrickx ME, Galvan-Villa CM (2015) A new species of the genus Typton Costa (Crustacea: Decapoda: Palaemonidae: Pontoniinae) from the eastern tropical Pacific. Zootaxa 3926, 430–438. doi:10.11646/zootaxa.3926.3.7 Baldari F, Mejía-Ortiz LM, López-Mejía M (2010) A new cave species of Cryphiops (Crustacea: Decapoda: Palaemonidae) from Southern Mexico. Zootaxa 2427, 47–54. doi:10.11646/zootaxa.2427.1.5 Banu R, Christianus A (2016) Giant freshwater prawn Macrobrachium rosenbergii farming: a review on its current status and prospective in Malaysia. Journal of Aquaculture Research & Development 7, 423. doi:10.4172/2155-9546.1000423 Berggren M (1990) Dasella herdmaniae (Lebour) (Decapoda: Natantia: Pontoniinae) from Moçambique and establishment of a new species, Dasella brucei. Journal of Crustacean Biology 10, 554–559. doi:10.2307/​1548347 Berggren M (1994) Periclimenes nomadophila and Tuleariocaris sarec, two new species of pontoniine shrimps (Decapoda: Pontoniinae), from Inhaca Island, Moçambique. Journal of Crustacean Biology 14, 782–802. doi:10.2307/1548872 Berggren M (1999) Notopontonia platycheles, a little-known shallowwater shrimp from Western Australia (Decapoda: Pontoniinae). Journal of Crustacean Biology 19, 180–187. doi:10.2307/1549559 Bos AR, Fransen CHJM (2018) Nocturnal cleaning of sleeping rabbitfish, Siganus canaliculatus, by the cleaner shrimp, Urocaridella antonbruunii (Decapoda, Palaemonidae). Crustaceana 91, 239–241. doi:10.1163/15685403-00003753 Bruce AJ (1966a) Notes on some Indo-Pacific Pontoniinae. II. Platycaris latirostris Holthuis. Crustaceana 11, 1–9. doi:10.1163/156854066X00397 Bruce AJ (1966b) Notes on some Indo-Pacific Pontoniinae. XI. A reexamination of Philarius lophos Barnard, with the designation of a new genus, Ischnopontonia. Bulletin of Marine Science 16, 584–598. Bruce AJ (1967) Notes on some Indo-Pacific Pontoniinae III–IX. Descriptions of some new genera and species from the western Indian Ocean and the south China Sea. Zoölogische Verhandelingen 87, 1–73. Bruce AJ (1969a) Notes on some Indo-Pacific Pontoniinae. XIII. Propontonia pellucida gen. nov., sp. nov., a new pontoniid shrimp from the Amirante Islands. Crustaceana 17, 141–150. doi:10.1163/156854068X00034 Bruce AJ (1969b) Notes on some Indo-Pacific Pontoniinae. XIV. Observations on Paratypton siebenrocki Balss. Crustaceana 17, 171–186. doi:10.1163/156854068X00061 Bruce AJ (1972a) Notes on Indo-Pacific Pontoniinae XIX. Allopontonia iaini gen. nov., sp. nov., a new echinoid associate from Zanzibar (Decapod, Caridea). Crustaceana 22, 1–12. doi:10.1163/156854072X00633 Bruce AJ (1972b) Notes on some Indo-Pacific Pontoniinae, XVIII. A redescription of Pontonia minuta Baker, 1907, and the occurrence of abbreviated development in the Pontoniinae (Decapoda Natantia, Palaemonidae). Crustaceana 23, 65–75. doi:10.1163/156854072X00075 Bruce AJ (1972c) Pycnocaris chagoae gen. nov., sp. nov., a new shrimp from the Chagos Archipelago (Decapoda Natantia, Gnathophyllidae). Crustaceana 23, 50–64. doi:10.1163/156854072X00066 Bruce AJ (1973a) Gnathophylloides robustus sp. nov., a new commensal gnathophyllid shrimp from Western Australia, with the designation of a new genus Levicaris (Decapoda, Caridea). Crustaceana 24, 17–32. doi:10.1163/156854073X00038

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Bruce AJ (1973b) Notes on some Indo-Pacific Pontoniinae, XXIII. Tectopontonia maziwiae gen. nov., sp. nov., a new coral associate from Tanganyika (Decapoda, Palaemonidae). Crustaceana 24, 169–180. doi:10.1163/​ 156854073X00344 Bruce AJ (1973c) Notes on some Indo-Pacific Pontoniinae, XXIV. Dasycaris zanzibarica sp. nov. from the Western Indian Ocean, with remarks on other species of Dasycaris Kemp, 1922 (Decapoda Natantia). Crustaceana 24, 247–260. doi:10.1163/156854073X00605 Bruce AJ (1974) Periclimenes insolitus sp. nov. (Decapoda Natantia, Pontoniinae), a New Commensal Shrimp from Waikiki Beach, Oahu, Hawaii. Crustaceana 26, 293–307. doi:10.1163/​156854074X00695 Bruce AJ (1975) Notes on some Indo-Pacific Pontoniinae. XXVI. Neoanchistus cardiodytes gen. nov., sp. nov., a new mollusc-associated shrimp from Madagascar (Decapoda, Palaemonidae). Crustaceana 29, 149–165. doi:10.1163/156854075X00162 Bruce AJ (1976a) A further occurrence of Lipkebe holthuisi Chace (Decapoda Natantia, Pontoniinae). Crustaceana 30, 310–312. doi:10.1163/​156854076X00675 Bruce AJ (1976b) Notes on some Indo-Pacific Pontoniinae, XXVII. Apopontona falcirostris gen. nov., sp. nov., from Madagascar. Crustaceana 31, 301–311. doi:10.1163/156854076X00080 Bruce AJ (1976c) A report on a small collection of shrimps from the Kenya National Marine Parks at Malindi, with notes on selected species. Zoölogische Verhandelingen 145, 3–72. Bruce AJ (1976d) A report on some pontoniid shrimps collected from the Seychelle Islands by the F.R.V. Manihine, 1972, with a review of the Seychelles pontoniid shrimp fauna. Zoological Journal of the Linnean Society 59, 89–153. doi:10.1111/j.1096-3642.1976.tb01012.x Bruce AJ (1977) Pontoniine shrimps in the collections of the Australian Museum. Records of the Australian Museum 31, 39–81. doi:10.3853/​ j.0067-1975.31.​1977.233 Bruce AJ (1979a) Ctenopontonia cyphastreophila, a new genus and species of coral associated pontoniine shrimp from Eniwetok Atoll. Bulletin of Marine Science 29, 423–435. Bruce AJ (1979b) A report on a small collection of pontoniine shrimps from Eniwetok Atoll. Crustaceana Supplement 5, 209–230. Bruce AJ (1980) Pontoniine shrimps from the Great Astrolabe Reef, Fiji. Pacific Science 34, 389–400. Bruce AJ (1981) Onycocaridella prima, new genus, new species, a new pontoniine sponge-associate from the Capricorn Islands, Australia (Decapoda, Caridea, Pontoniinae). Journal of Crustacean Biology 1, 241–250. doi:10.2307/1548162 Bruce AJ (1982a) Notes on some Indo-Pacific Pontoniinae, XLI. Orthopontonia, a new genus proposed for Periclimenaeus ornatus Bruce. Crustaceana 43, 163–176. doi:10.1163/156854082X00498 Bruce AJ (1982b) Notes on some Indo-Pacific Pontoniinae, XXXIX. Isopontonia platycheles gen. nov., sp. nov., from the Chesterfield Islands, New Caledonia (Decapoda, Caridea). Crustaceana 42, 54–64. doi:10.1163/156854082X00696 Bruce AJ (1982c) The shrimps associated with Indo-West Pacific echinoderms, with the description of a new species in the genus Periclimenes Costa, 1844 (Crustacea: Pontoniinae). Australian Museum Memoir 16, 191–216. doi:10.3853/j.0067-1967.16.1982.366 Bruce AJ (1983) Expédition Rumphius II (1975) Crustacés parasites, commensaux, etc. (Th. Monod éd.) IX. Crustacés Décapodes (1ère partie: Natantia Pontoniinae). Bulletin du Muséum National d’Histoire Naturelle , Paris, 4e série, A 5, 871–902. Bruce AJ (1985a) Decapod Crustacea: Pontoniinae (MUSORSTOM II). In: Forest J (Ed.) Résultats des Campagnes MUSORSTOM I et II Philippines (1976, 1980). Tome 2. Mémoires du Muséum National d’Histoire Naturelle. Série A, Zoologie 133, 229–260.

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Bruce AJ (1985b) Notes on some Indo-Pacific Pontoniinae, XLII. Miopontonia yongei gen. nov., sp. nov., from the Australian North West shelf (Decapoda, Caridea). Crustaceana 48, 167–178. doi:10.1163/156854085X00855 Bruce AJ (1986a) Chacella, a new palaemonid shrimp genus proposed for Dasycaris kerstitchi Wicksten, 1983. Journal of Crustacean Biology 6, 485–490. doi:10.2307/1548187 Bruce AJ (1986b) Diapontonia maranulus, new genus, new species, a pontoniine shrimp associate of a deep-water echinoid. Journal of Crustacean Biology 6, 125–133. doi:10.2307/1547936 Bruce AJ (1986c) Observations on the family Gnathophyllidae Dana, 1852 (Crustacea: Decapoda). Journal of Crustacean Biology 6, 463–470. doi:10.2307/1548185 Bruce AJ (1986d) A re-examination of Periclimenes (Hamiger) novae-zealandiae Borradaile, 1916, and recognition of the pontoniine shrimp genus Hamiger, Borradaile. Journal of Natural History 20, 911–919. doi:10.1080/00222938600770671 Bruce AJ (1987) Onycocaridites anomodactylus, new genus, new species (Decapoda: Palaemonidae), a commensal shrimp from the Arafura Sea. Journal of Crustacean Biology 7, 771–779. doi:10.2307/1548659 Bruce AJ (1988a) Exopontonia malleatrix, new genus, new species, a palaemonid shrimp from Ashmore Reef, Timor Sea. Journal of Crustacean Biology 8, 122–130. doi:10.2307/1548437 Bruce AJ (1988b) Two new palaemonid shrimps (Crustacea: Decapoda) from the Australian North West shelf. Journal of Natural History 22, 1263–1276. doi:10.1080/00222938800770771 Bruce AJ (1990) Crustacea Decapoda: Deep-sea palaemonoid shrimps from New Caledonian waters. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 6. Mémoires du Muséum National d’Histoire Naturelle, Paris 145, 149–215. Bruce AJ (1991a) Crustacea Decapoda: further deep-sea palaemonoid shrimps from New Caledonian waters. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 9. Mémoires du Muséum National d’Histoire Naturelle, Paris 152, 299–411. Bruce AJ (1991b) Shallow-water palaemonoid shrimps from New Caledonia (Crustacea: Decapoda). In Le benthos des fonds meubles des lagons de Nouvelle-Calédonie. (Ed. Richer de Forges B) pp. 221–279. ORSTOM Editions, Paris. Bruce AJ (1992) Designation of two new pontoniine shrimp genera (Decapoda: Palaemonidae). Journal of Natural History 26, 1273–1282. doi:10.1080/00222939200770721 Bruce AJ (1994) Leander plumosus sp. nov., a new palaemonine shrimp (Crustacea: Palaemonidae) from the Maldive Islands. The Beagle: Records of the Museums and Art Galleries of the Northern Territory 11, 39–51. Bruce AJ (1995) A synopsis of the Indo-West Pacific genera of the Pontoniinae (Crustacea: Decapoda: Palaemonidae). Theses Zoologicae 25, 1–172. Bruce AJ (1996) Crustacea Decapoda: palaemonid shrimps from the IndoWest Pacific region mainly from New Caledonia. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 15. Mémoires du Muséum National d’Histoire Naturelle, Paris 168, 197–267. Bruce AJ (1997) A new pontoniine shrimp genus (Crustacea: Decapoda) from the Yemen, with a note on other species. Journal of Natural History 31, 1213–1222. doi:10.1080/00222939700770651 Bruce AJ (1998a) New keys for the identification of Indo-West Pacific coral associated pontoniine shrimps, with observations on their ecology (Crustacea: Decapoda: Palaemonidae). Ophelia 49, 29–46. doi :10.1080/00785326.1998.10409371 Bruce AJ (1998b) A new species of the genus Brachycarpus (Decapoda, Caridea, Palaemonidae) from New Caledonia. Zoosystema 20, 157–165. Bruce AJ (2000a) The rediscovery of Leandrites stenopus Holthuis, 1950 (Crustacea: Palaemoninae) from Lucinda, Queensland. The Beagle: Records of the Museums and Art Galleries of the Northern Territory 16, 83–88.

Bruce AJ (2000b) Typton manningi and T. capricorniae, a new species, new pontonine shrimps from northern Queensland, with a review of the Indo-West Pacific species of Typton Costa (Decapoda: Palaemonidae). Journal of Crustacean Biology 20, 87–100. doi:10.1163/19372 40X-90000011 Bruce AJ (2004a) Balssia antipodarum sp. nov., the first occurrence of the genus Balssia Kemp in the Indo-West Pacific region (Crustacea: Decapoda: Pontoniinae). Cahiers de Biologie Marine 45, 365–372. Bruce AJ (2004b) A new pontoniine shrimp from Tahiti, French Polynesia (Crustacea, Decapoda, Palaemonidae). Zoosystema 26, 279–289. Bruce AJ (2004c) A partial revision of the genus Periclimenes Costa, 1884 (Crustacea: Decapoda: Palaemonidae). Zootaxa 582, 1–27. doi:10.11646/ zootaxa.582.1.1 Bruce AJ (2005a) A new genus Pontoniopsides, for Pontoniopsis paulae Gore, 1981 (Crustacea: Decapoda: Pontoniinae). Zootaxa 826, 1–4. doi:10.11646/zootaxa.826.1.1 Bruce AJ (2005b) Pontoniine shrimps from Papua New Guinea, with designation of two new genera, Cainonia and Colemonia (Crustacea: Decapoda: Palaemonidae). Memoirs of the Queensland Museum 51, 333–383. Bruce AJ (2006) A new species of the pontoniine shrimp genus Harpilius Dana, 1852, H. spinifer (Crustacea: Decapoda: Palaemonidae), from New Caledonia. Cahiers de Biologie Marine 47, 271–280. Bruce AJ (2007a) A re-definition of the genus Periclimenes Costa, 1844 and the designation of a new genus Margitonia (Crustacea: Decapoda: Pontoniinae). Cahiers de Biologie Marine 48, 403–406. Bruce AJ (2007b) The resurrection of the pontoniine genus Urocaris Stimpson, 1860, (Crustacea: Decapoda: Palaemonidae). Zootaxa 1632, 61–67. doi:10.11646/zootaxa.1632.1.5 Bruce AJ (2008a) Palaemonoid shrimps from the Australian North West shelf. Zootaxa 1815, 1–24. doi:10.11646/zootaxa.1815.1.1 Bruce AJ (2008b) Phycomenes zostericola gen. nov., sp. nov., a new pontoniine shrimp (Crustacea: Decapoda: Palaemonidae) from Moreton Bay, Queensland. Memoirs of the Queensland Museum 54, 219–232. Bruce AJ (2009a) Notes on some Indo-Pacific Pontoniinae, XLVII. Reevaluation of the genera Apopontonia Bruce, 1976, Paraclimenaeus Bruce, 1988 and Climeniperaeus Bruce, 1996. Crustaceana 82, 493–504. doi:10.1163/156854008X400568 Bruce AJ (2009b) Plesiomenaeus poorei gen. nov., sp nov., (Crustacea: Decapoda: Pontoniinae) from Zanzibar. Memoirs of Museum Victoria 66, 25–34. doi:10.24199/j.mmv.2009.66.4 Bruce AJ (2010a) Additions to the genus Phycomenes Bruce, 2008 (Crustacea: Decapoda: Pontoniinae). Zootaxa 2372, 367–368. doi:10.11646/ zootaxa.2372.1.28 Bruce AJ (2010b) Allopontonia alastairi sp. nov., a second species of the genus Allopontonia Bruce, 1972 (Crustacea: Decapoda: Pontoniinae), from the Australian Northwest Shelf. Zootaxa 2372, 33–36. doi:10.11646/zootaxa.2372.1.4 Bruce AJ (2010c) A revision of the systematic position of Periclimenaeus spinimanus Bruce, 1969 (Crustacea: Decapoda: Pontoniinae) and the designation of Anisomenaeus gen. nov. Zootaxa 2372, 405–414. doi:10.11646/​zootaxa.2372.1.32 Bruce AJ (2010d) Typtonoides nieli gen. nov., sp. nov., a new pontoniine shrimp (Crustacea: Decapoda: Palaemonidae) from the Chesterfield Island. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 26, 69–74. Bruce AJ (2011a) Notes on some Indo-Pacific Pontoniinae, L. Typtonychus spp. nov., from northwest Australia. Crustaceana 84, 1337–1351. doi:10.1163/156854011X594786 Bruce AJ (2011b) Notes on some Indo-Pacific Pontoniinae, XLIX. Onycocaris balsii sp. nov., from northern Australia, with the designation of O. fujinoi sp. nov. Crustaceana 84, 477–490. doi:10.1163/​ 001121611X554337

6 – Caridea – shrimps

Bruce AJ (2011c) Notes on some Indo-Pacific Pontoniinae, XLVIII. Onycocaris nieli sp. nov., a new pontoniine shrimp from Heron Island, Queensland. Crustaceana 84, 319–330. doi:10.1163/001121610X551836 Bruce AJ (2012) Notes on Indo-Pacific Pontoniinae, LII. A third species of the genus Manipontonia Bruce, Okuno & Li, 2005. Crustaceana 85, 1377–1383. doi:10.1163/156854012X651628 Bruce AJ (2013) Identification aid for the Indo-West Pacific species of Periclimenaeus Borradaile, 1915 (Crustacea: Decapoda: Caridea: Pontoniinae) using ambulatory dactyli. Memoirs of the Queensland Museum 56, 647–664. Bruce AJ (2014a) Nippontonia ashmoreiensis sp. nov., (Crustacea: Decapoda; Pontoniinae) from Ashmore Reef, Western Australia. Zootaxa 3847, 437–445. doi:10.11646/zootaxa.3847.3.8 Bruce AJ (2014b) Notes on some Indo-Pacific Pontoniinae, LIV. New Periclimenes species (Decapoda, Pontoniinae), from Cartier Island reef, Western Australia. Crustaceana 87, 64–82. doi:10.1163/1568540300003272 Bruce AJ, Okuno J (2010) Designation of a new genus Lipkemenes, with supplementary description and range extension of its type species, L. lanipes (Kemp, 1922) (Decapoda, Palaemonidae). In: Fransen C, De Grave S, Ng P (Eds) Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume. Crustaceana Monographs 14, 159–171. doi:10.1163/9789047427759_008 Carvalho FL, De Grave S, Mantelatto FL (2017) An integrative approach to the evolution of shrimps of the genus Palaemon (Decapoda, Palaemonidae). Zoologica Scripta 46, 473–485. doi:10.1111/zsc.12228 Chace FA, Bruce AJ (1993) The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 6: superfamily Palaemonoidea. Smithsonian Contributions to Zoology 543, 1–152. doi:10.5479/si.00810282.543 Chávez-Solís EM, Mejía-Ortíz LM, Simões N (2018) Predatory behavior of the cave shrimp Creaseria morleyi (Creaser, 1936) (Caridea: Palaemonidae), the blind hunter of the Yucatán cenotes, Mexico. Journal of Crustacean Biology 38, 1–7. doi:10.1093/jcbiol/rux098 Chow LH, De Grave S, Tsang LM (2020) The family Anchistioididae Borradaile, 1915 (Decapoda: Caridea) is a synonym of Palaemonidae Rafinesque, 1815 based on molecular and morphological evidence. Journal of Crustacean Biology 40, 277–287. doi:10.1093/jcbiol/ruaa012 Chow L, De Grave S, Ming Tsang L (2021) Evolution of protective symbiosis in palaemonid shrimps (Decapoda: Caridea) with emphases on host spectrum and morphological adaptations. Molecular Phylogenetics and Evolution 162, 107201. doi:10.1016/j.ympev.2021.107201 d’Udekem d’Acoz C (2001) Description of Gnathophylleptum tellei gen. nov., sp. nov., a remarkable new gnathophyllid shrimp from Canary Islands (Crustacea, Decapoda, Caridea). Bulletin de l’Institut Royal des Sciences Naturelles de Belgique 71, 113–125. De Grave S, Ashelby CW (2013) A re-appraisal of the systematic status of selected genera in Palaemoninae (Crustacea: Decapoda: Palaemonidae). Zootaxa 3734, 331–344. doi:10.11646/zootaxa.3734.3.3 De Grave S, Fransen CHJM, Page TJ (2015) Let’s be pals again: major systematic changes in Palaemonidae (Crustacea: Decapoda). PeerJ 3, e1167. doi:10.7717/peerj.1167 Ďuriš Z (2010) Periclimenes laevimanus sp. nov. from Vietnam, with a review of the Periclimenes granulimanus species group (Crustacea: Decapoda: Palaemonidae: Pontoniinae). In: De Grave S, Fransen CHJM (Eds) Contributions to shrimp taxonomy. Zootaxa 2372, 106–125. Ďuriš Z, Bruce AJ (1995) A revision of the ‘petitthouarsii’ species-group of the genus Periclimenes Costa, 1844 (Crustacea: Decapoda: Palaemonidae). Journal of Natural History 29, 619–671. doi:10.1080/00222939500770221 Ďuriš Z, Horká I (2008) A new species of the genus Leptomenaeus Bruce, 2007 (Crustacea: Decapoda: Pontoniinae) from Vietnam. Zootaxa 1872, 45–58. doi:10.11646/zootaxa.1872.1.4

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García-Dávila CR, Magalhães C (2003) Revisão taxonômica dos camarões de água doce (Crustacea: Decapoda: Palaemonidae, Sergestidae) da Amazônia Peruana. Acta Amazonica 33, 663–686. doi:10.1590/ S0044-59672003000400013 Gore RH (1981) Three new shrimps, and some interesting new records of decapod Crustacea from a deep-water coral reef in the Florida Keys. Proceedings of the Biological Society of Washington 94, 135–162. Grippa GB, d’Udekem d’Acoz C (1996) The genus Periclimenes Costa, 1844 in the Mediterranean Sea and the Northeastern Atlantic Ocean: review of species and description of Periclimenes sagittifer aegylios subsp. nov. (Crustacea, Decapoda, Caridea, Pontoniinae). Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano 135(1994), 401–412. Gurney R (1936) Notes on some decapod Crustacea of Bermuda, III: the larvae of the Palaemonidae. Proceedings of the Zoological Society of London 1936, 619–623, pls 1–4. Heard RW (1986) Pontoniine shrimps (Decapoda: Caridea: Palaemonidae) of the Northwest Atlantic. I. The genus Neopontonides Holthuis, 1951, with the description of N. chacei, new species, and the erection of Pseudopontonides, new genus, to receive N. principes Criales, 1980. Journal of Crustacean Biology 6, 471–484. doi:10.2307/1548186 Heard RW, Spotte S, Bubucis PM (1993) Pontoniine shrimps (Decapoda: Caridea: Palaemonidae) of the Northwest Atlantic. III. Neopericlimenes thornei, new genus, new species, from Pine Cay, Turks and Caicos Islands, British West Indies. Journal of Crustacean Biology 13, 793–800. doi:10.2307/1549109 Hipeau-Jacquotte R (1971) Notes de faunistique et de biologie marines de Madagascar, V. Platypontonia hyotis nov. sp. (Decapoda Natantia, Pontoniinae). Crustaceana 20, 125–140. doi:10.1163/156854069X00150 Hobbs HH (1973) Three new troglobitic decapod crustaceans from Oaxaca, Mexico. Bulletin of the Association for Mexican Cave Studies 5, 25–38. Holthuis LB (1951) A general revision of the Palaemonidae (Crustacea Decapoda Natantia) of the Americas. I. The subfamilies Euryrhynchidae and Pontoniinae. Allan Hancock Foundation Occasional Paper 11, 1–332. Holthuis LB (1952) A general revision of the Palaemonidae (Crustacea Decapoda Natantia) of the Americas. II. The subfamily Palaemoninae. Allan Hancock Foundation Occasional Paper 12, 1–396. Holthuis LB (1978) Redescription of Coutierea agassizi (Coutière, 1901) Ccrustacea Decapoda, Palaemonidae). Zoölogische Mededeelingen 52, 313–320. Holthuis LB (1986) Some Pontoniinae (Crustacea: Decapoda: Palaemonidae) from southern Oman. Zoölogische Mededeelingen 60, 263–272. Holthuis LB (1993) The Recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda) with an appendix on the Order Amphionidacea. revised edn. Nationaal Natuuristorisch Museum, Leiden. Horká I, Fransen CHJM, Ďuriš Z (2016) Two new species of shrimp of the Indo-West Pacific genus Hamodactylus Holthuis, 1952 (Crustacea: Decapoda: Palaemonidae). European Journal of Taxonomy 188, 1–26. doi:10.5852/ejt.2016.188 Jayachandran KV (1992) On the genus Leptocarpus Holthuis, 1950, with the description of a new species (Decapoda, Palaemonidae). Mahasagar 25, 129–134. Kemp S (1922) Notes on Crustacea Decapoda in the Indian Museum. XV. Pontoniinae. Records of the Indian Museum 24, 113–288, pls 1–9. Kensley B, Walker I (1982) Palaemonid shrimps from the Amazon Basin, Brazil (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology 362, 1–28. doi:10.5479/si.00810282.362 Komai T, Yamada Y (2015) A new species of the palaemonid shrimp genus Palaemonella Dana, 1852 (Crustacea: Decapoda: Caridea) from Okinawa Island, Ryukyu Islands, Japan. Zootaxa 3964, 352–362. doi:10.11646/​ zootaxa.3964.3.4

Kou Q, Li X (2016) A new palaemonid shrimp of the “Philarius gerlachei (Nobili, 1905) species complex” (Crustacea: Decapoda: Palaemonidae) from Hainan Island, South China Sea. Raffles Bulletin of Zoology 64, 269–277. Kou Q, Li X, Chan T-Y, Chu KH, Gan Z (2013) Molecular phylogeny of the superfamily Palaemonoidea (Crustacea: Decapoda: Caridea) based on mitochondrial and nuclear DNA reveals discrepancies with the current classification. Invertebrate Systematics 27, 502–514. doi:10.1071/IS13005 Kou Q, Li X, Bruce AJ (2016a) Designation of a new genus Bathymenes for the deep-sea pontoniine shrimps of the ‘Periclimenes alcocki species group’ (Decapoda, Caridea, Palaemonidae), with a checklist of the species assigned to the genus. Chinese Journal of Oceanology and Limnology 34, 170–176. doi:10.1007/s00343-015-4359-4 Kou Q, Li X, Gan Z (2016b) Re-evaluation of the systematic status of two deep-water recorded Periclimenes species (Decapoda : Palaemonidae), with a preliminary analysis on the evolution of certain echinoidassociated palaemonid shrimps and related taxa. Invertebrate Systematics 30, 421–430. doi:10.1071/IS15064 Li X (2000) Catalog of the genera and species of Pontoniinae Kingsley, 1878. Xueyuan Press, Beijing. Li X (2009) Sandimenes nov. gen., for Periclimenes hirsutus Bruce, 1971 (Decapoda, Caridea, Pontoniinae). Crustaceana 82, 881–896. doi:10.1163/156854009X427414 Li X, Bruce AJ (2006) Further Indo-West Pacific palaemonoid shrimps (Crustacea: Decapoda: Palaemonoidea), principally from the New Caledonian region. Journal of Natural History 40, 611–738. doi:10.1080/00222930600763627 Li X, Poupin J (2009) Report of some species of Palaemonidae (Decapoda: Caridea) from Clipperton Island. Journal of Crustacean Biology 29, 579–594. doi:10.1651/08-3084.1 Li X, Bruce AJ, Manning RB (2004) Some palaemonid shrimps (Crustacea: Decapoda) from Northern South China Sea, with descriptions of two new species. Raffles Bulletin of Zoology 52, 513–553. Li X, Mitsuhashi M, Chan T-Y (2008) Deep-sea pontoniines (Decapoda: Palaemonidae) from the Philippine “Panglao 2005” Expedition, with descriptions of four new species. Journal of Crustacean Biology 28, 385–411. doi:10.1651/0278-0372(2008)028[0385:DPDPFT]2.0.CO;2 Makombu JG, Stomeo F, Oben PM, Tilly E, Stephen OO, et al. (2019) Morphological and molecular characterization of freshwater prawn of genus Macrobrachium in the coastal area of Cameroon. Ecology and Evolution 9, 14217–14233. doi:10.1002/ece3.5854 Marin I (2006) Description of Crinotonia anastasiae, new genus, new species, a new crinoid associated pontoniine shrimp (Crustacea: Caridea) from Nha Trang Bay, Vietnam, with inclusion of Periclimenes attenuatus Bruce, 1971, in the new genus. Raffles Bulletin of Zoology 54, 321–340. Marin I (2007a) Diversity of the genus Pontonides (Caridea, Palaemonidae, Pontoniinae) in Nhatrang Bay, Vietnam, with description of new taxa. Zootaxa 1635, 1–21. doi:10.11646/zootaxa.1635.1.1 Marin I (2007b) Pontoniine shrimps (Decapoda: Caridea: Palaemonidae) inhabiting boring sponges (Porifera: Demospongia) from Nhatrang Bay, Vietnam, with descriptions of three new species. Zoölogische Mededeelingen 81, 217–240. Marin IN (2007c) Notes on taxonomy and biology of the symbiotic shrimp Vir euphyllius Marin et Anker, 2005 (Decapoda: Palaeomonidae: Pontoniinae), associated with scleractinian corals Euphyllia spp. (Cnidaria: Caryophyllidae). Invertebrate Zoology 4, 15–23. doi:10.15298/​ invertzool.04.1.02 Marin I (2008a) Description of two new species from the genera Palaemonella Dana, 1852 and Vir Holthuis, 1952 (Crustacea: Caridea: Palaemonidae: Pontoniinae). Zoölogische Mededeelingen 82, 375–390.

6 – Caridea – shrimps

Marin I (2008b) Notes on morphology and ecological difference between species of pontoniine shrimp genus Crinotonia Marin (Caridea: Palaemonidae) associated with shallow-water feather stars Phanogenia spp. (Crinoidea: Comasteridae). Zootaxa 1764, 19–24. doi:10.11646/ zootaxa.1764.1.2 Marin I (2008c) Taxonomic position of Pontonides sympathes De Ridder & Holthuis and species of the genus Veleronia Holthuis (Decapoda, Palaemonidae, Pontoniinae) with the description of a new genus. Zootaxa 1932, 1–17. doi:10.11646/zootaxa.1932.1.1 Marin I (2009a) Crinoid-associated shrimps of the genus Laomenes A.H. Clark, 1919 (Caridea: Palaemonidae: Pontoniinae): new species and probable diversity. Zootaxa 1971, 1–49. doi:10.11646/zootaxa.1971.1.1 Marin I (2009b) A review of the pontoniine shrimp genus Rapipontonia Marin, 2007 (Decapoda: Caridea: Palaemonidae), with the description of a new species from the Indo-West Pacific. Zootaxa 2289, 1–17. doi:10.11646/zootaxa.2289.1.1 Marin I (2009c) Sandyella gen. nov., a new shrimp genus for the Eastern Pacific species Chacella tricornuta Hendrickx, 1990 and Chacella mclaughlinae Li, 2006 (Caridea, Palaemonidae, Pontoniinae). Crustaceana 82, 913–918. doi:10.1163/156854009X427432 Marin I (2011) Fennera holthuisi sp. nov., a new coral-associated pontoniine shrimp (Decapoda: Palaemonidae: Pontoniinae) from Moorea, French Polynesia. Zootaxa 2890, 29–39. doi:10.11646/zootaxa.2890.1.3 Marin I (2012) New records and associations of palaemonid shrimps (Crustacea: Decapoda: Caridea: Palaemonidae: Pontoniinae) from the Nhatrang Bay, Vietnam; with taxonomic remarks on some species from the Indo-West Pacific region. In Benthic Fauna of the Bay of Nhatrang, Southern Vietnam. (Eds Britayev TA, Pavlov DS) pp. 345–405. KMK Scientific Press, Moscow. Marin I (2014a) A new species of the pontoniine shrimp genus Hamopontonia Bruce, 1970 associated with caryophyllid coral Euphyllia glabrescens (Chamisso & Eysenhardt, 1821) in Nhatrang Bay, Vietnam. Zootaxa 3815, 131–140. doi:10.11646/zootaxa.3815.1.9 Marin IN (2014b) The first record of an association between a pontoniine shrimp (Crustacea: Decapoda: Palaemonidae: Pontoniinae) and a thalassematid spoon worm (Echiura: Thalassematidae), with the description of a new shrimp species. Zootaxa 3847, 557–566. doi:10.11646/zootaxa.3847.4.5 Marin IN (2015) New records of pontoniine shrimp genus Onycocaris Nobili, (Crustacea: Decapoda: Palaemonidae) from the Indo-Pacific with the description of two new species from French Polynesia. Arthropoda Selecta 24, 283–302. doi:10.15298/arthsel.24.3.06 Marin I, Anker A (2011) A partial revision of the Philarius gerlachei (Nobili, 1905) species complex (Crustacea, Decapoda, Palaemonidae), with description of four new species. Zootaxa 2781, 1–28. doi:10.11646/ zootaxa.2781.1.1 Marin I, Chan T-Y (2006) Two new genera and a new species of crinoidassociated pontoniine shrimps (Decapoda: Caridea: Palaemonidae). Journal of Crustacean Biology 26, 524–539. doi:10.1651/S-2705.1 Marin I, Chan T-Y (2013) Typtonomenaeus formosanus gen. et sp. nov., a new sponge-associated pontoniine shrimp (Crustacea: Decapoda: Palaemonidae: Pontoniinae) from northern Taiwan. Zootaxa 3664, 45–56. doi:10.11646/zootaxa.3664.1.2 Marin I, Chan T-Y (2014) Deep water echinoid-associated pontoniine shrimp “Periclimenes hertwigi Balss, 1913” species group (Crustacea: Decapoda: Caridea: Palaemonidae): species review, description of a new genus and species from Philippines. Zootaxa 3835, 301–324. doi:10.11646/zootaxa.3835.3.1 Marin I, Paulay G (2010) Pinnotherotonia rumphiusi gen. et sp. nov., a new furry bivalve-associated pontoniine shrimp (Crustacea: Decapoda: Palaemonidae) from Palau. Zootaxa 2636, 37–48. doi:10.11646/zootaxa.​ 2636.1.3

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Marin I, Chan T-Y, Okuno J (2012) Crinoid-associated pontoniine shrimps of the genus Laomenes Clark, 1919 (Decapoda, Caridea, Palaemonidae) from PANGLAO 2004 and KUMEJIMA 2009 Expeditions, with description of two new species. Zootaxa 3367, 103–114. doi:10.11646/zootaxa.3367.1.10 Martínez-Mayén M, Román-Contreras R, Boyko CB (2006) A new species of Periclimenes Costa, 1844 (Crustacea: Decapoda: Palaemonidae) from the Caribbean coast of Quintana Roo, Mexico, and a key for the “iridescens” complex. Proceedings of the Biological Society of Washington 119, 32–42. doi:10.2988/0006-324X(2006)119[32:ANSOPC]2.0.CO;2 Mitsuhashi M, Chan T-Y (2006) A new genus and species of deep-water pontoniine shrimp (Decapoda, Caridea, Palaemonidae) from Taiwan. Zoosystema 28, 389–398. Mitsuhashi M, Chan T-Y (2007) A new pontoniine shrimp genus and species (Crustacea: Decapoda: Palaemonidae) from the Philippine Panglao 2004 expedition. Raffles Bulletin of Zoology Supplement 16, 1–6. Mitsuhashi M, Takeda M (2008) Identity of the coral-associated pontoniine shrimp species, Coralliocaris nudirostris (Heller, 1816) and C. venusta Kemp, 1922 (Crustacea: Decapoda: Palaemonidae), with descriptions of two new species. Zootaxa 1703, 1–24. doi:10.11646/ zootaxa.1703.1.1 Mitsuhashi M, Chan T-Y, Jeng M-S (2005) First records of the two caridean families, Gnathophyllidae Dana, 1852 and Hymenoceridae Ortmann, 1890 (Decapoda, Palaemonoidea) from Taiwan. Crustaceana 78, 987–1000. doi:10.1163/156854005775197271 Mitsuhashi M, Li X, Chan T-Y (2010) Neoclimenes holthuisi n. gen., n. sp., a new deep-sea pontoniine shrimp from the South China Sea (Decapoda, Palaemonidae). In: Fransen C, De Grave S, Ng P (Eds) Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume. Crustaceana Monographs 14, 515–527. doi:10.1163/9789047427759_035 Odijk M, Fransen CHJM (2017) A new sponge associated shrimp species of the Indo-West Pacific genus Paraclimenaeus (Decapoda, Caridea, Palaemonidae). Crustaceana 90, 1099–1115. doi:10.1163/1568540300003660 Okuno J (1999) Izucaris masudai, new genus, new species (Decapoda: Caridea: Palaemonidae), a sea anemone associate from Japan. Journal of Crustacean Biology 19, 397–407. doi:10.2307/1549247 Okuno J (2009) Pliopontonia harazakii sp. nov., a new species of coralassociated shrimp (Decapoda: Caridea: Palaemonidae) from Yakushima Island, southern Japan. Bulletin of the National Museum of Nature and Science. Series A, Zoology 3(Supplement), 105–113. Okuno J (2017) Designation of a new genus Michaelimenes (Decapoda: Caridea: Palaemonidae), with new host record and range extension of its type species, M. perlucidus (Bruce, 1969). Nauplius 25, e2017013. doi:10.1590/2358-2936e2017013 Okuno J, Bruce AJ (2010) Designation of Ancylomenes gen. nov., for the ‘Periclimenes aesopius species group’ (Crustacea: Decapoda: Palaemonidae), with the description of a new species and a checklist of congeneric species. In: De Grave S, Fransen CHJM (Eds) Contributions to shrimp taxonomy. Zootaxa 2372, 85–105. Olthof G, Becking LE, Fransen CHJM (2018) On a collection of deepwater shrimp (Crustacea, Decapoda) from the Dutch Caribbean, with the description of a new species of Pseudocoutierea. Zootaxa 4415, 533–548. doi:10.11646/zootaxa.4415.3.7 Park JH, De Grave S, Kim W (2019) A new species of the genus Cristimenes Ďuriš & Horká, 2017 (Decapoda, Caridea, Palaemonidae). ZooKeys 852, 53–71. doi:10.3897/zookeys.852.34959 Pereira G (1997) A cladistic analysis of the freshwater shrimps of the family Palaemonidae (Crustacea, Decapoda, Caridea). Acta Biologica Venezuelica 17(Supplement 1), 1–69. Prakash S, Baeza JA (2018) A new species of shrimp of the genus Urocaridella Borradaile, 1915 (Decapoda: Caridea: Palaemonidae) from

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Papua New Guinea. Journal of Crustacean Biology 38, 206–214. doi:10.1093/jcbiol/rux113 Sankolli KN, Shenoy S (1979) On a new genus and a new species of a subterranean prawn Troglindicus phreaticus (Caridea, Palaemonidae). Bulletin of Fisheries, Faculty of Konkan Agricultural University, India 1, 83–91. Shaw JK, Heard RW, Hopkins TS (1977) Notes on the biology of the pontoniine shrimp Lipkebe holthuisi Chace, with a description of the male. Proceedings of the Biological Society of Washington 90, 284–290. Short JW, Humphrey CL, Page TJ (2013) Systematic revision and reappraisal of the Kakaducarididae Bruce (Crustacea: Decapoda: Caridea) with the description of three new species of Leptopalaemon Bruce & Short. Invertebrate Systematics 27, 87–117. doi:10.1071/IS12016 Snijders N, Fransen CHJM (2010) Pseudopontonides plumosus sp. nov., a new cnidarian-associated pontoniine shrimp (Crustacea, Decapoda, Palaemonidae) from Curaçao. Zootaxa 2372, 7–14. doi:10.11646/zootaxa.2372.1.2 Šobáňová A, Ďuriš Z (2018) Kaviengella jeffkinchi, a new genus and species of symbiotic shrimp (Crustacea: Decapoda: Palaemonidae) from Papua New Guinea. Zootaxa 4415, 118–134. doi:10.11646/zootaxa.4415.1.5 Šobáňová A, Duriš Z (2021) Unexpected diversity in the sponge-associated shrimps Onycocaridella Bruce, 1981 (Crustacea : Decapoda : Palaemonidae) revealed by bulk collecting techniques and molecular tools. Invertebrate Systematics 35, 361–393. Šobáňová A, Lin CW, Ďuriš Z (2017) Periclimenoides tyrannodentatus, a new species of symbiotic shrimps (Crustacea: Decapoda: Palaemonidae) from Taiwan. Zootaxa 4363, 281–290. doi:10.11646/zootaxa.4363.2.7 Soledade GO, Santos GG, Pinheiro U, Almeida AO (2017) New records of association between caridean shrimps (Decapoda) and sponges (Porifera) in Abrolhos Archipelago, northeastern Brazil. Nauplius 25, e201702. doi:10.1590/2358-2936e2017027 Suzuki H (1971) On some commensal shrimps found in the western region of Sagami Bay. Crustacean Research 4, 1–31. Villalobos JL, Alvarez F, Iliffe TM (1999) New species of troglobitic shrimps from Mexico, with the description of Troglomexicanus, new genus (Decapoda: Palaemonidae). Journal of Crustacean Biology 19, 111–122. doi:10.2307/1549553 Villalobos Hiriart JL, Nates Rodriguez JC, Cantú Diaz Barriga A (1989) Revisión de los géneros Cryphiops Dana, 1852 y Bithynops Holthuis, 1973, de la familia Palaemonidae (Crustacea, Decapoda), y descripción de una especie nueva para el Estado de Chiapas, México. Anales del Instituto de Biologiá, Universidad Nacional Autónoma de México 60, 159–184. Wowor D, Choy SC (2001) The freshwater prawns of the genus Macrobrachium Bate, 1868 (Crustacea: Decapoda: Palaemonidae) from Brunei Darussalam. Raffles Bulletin of Zoology 49, 269–289. Wowor D, Ng PKL (2010) On two new genera of Asian prawns previously assigned to Macrobrachium (Crustacea: Decapoda: Caridea: Palaemonidae). Zootaxa 2372, 37–52. doi:10.11646/zootaxa.2372.1.5

Superfamily Pandaloidea Haworth, 1825 Pandaloid shrimps are most easily recognised by a non-chelate or microscopically chelate first pereopod. The superfamily comprises two families, Chlorotocellidae inhabiting shallow marine environments, and Pandalidae the other usually living in the cold-water deep sea. The classification of

the group was rearranged on the basis of molecular and morphological evidence by Liao et al. (2019). Using this framework Komai et  al. (2019) introduced Chlorotocellidae and synonymised the family Thalassocarididae, which was recognised in Holthuis’s (1993) keys, with Pandalidae. Diagnosis. Pereopods 1, 2 dissimilar, first simple (Figs 6.46g, 6.48k) or with minute chela (Fig. 6.48m). Pereopod 2 carpus multiarticulate (Fig.  6.48n–r). Exopod sometimes present on maxilliped 3, present on pereopods 1–5. References Holthuis LB (1993) The Recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda) with an appendix on the Order Amphionidacea. revised edn. Nationaal Natuuristorisch Museum, Leiden. Komai T, Chan T-Y, De Grave S (2019) Establishment of a new shrimp family Chlorotocellidae for four genera previously assigned to Pandalidae (Decapoda, Caridea, Pandaloidea). Zoosystematics and Evolution 95, 391–402. doi:10.3897/zse.95.35999 Liao Y, Ma KY, De Grave S, Komai T, Chan T-Y, et al. (2019) Systematic analysis of the caridean shrimp superfamily Pandaloidea (Crustacea: Decapoda) based on molecular and morphological evidence. Molecular Phylogenetics and Evolution 134, 200–210. doi:10.1016/j.ympev.​ 2019.02.006

Chlorotocellidae Komai, Chan & De Grave, 2019 Figure 6.46, Plate 11h, i Chlorotocellids are small species inhabiting shallow subtidal tropical to temperate waters and are often associated with cnidarians (Hayashi 1975; Komai et al. 2019). The establishment of the family was based on molecular and morphological evidence (Liao et  al. 2019). Chlorotocellids are most easily differentiated from pandalids by pereopod 2 carpus of three segments and the absence of ventral rostral teeth. Komai et al. (2019) diagnosed all the genera with distributions, and listed the species. Diagnosis. Rostrum absent, or ventral margin with no or few teeth or fringe of setae. Thoracic sternites unornamented. Pleonite 6 without posteroventral tooth; posterolateral process rounded or truncate. Eyestalks subcylindrical, cornea distinctly shorter than eyestalk. Pereopod 2 carpus of 3 segments (Fig.  6.46j). Telson with dorsolateral spiniform setae adjacent to lateral margins (Fig. 6.46e). Implicit generic attributes. Carapace without supraorbital spine. Pleonite 3 evenly rounded dorsally. Mandibular palp absent. Pereopods 3–5 propodus not widened distally, simple.

Key to genera of Chlorotocellidae 1. –

Carapace with supraorbital spine (Fig. 6.46a). Mandibular palp 2-articled (Fig. 6.46f). Rostrum long, slender (Fig. 6.46a)������������������������������������������������������������������������������������������������������������������������������������������������������� Chlorotocella Carapace without supraorbital spine (Fig. 6.46b–d). Mandible without palp. Rostrum short, deep or absent����������������2

6 – Caridea – shrimps

159

Fig. 6.46.  Chlorotocellidae. Habitus: a, Chlorotocella spinicaudus (H. Milne Edwards, 1837). Carapace: b, Anachlorocurtis; c, Chlorocurtis. Carapace, pleon: d, Miropandalus. Telson: e, Chlorocurtis. Mandible: f, Chlorotocella. Maxilliped 1: g, Chlorotoceella. Pereopod 1: h, i, Anachlorocurtis (with detail of end of dactylus). Pereopods 2, 3: j, k,Chlorocurtis.

2. – 3. –

Pereopods 3–5 propodus widened distally, subchelate (Fig. 6.46k). Rostrum dorsal margin crested, with 5–7 regular fixed or movable teeth. Carapace midline without prominent processes (Fig. 6.46c)������������Chlorocurtis Pereopods 3–5 propodus not widened, simple. Rostrum absent or with irregular teeth. Carapace midline with 2 prominent processes�������������������������������������������������������������������������������������������������������������������������������������������������������3 Rostrum present, formed by obliquely truncate tooth, with 1–3 teeth on oblique anterior margin, none on dorsal margin (Fig. 6.46b). Pleonite 3 evenly convex dorsally������������������������������������������������������������������ Anachlorocurtis Rostrum absent. Pleonite 3 with high triangular crest on posterior half of dorsal midline (Fig. 6.46d)����Miropandalus Anachlorocurtis Hayashi, 1975

Diagnosis. Rostrum short, deep, with irregular teeth; carapace midline with 2 prominent processes, postrostral and cardiac. Maximum cl. 2.2 mm. Subtidal (0–40 m; associated with antipatharian corals). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Australasia (Kermadec Is). 3 species (Ahyong 2015; Horká et al. 2014: rediagnosis).

Chlorocurtis Kemp, 1925 Diagnosis. Rostrum short, deep, crested, with 5–7 teeth including 2 or 3 postrostral; carapace midline without prominent processes. Pereopods 3–5 propodus widened distally, subchelate. Maximum cl. 2 mm (Pl. 11h). Subtidal (0–10 m; seagrasses). Indo-West Pacific. 2 species (Gan and Li 2018: redescription).

Chlorotocella Balss, 1914 Diagnosis. Rostrum long, slender, with dorsal teeth near base; carapace with supraorbital spine; carapace midline without

prominent processes. Mandibular palp 2-articled. Maximum cl. 6 mm (Pl. 11i). Intertidal–shelf (0–80 m; free living or associated with gorgonarians and hydroids). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia. 2 species (Komai et al. 2019: discussions of species; Li and Davie 2006).

Miropandalus Bruce, 1983 Diagnosis. Rostrum short, deep, crested, with 5–7 teeth including 2 or 3 postrostral; carapace midline with 2 prominent processes, postrostral and cardiac. Pleonite 3 with triangular crest on posterior half of dorsal midline. Maximum cl. 5 mm. Intertidal–shelf (0–58 m; associated with antipatharian corals). Central Indo-Pacific. 1 species (Bruce 1983, 1991). References Ahyong ST (2015) Decapod crustacea of the Kermadec Biodiscovery Expedition 2011. Bulletin of the Auckland Museum 20, 405–442. Bruce AJ (1983) Miropandalus hardingi, new genus, new species, a bizarre commensal pandalid shrimp from the Marshall Islands. Journal of Crustacean Biology 3, 482–490. doi:10.2307/1548149

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Marine Decapod Crustacea

Bruce AJ (1991) A second occurrence of the bizarre shrimp Miropandalus hardingi (Crustacea: Decapoda: Pandalidae). The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 8, 11–14. Gan Z, Li X (2018) Four new records of caridean shrimp (Crustacea: Decapoda: Caridea) from the East China Sea and South China Sea. Acta Oceanologica Sinica 37, 212–217. doi:10.1007/s13131-018-1325-y Hayashi K-I (1975) Anachlorocurtis commensalis gen. nov., sp. nov. (Crustacea, Decapoda, Pandalidae), a new pandalid shrimp associated with antipatharian corals from Central Japan. Annotationes Zoologicae Japonenses 48, 172–182. Horká I, De Grave S, Ďuriš Z (2014) A new species of shrimp of the genus Anachlorocurtis Hayashi, 1975 from the Red Sea, with range extension of A. commensalis Hayashi, 1975 (Crustacea, Decapoda, Pandalidae). ZooKeys 407, 9–28. doi:10.3897/zookeys.407.7457 Komai T, Chan TY, De Grave S (2019) Establishment of a new shrimp family Chlorotocellidae for four genera previously assigned to Pandalidae (Decapoda, Caridea, Pandaloidea). Zoosystematics and Evolution 95, 391–402. doi:10.3897/zse.95.35999 Li X, Davie PJF (2006) An account of the pandaloid shrimps (Crustacea: Decapoda: Caridea) in the collections of the Queensland Museum. Memoirs of the Queensland Museum 52, 151–170. Liao Y, Ma KY, De Grave S, Komai T, Chan T-Y, et al. (2019) Systematic analysis of the caridean shrimp superfamily Pandaloidea (Crustacea: Decapoda) based on molecular and morphological evidence. Molecular Phylogenetics and Evolution 134, 200–210. doi:10.1016/j. ympev.2019.02.006

Pandalidae Haworth, 1825 Figures 6.47, 6.48, Plate 12a–h In pandalids, as in chlorotocellids, pereopod 1 is simple (Fig.  6.48k) or minutely chelate (Fig.  6.48l–m). Unlike the

small coastal chlorotocellids, most pandalids are large deep-water, sometimes, mesopelagic shrimps. A few species are sufficiently common, especially of the most speciose genera Pandalus and Plesionika, to be fished commercially. The key, based on those in Chace (1985) and Holthuis (1993), relies on strap-like epipods, features difficult to see. Epipods are short posteriorly-directed curved appendages at the base of the legs and covered by the carapace (Fig. 6.48u). Diagnosis. Rostrum usually well developed, ventral margin with few or series of teeth accompanied by row(s) of short setae. Thoracic sternites 6 and 7 each with paired prominences on either side of median keel; thoracic sternite 8 with transverse carina bearing submedian spines or teeth (Fig.  6.48w). Pleonite 6 with small posteroventral tooth; posterolateral process terminating in small tooth (Fig. 6.47a, b, r). Eyestalks pyriform or subpyriform, cornea usually distinctly wider than eyestalk. Pereopod 2 carpus variable, never of 3 segments. Telson with dorsolateral spiniform setae on dorsolateral ridges (Fig. 6.48s, t). Implicit generic attributes. Dioecious. Rostrum fixed. Carapace without supraorbital spine, without lateral carinae. Pleonites without posterodorsal keels. Telson acute. Cornea much wider than eyestalk; eye with ocellus. Maxilla scaphognathite posterior lobe broadly rounded. Pereopod 2 carpus multiarticulate; pereopod 2 palm barely swollen if at all. Male pleopod 1 endopod laminar, not convoluted. Maxilliped 3 without exopod; epipod on pereopods 1–4; arthrobranchs on pereopods 1–4.

Key to genera of Pandalidae 1. – 2. – 3. – 4. – 5. – 6. – 7. –

Pereopod 2 carpus undivided or of 2 segments. Male pleopod 1 endopod enlarged, convoluted, spinulose (Fig. 6.48u)������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Pereopod 2 carpus usually composed of 2 or more segments (Fig. 6.48p). Male pleopod 1 endopod laminar, not convoluted�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Carapace without supraorbital spine (Fig. 6.47a–n). Pleonite 6 without dorsomedian spine on posterior margin. Telson bifurcate (Fig. 6.48s). Scaphocerite unarmed laterally. Pereopods 1–4 with epipods. Pereopod 2 carpus with 2 segments�����������������������������������������������������������������������������������������������������������������Chlorotocoides Carapace with supraorbital spine (Fig. 6.47 p, q). Pleonite 3 with dorsomedian spine on posterior margin (Fig. 6.47r). Telson not bifurcate (Fig. 6.48t). Scaphocerite with 2–4 lateral teeth (Fig. 6.47q). Pereopods 1–3 with epipods. Pereopod 2 carpus undivided���������������������������������������������������������������������������������������������Thalassocaris Pereopod 2 carpus of 2 (Fig. 6.48p) or 3 segments���������������������������������������������������������������������������������������������������Chlorotocus Pereopod 2 carpus of more than 3 segments (Fig. 6.48n, o, q, r)����������������������������������������������������������������������������������������������4 Carapace without longitudinal carinae except postrostral crest (Fig. 6.47b)�������������������������������������������������������������������������5 Carapace with longitudinal carinae. Integument firm (Fig. 6.47a)����������������������������������������������������������������������������������������17 Rostrum movably connected to carapace (Fig. 6.47n)������������������������������������������������������������������������������������������������ Pantomus Rostrum not movable����������������������������������������������������������������������������������������������������������������������������������������������������������������������6 Cornea narrower than eyestalk (Fig. 6.47h)�����������������������������������������������������������������������������������������������������������������Dorodotes Cornea much wider than eyestalk (Fig. 6.48a)����������������������������������������������������������������������������������������������������������������������������7 Maxilliped 3 with exopod���������������������������������������������������������������������������������������������������������������������������������������������������������������8 Maxilliped 3 without exopod�������������������������������������������������������������������������������������������������������������������������������������������������������10

6 – Caridea – shrimps

161

Fig. 6.47.  Pandalidae. Habitus: a, Heterocarpus sibogae De Man, 1917; b, Plesionika martia (A. Milne-Edwards, 1883). Carapace, rostrum: c, Austropandalus; d, Bitias; e, Chelonika; f, Chlorotocoides; g, Dichelopandalus; h, Dorodotes; i, Heteronika; j, Notopandalus; k, Pandalina; l, Pandalus; m, Procletes; n, Pantomus; o, Pseudopandalus; p, q, Thalassocaris (lateral and dorsal, with scaphocerites). Pleonites 1–6: r, Thalassocaris.

8.

Scaphognathite posterior lobe acutely produced (Fig. 6.48j). Stylocerite broad, rounded. Rostrum with only movable dorsal spines (Fig. 6.47g)���������������������������������������������������������������������������������������������������������������� Dichelopandalus – Scaphognathite posterior lobe broadly rounded (Fig. 6.48i) or truncate (Fig. 6.48h). Stylocerite acute or truncate. Rostrum with all or some dorsal teeth fixed���������������������������������������������������������������������������������������������������������9 9. Pereopod 2 chela not swollen (Fig. 6.48q). Stylocerite as long as first antennular article, acute or blunt (Fig. 6.48e, f)����������������������������������������������������������������������������������������������������������������������������������������������������������������Plesionika – Pereopod 2 chela swollen (Fig. 6.48o). Stylocerite less than half as long as first antennular article, truncate (Fig. 6.48c)��������������������������������������������������������������������������������������������������������������������������������������������������������������������Chelonika 10. Pereopods without epipods��������������������������������������������������������������������������������������������������������������������������������������� Peripandalus – Pereopods 1, 2, at least with epipods (Fig. 6.48w)���������������������������������������������������������������������������������������������������������������������11 11. Pereopods without arthrobranchs�������������������������������������������������������������������������������������������������������������������������������� Pandalina – Pereopods 1–4 with arthrobranchs (Fig. 6.48w)���������������������������������������������������������������������������������������������������������������������� 12 12. Pereopods 1, 2 with epipods. Pereopods 2 subequal��������������������������������������������������������������������������������������������Notopandalus – Pereopods 1–4 with epipods (Fig. 6.48v). Pereopods 2 equal or unequal������������������������������������������������������������������������������13 13. Rostrum short, not reaching end of antennular peduncle; all dorsal teeth movable (Fig. 6.47d). Pereopods 2 equal. Stylocerite short, acute (Fig. 6.48b)�����������������������������������������������������������������������������������������������������������������������Bitias

162

Marine Decapod Crustacea

Fig. 6.48.  Pandalidae. Eye, with ocellus: a, Pandalus. Antenna 1 peduncle, stylocerite: b, Bitias; c, Chelonika; d, Pandalus; e, f, Plesionika; g, Pseudopandalus. Maxilla, scaphognathite (posterior lobe arrowed): h, Austropandalus; i, Chelonika; j, Pandalus. Pereopod 1, carpus, propodus: k, Chlorotocus. Pereopod 1, propodus, dactylus: l, m: Pandalus montagui Leach, 1814 (detail of chela). Pereopod 2, carpus, chela: n, Atlantopandalus; o, Chelonika; p, Chlorotocus; q, Plesionika; r, Procletes. Telson: s, u, Chlorotocoides; t, Thalassocaris. Male pleopod 1: u, Chlorotocoides. Thoracic sternites 6–8 and coxae: v, Chlorotocus. Left branchial cavity showing base of maxilliped 2, coxae of maxilliped 3, pereopods 1–5, one epipod each on mp3–p4, two arthrobranchs on mp3, one each on p1–p4, one pleurobranch each on p1–p5: w, Bitias

– 14. – 15. – 16. – 17. – 18. –

Rostrum long, reaching beyond antennular peduncle (Fig. 6.47c, l, o). Pereopods 2 unequal. Stylocerite short or long, acute or rounded�����������������������������������������������������������������������������������������������������������������������������������������������14 Rostrum dorsal margin with movable spines and fixed teeth (Fig. 6.47c). Scaphognathite posterior lobe truncate (Fig. 6.48h)������������������������������������������������������������������������������������������������������������������������������������������Austropandalus Rostrum dorsal margin with movable spines only (Fig. 6.47l, o). Scaphognathite posterior lobe rounded or acutely produced�����������������������������������������������������������������������������������������������������������������������������������������������������������������������15 Scaphognathite posterior lobe short, rounded (Fig. 6.48i). Stylocerite long, sharp (Fig. 6.48g)��������������� Pseudopandalus Scaphognathite posterior lobe acutely produced (Fig. 6.48j). Stylocerite short, rounded or truncate (Fig. 6.48d)���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������16 Pereopods 2 with right carpus with > 6 segments (Fig. 6.48j). Eye with ocellus (Fig. 6.48a). Protandrous hermaphroditic������������������������������������������������������������������������������������������������������������������������������������������������������������� Pandalus Pereopods 2 with right carpus with 5 segments, first and last longest (Fig. 6.48n). Eye without ocellus. Dioecius������������������������������������������������������������������������������������������������������������������������������������������������������������ Atlantopandalus Carapace with lateral carina bifurcating over posterior half (Fig. 6.47m. Pereopods 2 equal, carpus with 6 segments, first and last longest (Fig. 6.48r)��������������������������������������������������������������������������������������������������������������� Procletes Carapace with 1 or 2 carinae extending at least two-thirds carapace length. Pereopods 2 unequal��������������������������������18 Carapace with 1 lateral carina (Fig. 6.47i). Pleonites without posterodorsal keels���������������������������������������������� Heteronika Carapace with 2 lateral carinae. Pleonites often with posterodorsal keels produced as spine (Fig. 6.47a)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Heterocarpus Atlantopandalus Komai, 1999

Diagnosis. Rostrum long, reaching beyond antennular peduncle, with dorsal teeth movable, except subapical tooth. Eye without ocellus. Stylocerite short, rounded. Maxilla scaphognathite

posterior lobe acutely produced. Pereopods 2 unequal; pereopod 2 left carpus with > 10 segments, right carpus with 5 segments, most proximal occupying half carpus. Maximum cl. 22 mm. Shelf–bathyal (20–2180 m). Arctic, Temperate Northern Atlantic. 1 species (Komai 1999).

6 – Caridea – shrimps

Austropandalus Holthuis, 1952 Diagnosis. Rostrum long, reaching beyond antennular peduncle, with movable and fixed dorsal teeth. Pleonite 3 with dorsomedian spine on posterior margin. Stylocerite short, rounded. Maxilla scaphognathite posterior lobe truncate. Pereopods 2 unequal. Maximum cl. 20 mm. Shelf, slope (24–414 m). Tropical Atlantic, Temperate South America. 1 species (Holthuis 1952).

Bitias Fransen, 1990 Diagnosis. Rostrum short, not reaching end of antennular peduncle, with dorsal teeth movable. Eye without ocellus. Stylocerite about as long as first antennular peduncle article, acute or blunt. Pereopods 2 subequal. Maximum cl. 8 mm. Slope (620–1350 m). Tropical Atlantic, Indo-West Pacific. 2 species (Anker et al. 2014; Fransen 1990).

Chelonika Fransen, 1997 Diagnosis. Rostrum slightly overreaching scaphocerite, with all dorsal teeth fixed. Stylocerite less than half as long as first antennular peduncle article, truncate. Maxilla scaphognathite posterior lobe broadly rounded. Pereopods 2 subequal; pereopod 2 palm swollen, twice as wide as carpus. Maxilliped 3 with exopod. Maximum cl. 7.3 mm. Shelf (144–155 m). Central Indo-Pacific. 1 species (Fransen 1997).

Chlorotocoides Kemp, 1925 Diagnosis. Rostrum slightly overreaching scaphocerite, with all dorsal teeth fixed. Pleonite 6 with dorsomedian spine on posterior margin. Telson bifurcate. Stylocerite overreaching second antennular article. Pereopod 2 carpus of 2 segments. Male pleopod 1 endopod enlarged, convoluted, spinulose. Epipods on pereopods 1–4. Maximum cl. 9 mm (Pl. 12a). Subtidal, shelf (9–32 m). Western and Central Indo-Pacific. 1 species (Chace 1985: rediagnosis; De Man 1902: figures).

Chlorotocus A. Milne-Edwards, 1882 Diagnosis. Rostrum short, not reaching end of antennular peduncle, with all dorsal teeth fixed. Pereopods 2 subequal; pereopod 2 carpus of 2 segments, or carpus of 3 segments. Maxilliped 3 with exopod (Pl. 12b). Subtidal–slope (3–597 m). Temperate Northern Atlantic, Western and Central Indo-Pacific, Temperate Australasia. 2 species (Chace 1985: rediagnosis).

Dichelopandalus Caullery, 1896 pink shrimp Diagnosis. Rostrum long, reaching beyond antennular peduncle, with only movable dorsal spines. Stylocerite broad, rounded.

163

Maxilla 2, scaphognathite posterior lobe acutely produced. Maxilliped 3 with exopod. Maximum cl. 30 mm. Shelf, slope (20–850 m). Temperate Northern and Tropical Atlantic (rare). 2 species (Al-Adhub and Bowers 1977: biology; Calman 1899: figures of limbs; Squires 1990: figures). One species is fished commercially.

Dorodotes Bate, 1888 Diagnosis. Rostrum short, not reaching end of antennular peduncle, with movable and fixed dorsal teeth. Cornea narrower than eyestalk. Pereopods 2 subequal; pereopod 2 carpus multiarticulate (8–10 segments). Maxilliped 3 with exopod. Maximum cl. 25 mm (Pl. 12c). Bathyal, abyssal (1920–4114 m). GOODS bathyal provinces: Indian, W Pacific. 1 species (Chace 1985: rediagnosis).

Heterocarpus A. Milne-Edwards, 1881 Diagnosis. Rostrum long, reaching beyond antennular peduncle, with all dorsal teeth fixed. Carapace with 2 lateral carinae, sometime short third more dorsal. Pleonites often with posterodorsal keels produced as spine. Pereopods 2 unequal; pereopod 2 right carpus with 7–12 segments, right carpus with 18–25 segments. Maximum cl. 37 mm (Pl. 12f). Shelf, slope (73–2834 m). Cosmopolitan. 30 species (Crosnier 1988: key to 25 species and subspecies; Crosnier 1999: supplement to key).

Heteronika Hendrickx, 2019 Diagnosis. Rostrum long, reaching beyond antennular peduncle, with all dorsal teeth fixed. Carapace with 1 lateral carina extending at least two-thirds carapace length. Pleonites without posterodorsal keels. Pereopods 2 unequal; pereopod 2 right carpus of 5 segments, first and last longest, left carpus with 17–18 segments. Maximum cl. 31 mm. Slope (750–850 m). Tropical Atlantic, Eastern Indo-Pacific (possible). 2 species (Hendrickx 2019).

Notopandalus Yaldwyn, 1960 Diagnosis. Rostrum long, reaching beyond antennular peduncle, with dorsal teeth movable, except subapical tooth. Pleonite 3 with dorsomedian spine on posterior margin. Stylocerite half as long as first antennular peduncle article, acute. Pereopods 2 subequal. Epipods on pereopods 1, 2. Maximum cl. 13 mm. Slope, mesopelagic (275–600 m). Temperate New Zealand. 1 species (Webber et al. 1990; Yaldwyn 1960).

Pandalina Calman, 1899 Diagnosis. Rostrum short, not reaching end of antennular peduncle, with movable and fixed dorsal teeth. Pleonite 3 with dorsomedian spine on posterior margin. Stylocerite less than half as long as first antennular peduncle article, truncate. Pereopods 2

164

Marine Decapod Crustacea

unequal. Arthrobranchs absent from pereopods 1–4. Maximum cl. 7 mm. Shelf– bathyal (50–1068 m). Temperate Northern and Tropical Atlantic, Western Indo-Pacific, Temperate Southern Africa. GOODS bathyal provinces: Nazca Plate (doubtful). 5 species (possibly 4) (Komai and Chan 2010: rediagnosis, discussion of species).

Pandalus Leach, 1814 Diagnosis. Protandric hermaphroditic. Rostrum long, reaching beyond antennular peduncle, with dorsal teeth movable, except subapical tooth. Stylocerite short, rounded. Maxilla 2, scaphognathite posterior lobe acutely produced. Pereopods 2 unequal; pereopod 2 left carpus with > 10 segments, right carpus with > 6 segments. Maximum cl. 48 mm (Pl. 12e). Intertidal–bathyal (0–1380 m). Temperate Northern Atlantic, Temperate Northern Pacific, Tropical Eastern Pacific. 43 species (Komai 1999: key to species). The genus contains some large shrimps of current or potential economic importance (Holthuis 1980; Park et al. 2012).

1997). Plesionika has not been treated taxonomically world-wide but local studies are available (e.g. Chan and Crosnier 1997; Hanamura and Evans 1996; Li and Davie 2006).

Procletes Bate, 1888 Diagnosis. Rostrum long, reaching beyond antennular peduncle, with all dorsal teeth fixed. Carapace with lateral carina bifurcating over posterior half. Pleonite 3–5 each with dorsomedian spine on posterior margin. Pereopods 2 subequal; pereopod 2 carpus of 6 segments, first and last longest. Maximum cl. 11 mm (Pl. 12d). Subtidal–slope (14–393 m). Western and Central Indo-Pacific. 1 species (De Man 1920).

Pseudopandalus Crosnier, 1997 Diagnosis. Rostrum long, reaching beyond antennular peduncle, with dorsal teeth movable, except subapical tooth. Pleonite 3 with dorsal truncated crest at midpoint. Stylocerite overreaching second antennular article. Pereopods 2 unequal. Maximum cl. 25 mm. Slope (420–570 m). Central Indo-Pacific. 11 species (Crosnier 1997).

Pantomus A. Milne-Edwards, 1883 Diagnosis. Rostrum movably connected to carapace, long, reaching beyond antennular peduncle, with dorsal teeth movable, except subapical tooth, 2 movable teeth on carapace. Pereopods 2 unequal; pereopod 2 carpus multiarticulate (8, 18 articles). Maxilliped 3 with exopod. Maximum cl. 7.3 mm. Shelf, slope (110–474 m). Tropical Atlantic, Eastern IndoPacific. 2 species (Chace 1937).

Peripandalus De Man, 1917 Diagnosis. Rostrum long, reaching beyond antennular peduncle, with all dorsal teeth fixed. Pereopods 2 unequal. Epipods absent from pereopods 1–4. Shelf. Central Indo-Pacific. 1 species (Holthuis 1993: only figure).

Plesionika Bate, 1888 Diagnosis. Rostrum long, reaching beyond antennular peduncle or slightly overreaching scaphocerite, with all or some dorsal teeth fixed. Pleonite 3 with dorsomedian spine on posterior margin, or pleonites without posterodorsal keels. Stylocerite about as long as first antennular peduncle article, acute or blunt. Maxilla 2, scaphognathite posterior lobe broadly rounded, or truncate. Pereopods 2 subequal. Maxilliped 3 with exopod. Maximum cl. 33 mm (Pl. 12g). Subtidal–abyssal (14–3600 m). Cosmopolitan (except Temperate S America). 96 species, divided into species-groups (Chan 2004; Chan and Chuang 2002; Chan and Crosnier 1991). The genus is common and diverse in the catches of deep-sea commercial trawlers in Taiwan and elsewhere in the NW Pacific where some species have been studied (e.g. Chilari et al. 2005; Ohtomi

Thalassocaris Stimpson, 1860 Diagnosis. Rostrum slightly overreaching scaphocerite, with all dorsal teeth fixed. Carapace with supraorbital spine. Pleonite 3 with dorsomedian spine on posterior margin. Eye without ocellus. Pereopod 2 carpus undivided. Male pleopod 1 endopod enlarged, convoluted, spinulose. Epipods on pereopods 1–3. Maximum cl. 7 mm (Pl. 12h). Subtidal–abyssal, pelagic (0–200 m over depths of 13–4000 m). Indo-West Pacific, Temperate Southern Africa. 3 species (Chace 1985: rediagnosis, key to species). References Al-Adhub AHY, Bowers AB (1977) Growth and breeding of Dichelopandalus bonnieri in Isle of Man waters. Journal of the Marine Biological Association of the United Kingdom 57, 229–238. doi:10.1017/ S0025315400021378 Anker A, Pachelle PPG, Tavares M (2014) Two new species and two new records of deep-water caridean shrimps from Brazil (Decapoda: Pandalidae, Palaemonidae, Crangonidae). Zootaxa 3815, 263–278. doi:10.11646/zootaxa.3815.2.6 Calman WT (1899) On the British Pandalidae. Annals and Magazine of Natural History (ser. 7) 3, 27–39, pls 1–4. Chace FA (1937) The Templeton Crocker Expedition. VII. Caridean decapod Crustacea from the Gulf of California and the west coast of Lower California. Zoologica 22, 109–138. Chace FA (1985) The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 3: families Thalassocarididae and Pandalidae. Smithsonian Contributions to Zoology 411, 1–143. doi:10.5479/si.00810282.411 Chan T-Y (2004) The “Plesionika rostricrescentis (Bate, 1888)” and “P. lophotes Chace, 1985” species groups of Plesionika Bate, 1888, with descriptions of five new species (Crustacea: Decapoda: Pandalidae). In: Marshall BA, Richer de Forges B (Eds), Tropical Deep-Sea Benthos, Vol. 23. Mémoires du Muséum National d’Histoire Naturelle, Paris 191, 293–318.

6 – Caridea – shrimps

Chan T-Y, Chuang S-C (2002) A new shrimp species of Plesionika Bate, 1888 with high basal rostral crest (Crustacea: Decapoda: Pandalidae) from Taiwan. Proceedings of the Biological Society of Washington 115, 611–615. Chan T-Y, Crosnier A (1991) Crustacea Decapoda: Studies of the Plesionika narval (Fabricius, 1787) group (Pandalidae) with descriptions of six new species. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 9. Mémoires du Muséum National d’Histoire Naturelle, Paris 152, 413–461. Chan T-Y, Crosnier A (1997) Crustacea Decapoda: Deep-sea shrimps of the genus Plesionika Bate, 1888 (Pandalidae) from French Polynesia, with descriptions of five new species. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 18. Mémoires du Muséum National d’Histoire Naturelle, Paris 176, 187–234. Chilari A, Thessalou-Legaki M, Petrakis G (2005) Population structure and reproduction of the deep-water shrimp Plesionika martia (Decapoda: Pandalidae) from the Eastern Ionian Sea. Journal of Crustacean Biology 25, 233–241. doi:10.1651/C-2513 Crosnier A (1988) Sur les Heterocarpus (Crustacea, Decapoda, Pandalidae) du sud-ouest de l’océan Indien. Remarques sur d’autres espèces ouestpacifiques du genre et description de quatre taxa nouveaux. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 10, 57–103. Crosnier A (1997) Crustacea Decapoda: Pseudopandalus curvirostris, genre et espèce nouveaux (Pandalidae) de Nouvelle-Calédonie. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 18. Mémoires du Muséum National d’Histoire Naturelle, Paris 176, 169–176. Crosnier A (1999) Un Heterocarpus nouveau (Crustacea, Decapoda, Pandalidae) du Pacifique Sud-Ouest. Zoosystema 21, 345–351. De Man JG (1902) Die von herrn Professor Kükenthal im Indischen Arrchipel gesammelten Dekapoden und Stomatopoden. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 25, 465–929. De Man JG (1920) Pasiphaeidae, Stylodactylidae, Hoplophoridae, Nematocarcinidae, Thalassocaridae, Pandalidae, Psalidopodidae, Gnathophyllidae, Processidae, Glyphocranconidae and Crangonidae. Siboga-Expéditie 39, 1–318, pls 1–25. Fransen CHJM (1990) Bitias stocki, a new genus and new species of pandalid shrimp (Crustacea, Decopoda, Caridea) in the eastern Atlantic Ocean. Beaufortia 41, 67–73. Fransen CHJM (1997) Crustacea Decapoda: Chelonika macrochela, a new genus and new species of pandalid shrimp (Caridea) from New Caledonian waters. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 18. Mémoires du Muséum National d’Histoire Naturelle, Paris 176, 177–185. Hanamura Y, Evans DR (1996) Deepwater caridean shrimps of the families Nematocarcinidae, Stylodactylidae, Pandalidae and Crangonidae (Crustacea: Decapoda) from Western Australia. Bulletin of the Nansei National Fisheries Research Institute 29, 1–18. Hendrickx M (2019) Redescription of the rare shrimp Heterocarpus nesisi (Burukovsky, 1986 (Crustacea: Caridea: Pandalidae) rediscovered off western Mexico, with the proposal of a new genus. Zootaxa 4565, 49–60. doi:10.11646/zootaxa.4565.1.3 Holthuis LB (1952) Reports of the Lund Univeristy Chile Expedition 1948–49, 5. The Crustacea Decapoda Macrura of Chile. Lunds Universitets Årsskrift N.F. Avd. 2(47), 1–109. Holthuis LB (1980) FAO species catalogue. Vol. 1. Shrimps and prawns of the world. An annotated catalogue of species of interest to fisheries. FAO Fisheries Synopsis 125, 1–261. Holthuis LB (1993) The recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda) with an appendix on the Order Amphionidacea. revised edn. Nationaal Natuuristorisch Museum, Leiden. Komai T (1999) A revision of the genus Pandalus (Crustacea: Decapoda: Caridea: Pandalidae. Journal of Natural History 33, 1265–1372. doi:10.1080/002229399299914

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Komai T, Chan T-Y (2010) Two new pandalid shrimps and the discovery of the second specimen of the rare hippolytid shrimp Leontocaris bulga Taylor & Poore, 1998 (Crustacea, Decapoda) from the Mozambique MAINBAZA Cruise. Zoosystema 32, 625–641. doi:10.5252/ z2010n4a6 Li X, Davie PJF (2006) An account of the pandaloid shrimps (Crustacea: Decapoda: Caridea) in the collections of the Queensland Museum. Memoirs of the Queensland Museum 52, 151–170. Ohtomi J (1997) Reproductive biology and growth of the deep-water pandalid shrimp Plesionika semilaevis (Decapoda: Caridea). Journal of Crustacean Biology 17, 81–89. doi:10.2307/1549465 Park H-M, Oh C-W, Sohn MH (2012) Distribution and reproductive aspects of the pandalid shrimp, Pandalus eous, in the deep sea of the East Sea, Korea. Animal Cells and Systems 16, 77–84. doi:10.1080/1976 8354.2011.620173 Squires HJ (1990) Decapod Crustacea of the Atlantic coast of Canada. Canadian Bulletin of Fisheries and Aquatic Sciences 221, 1–532. Webber WR, Fenaughty CM, Clark MR (1990) A guide to some common offshore shrimp and prawn species of New Zealand. New Zealand Fisheries Occasional Publication 6, 1–42. Yaldwyn JC (1960) Biological results of the Chatham Islands 1954 Expedition, Part I—Crustacea Decapoda Natantia from the Chatham rise: a deep water bottom fauna from New Zealand. New Zealand Department of Scientific and Industrial Research Bulletin 139, 13–53.

Superfamily Pasiphaeoidea Dana, 1852 The only family, Pasiphaeidae, is distributed throughout the world’s oceans including polar regions. The shrimps are sufficiently abundant in some parts of Asia to be fished commercially. Most are mesopelagic or bathypelagic and migrate vertically, feeding on benthic animals during the day and plankton at night (Cartes 1993). Some are bioluminescent (Herring 1985). Pasiphaeids are instantly recognised by their long slender pectinate fingers on the first two pairs of pereopods. Molecular analysis has suggested that the family Pasiphaeidae is not monophyletic, with Psathyrocaris more closely related to other carideans (Liao et  al. 2017). These authors reviewed the morphology of the genera and identified morphological characters shared by the genera, or not, and others unique to Psathyrocaris. The key is based on those of Crosnier (1988) and Holthuis (1993) with the addition of a seventh genus. Diagnosis. Rostrum small, obsolete, foten upright. Mandible molar absent (Fig.  6.49h, i). Maxilliped 3 with fewer than 7 articles. Pereopods 1, 2 similar, with long slender pectinate fingers (Fig. 6.49l). Pereopod 2 carpus entire. Exopod present on maxilliped 3, pereopods 1–5.

Pasiphaeidae Dana, 1852 Figure 6.49, Plate 12i–m Implicit generic attributes. Rostrum directed forwards. Carapace without dorsal teeth. Mandibular palp present in adults.

166

Marine Decapod Crustacea

Key to genera of Pasiphaeidae 1. – 2. – 3. – 4. – 5. – 6. –

Mandible without palp (Fig. 6.49h). Rostrum an erect postfrontal spine or tooth (Fig. 6.49b, c)���������������������������������������2 Mandible with palp in adult (Fig. 6.49i). Rostrum directed forwards (Fig. 6.49d–g)������������������������������������������������������������3 Telson posterior margin with 4 or more pairs of spiniform setae (Fig. 6.49k). Arthrobranchs present. Pereopod 5 with pleurobranch����������������������������������������������������������������������������������������������������������������������������������Pasiphaea Telson posterior margin with 3 pairs of spiniform setae (Fig. 6.49j). Arthrobranchs absent. Pereopod 5 without pleurobranch���������������������������������������������������������������������������������������������������������������������������������� Alainopasiphaea Pereopod 4 distinctly shorter than both pereopods 3, 5������������������������������������������������������������������������������������������������������������4 Pereopod 4 longer than pereopod 5 (Fig. 6.49a)��������������������������������������������������������������������������������������������������������������������������6 Carapace without antennal and branchiostegal spines, usually without dorsal teeth (Fig. 6.49f)����������������Parapasiphae Carapace with antennal and branchiostegal spines, with dorsal teeth (Fig. 6.49d, e)�����������������������������������������������������������5 Maxilliped 3 with 1 arthrobranch������������������������������������������������������������������������������������������������������������������������������������ Glyphus Maxilliped 3 with 2 arthrobranchs���������������������������������������������������������������������������������������������������������������������������� Eupasiphae Pereopods 3, 4 slender, about as long as pereopod 1. Rostrum with dorsal teeth (Fig. 6.49g)�����������������������Psathyrocaris Pereopod 3 longer than pereopod 4, both shorter than pereopod 1. Rostrum without dorsal teeth (Fig. 6.49a)�������������������������������������������������������������������������������������������������������������������������������������������������������������������Leptochela Alainopasiphaea Hayashi, 1999

Diagnosis. Rostrum an erect postfrontal spine. Carapace with branchiostegal spine. Telson posterior margin with 3 pairs of spiniform setae. Mandibular palp absent in adults. Pereopod 4 distinctly

shorter than both pereopods 3, 5. Arthrobranchs absent; pereopod 5 without pleurobranch. Maximum cl. 12 mm (Pl. 12k). Shelf, slope (32–797 m). Western and Central Indo-Pacific, Temperate Australasia. 2 species (Hanamura 1989: as Pasiphaea australis; Hayashi 1999).

Fig. 6.49.  Pasiphaeidae. Habitus: a, Leptochela sydniensis Dakin & Colefax, 1940; b, Pasiphaea barnardi Yaldwyn, 1971. Carapace: c, Alainopasiphaea; d, Eupasiphae; e, Glyphus; f, Parapasiphae; g, Psathyrocaris. Mandible: h, Alainopasiphaea; i, Pasiphaea. Telson, with posterior margin: j, Alainopasiphaea; k, Pasiphaea. Pereopod 1, chela: l, Pasiphaea.

6 – Caridea – shrimps

167

Eupasiphae Wood-Mason, 1893

Psathyrocaris Wood-Mason, 1893

Diagnosis. Rostrum with dorsal teeth. Carapace with antennal and branchiostegal spines. Telson posterior margin with 4 or 5 pairs of spiniform setae. Maxilliped 3 with 2 arthrobranchs. Pereopod 4 distinctly shorter than both pereopods 3, 5. Maximum cl. 46 mm. Slope–abyssal (790–4740 m). Temperate Northern Atlantic, Temperate Northern Pacific, Tropical Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate Australasia. 5 species (Crosnier 1988: key to 3 species; Hanamura and Evans 1994: compared 2 species; Rodrigues and Cardoso 2018: fifth species).

Diagnosis. Rostrum with small dorsal fixed or movable teeth. Carapace without antennal and branchiostegal spines. Telson posterior margin with 2–4 pairs of spiniform setae. Pereopods 3, 4 slender, about as long as pereopod 1. Pleopodal exopods long, narrow, endopods shorter. Maximum cl. 32 mm. Slope (315–2000 m depth). Cosmopolitan. 5 species (Lin and Chan 2001).

Glyphus Filhol, 1884 Diagnosis. Rostrum without dorsal teeth. Carapace with antennal and branchiostegal spines. Telson posterior margin with 4 pairs of spiniform setae. Maxilliped 3 with 1 arthrobranch. Pereopod 4 distinctly shorter than both pereopods 3, 5. Maximum cl. 61 mm (Pl. 12m). Slope (800–1160 m). Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific, Temperate South America. 1 species (Crosnier and Forest 1973: redescription).

Leptochela Stimpson, 1860 Diagnosis. Rostrum without dorsal teeth. Carapace without antennal and branchiostegal spines. Telson posterior margin with 5 pairs of spiniform setae. Pereopod 3 longer than pereopod 4, both shorter than pereopod 1. Pleopodal exopods and endopods short, subequal. Maximum cl. 13 mm (Pl. 12i). Subtidal–slope (6–525 m). Temperate Northern Atlantic (Mediterranean invader), Temperate Northern Pacific, Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific, Temperate Australasia. 16 species (Chace 1976: key to 12 species; Hayashi 1995: recognised 2 species groups).

Parapasiphae Smith, 1884 Diagnosis. Rostrum without dorsal teeth. Carapace without antennal and branchiostegal spines; usually with dorsal teeth. Telson posterior margin with c. 8 pairs of spiniform setae. Pereopod 4 distinctly shorter than both pereopods 3, 5. Maximum cl. 21 mm. Slope–abyssal (944–5397 m). Cosmopolitan including polar oceans. 4 species (Hanamura 1989: 2 species compared; Wasmer 2005: key to species).

Pasiphaea Savigny, 1816 Diagnosis. Rostrum an erect postfrontal spine, or tooth-like. Carapace with branchiostegal spine. Telson posterior margin with 4 pairs of spiniform setae. Mandibular palp absent in adults. Arthrobranchs present at least on pereopods 1–3; pereopod 5 with pleurobranch. Maximum cl. 37 mm (Pl. 12j). Shelf, slope, bathyal, mesopelagic (100–2452 m). Cosmopolitan including polar oceans. 69 species (Hayashi 1999: key to 9 spp. of P. sivado group; Hayashi 2004: key to 22 spp. of P. cristata species group; Hayashi 2006: key to 17 spp. of P. alcocki species group; Komai et al. 2012: compared 10 species).

References Cartes JE (1993) Feeding habits of pasiphaeid shrimps close to the bottom on the Western Mediterranean slope. Marine Biology 117, 459–468. Chace FA (1976) Shrimps of the pasiphaeid genus Leptochela with descriptions of three new species (Crustacea: Decapoda: Caridea). Smithsonian Contributions to Zoology 222, 1–51. doi:10.5479/si.00810282.222 Crosnier A (1988) Les Eupasiphae (Crustacea Decapoda Pasiphaeidae) du sud-ouest de l’océan Indien. Description d’E. paucidentata sp. nov. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 10, 785–797. Crosnier A, Forest J (1973) Les crevettes profondes de l’Atlantique oriental tropical. Faune Tropicale 19, 1–409. Hanamura Y (1989) Deep-sea shrimps (Crustacea: Decapoda) collected by the R.V. “Soela” from southern Australia. Nihon Seibutsu Chiri Gakkai Kaiho 44, 51–69. Hanamura Y, Evans DR (1994) Deepwater caridean shrimps of the families Oplophoridae and Pasiphaeidae (Crustacea: Decapoda) from Western Australia, with an appendix on a lophogastridan mysid (Mysidacea). Crustacean Research 23, 46–60. doi:10.18353/crustacea.23.0_46 Hayashi K-I (1995) Brief revision of the genus Leptochela, with description of two new species (Crustacea, Decapoda, Pasiphaeidae). In Les fonds meubles des lagons de Nouvelle-Calédonie (Sédimetnologie, benthos). (Ed. Richer de Forges B) pp. 83–99. ORSTOM, Paris. Hayashi K-I (1999) Crustacea Decapoda: Revision of Pasiphaea sivado (Risso, 1816) and related species, with descriptions of one new genus and five new species (Pasiphaeidae). In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 20. Mémoires du Muséum National d’Histoire Naturelle, Paris 180, 267–302. Hayashi K-I (2004) Revision of the Pasiphaea cristata Bate, 1888 species group of Pasiphaea Savigny, 1816, with descriptions of four new species and referral of P. australis to Alainopasiphaea Hayashi, 1999 (Crustacea: Decapoda: Pasphaeidae). In: Marshall BA, Richer de Forges B (Eds), Tropical Deep-Sea Benthos, Vol. 23. Mémoires du Muséum National d’Histoire Naturelle, Paris 191, 319–373. Hayashi K-I (2006) Revision of the Pasiphaea alcocki species group (Crustacea, Decapoda, Pasiphaeldae). In: Richer de Forges B, Justine J-L (Eds), Tropical Deep-Sea Benthos, Vol. 24. Mémoires du Muséum National d’Histoire Naturelle, Paris 193, 193–241. Herring PJ (1985) Bioluminescence in the Crustacea. Journal of Crustacean Biology 5, 557–573. doi:10.2307/1548235 Holthuis LB (1993) The Recent genera of the caridean and stenopodidean shrimps (Crustacea, Decapoda) with an appendix on the Order Amphionidacea. revised edn. Nationaal Natuuristorisch Museum, Leiden. Komai T, Lin WC, Chan T-Y (2012) Bathypelagic shrimp of the genus Pasiphaea (Decapoda: Caridea: Pasiphaeidae) from waters around Taiwan, with description of four new species. Journal of Crustacean Biology 32, 295–325. doi:10.1163/193724011X615550 Liao Y, De Grave S, Ho TW, Ip BHY, Tsang LM, et al. (2017) Molecular phylogeny of Pasiphaeidae (Crustacea, Decapoda, Caridea) reveals systematic incongruence of the current classification. Molecular Phylogenetics and Evolution 115, 171–180. doi:10.1016/j.ympev.2017.07.021

168

Marine Decapod Crustacea

Lin C-W, Chan T-Y (2001) First record of the deep-sea caridean shrimp genus Psathyrocaris Wood-Mason and Alcock, 1893 (Decapoda: Pasiphaeidae) from Taiwan. Raffles Bulletin of Zoology 49, 51–56. Rodrigues TGA, Cardoso IA (2018) A new species of Eupasiphae WoodMason & Alcock, 1893 (Crustacea, Decapoda, Pasiphaeidae) from the southwestern Atlantic. Zootaxa 4444, 189–194. doi:10.11646/ zootaxa.4444.2.7 Wasmer RA (2005) A remarkable new species of the pelagic shrimp genus Parapasiphae Smith, 1884 (Crustacea: Decapoda: Pasiphaeidae) with double eyes. Proceedings of the Biological Society of Washington 118, 165–175. doi:10.2988/0006-324X(2005)118[165:ARNSOT]2.0.CO;2

Superfamily Physetocaridoidea Chace, 1940 The superfamily comprises a single species. Diagnosis. Rostrum small, inflated, merging with carapace (Fig. 6.50a). Mandible without incisor or palp. Pereopods 1, 2 dissimilar, second with fixed finger curving subrectangularly around short dactylus. Pereopod 2 carpus multiarticulate (Fig.  6.50b, c). Exopod absent from pereopods 1–5.

Physetocarididae Chace, 1940 Figure 6.50 Physetocaris Chace, 1940 Bathypelagic. N and mid-Atlantic, North Pacific, W Tasman Sea, SE Pacific. Maximum cl. 16 mm (Chace 1940). The only species P. microphthalma Chace, 1940 is immediately recognised by the inflated rostrum which dominates the carapace (Foxton and Herring 1970). The species is oceanic, probably a vertical migrator, rarely at the surface, but more common between 900 and 1720 m depth (Wasmer 1985). References Chace FA (1940) Plankton of the Bermuda Oceanographic Expeditions. IX. The bathypelagic caridean Crustacea. Zoologica 25, 117–209. Foxton P, Herring PJ (1970) Recent records of Physetocaris microphthalma Chace with notes on the male and description of the early larvae (Decapoda, Caridea). Crustaceana 18, 93–104. doi:10.1163/​ 156854070X00103 Wasmer RA (1985) New records for Physetocaris microphthalma Chace (Decapoda, Caridea, Physetocarididae) from the South Pacific. Crustaceana 49, 315–318. doi:10.1163/156854085X00648

Fig. 6.50.  Physetocarididae. Physetocaris microphthalma Chace, 1940. a, habitus; b, c, pereopods 1, 2.

Superfamily Processoidea Ortmann, 1896 Processidae is the only family. Processid shrimps range from intertidal rockpools to shelf environments (Chace 1997). Some are mesopelagic but most live in shallow water where they are nocturnal, associated with seagrass and algae (Aguzzi et al. 2008; Manning and Chace 1971). Diagnosis. Rostrum short, with 1 or 2 small terminal teeth (Fig.  6.51b, e). Maxilliped 1 exopod abutting endite, displacing palp (Fig. 6.51k). Pereopods 1, 2 dissimilar; right pereopod 1 chelate, left simple, ending in claw-like dactylus (Fig.  6.51l–m) (both chelate in Ambidexter). Pereopod 2 carpus multiarticulate. Exopod usually present on maxilliped 3, pereopods 2–5 (rarely 1–5).

Processidae Ortmann, 1896 Figure 6.51 Implicit generic attributes. Rostrum narrowly triangular in dorsal review. Pleonite 3 without posterodorsal lobe. Pleonite 6 with pair of posterolateral rounded lobes. Telson dorsolateral spiniform setae distinct. Mandibular molar longer than wide. Pereopods 1 asymmetrical, right chelate, left with simple dactylus. Carpus with numerous segments. Pereopod 1 without exopod.

Key to genera of Processidae 1. –

Rostrum broad, subequilaterally triangular in dorsal view, reaching middle of cornea (Fig. 6.51c). Telson dorsolateral spiniform setae minute or absent (Fig. 6.51g)���������������������������������������������������������������������������Hayashidonus Rostrum usually longer than wide in dorsal view, short or overreaching cornea (Fig. 6.51a, b, d, f). Telson dorsolateral spiniform setae distinct (Fig. 6.51h)�������������������������������������������������������������������������������������������������������������������2

6 – Caridea – shrimps

169

Fig. 6.51.  Processidae. a, Processa australiensis Baker, 1907. Carapace, pleon: b, Maryprocessa pippinae (Wicksten & Méndez, 1985). Rostrum, eyes: c, Hayashidonus japonicus (De Haan, 1844); d, e, Processa longirostris Hayashi, 1975; f, P. macrognatha Hayashi, 1975; Telson: g, Hayashidonus; h, Processa. Mandible: i, Clytomanningus j, Processa. Maxilliped 1: k, Processa. Pereopods 1: l, m, Processa (right, with chela); n, Processa (left); o, Ambidexter. Pereopods 2: p, Clytomanningus; q, r, Processa (right, left).

2. – 3. – 4. – 5. –

Mandible massive, as long as scaphocerite, molar wider than long (Fig. 6.51i). Pereopods 2 subequal, carpus with 6 segments (Fig. 6.51p)��������������������������������������������������������������������������������������������������������������Clytomanningus Mandible not massive, molar longer than wide (Fig. 6.51j). Pereopods 2 unequal, carpus of shorter member with numerous segments (Fig. 6.51q, r)�����������������������������������������������������������������������������������������������������������������������������������3 Pereopod 1 with exopod���������������������������������������������������������������������������������������������������������������������������������������������������� Nikoides Pereopod 1 without exopod������������������������������������������������������������������������������������������������������������������������������������������������������������4 Pereopods 1 symmetrical, chelate (Fig. 6.51o)����������������������������������������������������������������������������������������������������������Ambidexter Pereopods 1 asymmetrical, one chelate, other with simple dactylus (Fig. 6.51l–n)���������������������������������������������������������������5 Rostrum rarely overreaching cornea (Fig. 6.51d–f). Pleonite 3 without posterodorsal lobe overlapping pleonite 4 (Fig. 6.51a)�����������������������������������������������������������������������������������������������������������������������������������������������������Processa Rostrum overreaching cornea. Pleonite 3 with posterodorsal lobe overlapping pleonite 4 (Fig. 6.51b)������ Maryprocessa Ambidexter Manning & Chace, 1971

Hayashidonus Chace, 1997

Diagnosis. Pereopods 1 symmetrical, both chelate. Pereopods 2 unequal. Maximum cl. 6.7 mm. Intertidal, subtidal (0–6 m). Temperate Northern and Tropical Atlantic, Tropical Eastern Pacific. 3 species (Abele 1972: key to species).

Diagnosis. Rostrum broad, subequilaterally triangular in dorsal review; reaching base of cornea. Telson dorsolateral spiniform setae minute or absent. Pereopods 2 unequal. Maximum cl. 16 mm. Shelf (27–150 m). Western and Central Indo-Pacific. 1 species (Chace 1997; Hayashi 1975).

Clytomanningus Chace, 1997

Maryprocessa Hendrickx, 2012

Diagnosis. Mandibles massive, as long as scaphocerite, molar wider than long. Pereopods 2 subequal; carpus with 6 segments. Maximum cl. 3 mm. Intertidal, subtidal (0–15 m). Western and Central IndoPacific. 2 species (Chace 1997).

Diagnosis. Rostrum laterally compressed; overreaching cornea. Pleonite 3 with posterodorsal lobe. Pleonite 6 with pair of posterolateral spines. Pereopods 2 subequal. Maximum cl. 8.6 mm. Mesopelagic (265–1165 m). Gulf of California. 1 species (Hendrickx 2012; Wicksten and Méndez 1985).

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Marine Decapod Crustacea

Nikoides Paul’son, 1875 Diagnosis. Pereopods 2 unequal; carpus with numerous segments. Pereopod 1 with exopod. Maximum cl. 13.6 mm. Intertidal–shelf (0–164 m). Tropical Atlantic, Indo-West Pacific. 11 species (Komai and Hirabayashi 2021: key to species).

Processa Leach, 1815 Diagnosis. Pereopods 2 unequal. Maximum cl. 11 mm. Intertidal–slope (0–619 m). Cosmopolitan except polar seas. 51 species (Noël 1986: key to species).

Psalidopus is the only genus in the superfamily. Its species are commonly referred to as scissor-foot shrimps because of the articulation of both of the fingers of the first cheliped. Besides this, they are easily recognised by the strong spines on the rostrum, along the midline and dorsolateral carinae of the carapace and pleon giving a strongly ‘prickly’ appearance to the whole animal. Two species have been described in detail (Chace and Holthuis 1978). All three species were compared by Toriyama and Horikawa (1993). Psalidopus Wood-Mason, 1892

References Abele LG (1972) A review of the genus Ambidexter (Crustacea: Decapoda; Processidae) in Panama. Bulletin of Marine Science 22, 365–380. Aguzzi J, Company JB, García JA (2008) The circadian behavioural regulation of the shrimp, Processa canaliculata Leach, 1815 (Decapoda, Processidae) in relation to depth, ontogeny, and the reproductive cycle. Crustaceana 81, 1301–1316. doi:10.1163/156854008X369492 Chace FA (1997) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907–1910, Part 7: Families Atyidae, Eugonatonotidae, Rhynchocinetidae, Bathypalaemonellidae, Processidae, and Hippolytidae. Smithsonian Contributions to Zoology 587, 1–106. Hayashi K-I (1975) The Indo-West Pacific Processidae (Crustacea, Decapoda, Caridea). Suisan Daigakko Kenkyu Hokoku 24, 47–145. Hendrickx ME (2012) Distribution and abundance of the pelagic processid, Processa pippinae Wicksten & Méndez, 1985 (Decapoda, Caridea, Processidae), collected during the Talud XIV Cruise in the Gulf of California, Mexico, and description of a new genus. Crustaceana 85, 463–472. doi:10.1163/156854012X623773 Komai T, Hirabayashi I (2021) A new species of the processid shrimp genus Nikoides Paul’son, 1875 (Decapoda: Caridea) from Japan. Zootaxa 5057, 114–126. doi:10.11646/zootaxa.5057.1.7 Manning RB, Chace FA (1971) Shrimps of the family Processidae from the northwestern Atlantic Ocean (Crustacea: Decapoda: Caridea). Smithsonian Contributions to Zoology 89, 1–41. doi:10.5479/si.00810282.89 Noël P (1986) Crustacés Décapodes: Processidae de l’indo-ouest-Pacifique. In: Forest J (Ed.), Résultats des Campagnes MUSORSTOM I & II, Philippines, vol. 2. Mémoires du Muséum National d’Histoire Naturelle, Paris 133, 261–301. Wicksten MK, Méndez MG (1985) Processa pippinae, a new species of deep-sea shrimp from the Gulf of California (Decapoda, Caridea). Crustaceana 49, 16–21. doi:10.1163/156854085X00152

Maximum cl. 32 mm (Pl. 11j). Slope, bathyal (412–2881 m). Tropical Atlantic, Western and Central Indo-Pacific. GOODS bathyal province: Northern N Pacific Ocean. 3 species. References Chace FA, Holthuis LB (1978) Psalidopus: the scissor-foot shrimps (Crustacea: Decapoda: Caridea). Smithsonian Contributions to Zoology 277, 1–22. doi:10.5479/si.00810282.277 Toriyama M, Horikawa H (1993) A new caridean shrimp, Psalidopus tosaensis, from Tosa Bay, Japan (Decapoda: Caridea, Psalidopodidae). Bulletin of the Nansei National Fisheries Research Institute 26, 1–8.

Superfamily Stylodactyloidea Bate, 1888 Stylodactylids are deep-water benthic shrimps notable for the exceptionally long fingers on the first two pairs of pereopods. There is only one family. All have long a long rostrum with numerous teeth with a basal suture. Observations from a submersible at over 4800 m depth led Wicksten et  al.

Superfamily Psalidopodoidea Wood-Mason, 1892 Diagnosis. Rostrum longer than carapace, ascendant, with dorsal, lateral and ventral series of spines. Mandible incisor conical, deeply separate from molar, with palp. Pereopods 1, 2 dissimilar, each pair symmetrical, first more robust than second. Pereopod 1 chelate, both fingers movable. Pereopod 2 carpus entire. Exopods absent from pereopods 1–5.

Psalidopodidae Wood-Mason, 1892 Figure 6.52, Plate 11j

Fig. 6.52.  Psalidopodidae. Psalidopus barbouri Chace, 1939: a, carapace, pleonites 1, 2; b, pereopod 1.

6 – Caridea – shrimps

(2017) to conclude that Bathystylodactylus bathyalis is a filter feeder. Diagnosis. Maxilliped 2 with distal 2 articles attached side by side (Fig. 6.53i). Pereopods 1, 2 similar, fingers extremely long, more than 10 times as long as wide, more than 5 times as long as palm, with long setae (Fig. 6.53j). Exopod present on maxilliped 3, pereopods 1–5.

171

Stylodactylidae Bate, 1888 Figure 6.53, Plate 11k Implicit generic attributes. Carapace without suprabranchial ridge. Pereopods 3–5 about as long as anterior pereopods. Arthrobranchs at base of pereopods 1–4 in both sexes.

Key to genera of Stylodactylidae 1. – 2. – 3. – 4. –

Mandibular palp absent������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Mandibular palp present (Fig. 6.53h)��������������������������������������������������������������������������������������������������������������������������������������������3 Supraorbital tooth distinct (Fig. 6.53b). Arthrobranchs at base of pereopods 1–4 in male only������������� Neostylodactylus Supraorbital tooth minute or absent (Fig. 6.53c). Arthrobranchs at base of pereopods 1–4 in both sexes��������������������������������������������������������������������������������������������������������������������������������������������������������Parastylodactylus Mandibular palp uniarticulate, at least in adult����������������������������������������������������������������������������������������������� Stylodactyloides Mandibular palp 2-articled (Fig. 6.53h)����������������������������������������������������������������������������������������������������������������������������������������4 Stylocerite acute, extending to or well beyond end of antennular peduncle article 1 (Fig. 6.53g). Pereopods 3–5 about as long as anterior pereopods. Carapace without suprabranchial ridge (Fig. 6.53e)��������������� Stylodactylus Stylocerite acute, extending nearly to midlength of antennular peduncle article 1 (Fig. 6.53f). Pereopods 3–5 ~1.5 times as long as anterior pereopods. Carapace with distinct suprabranchial ridge (Fig. 6.53a)�����������������������������������������������������������������������������������������������������������������������������������������������������Bathystylodactylus Bathystylodactylus Hanamura & Takeda, 1996

Diagnosis. Carapace with suprabranchial ridge; supraorbital tooth absent. Stylocerite acute, extending nearly to midlength of antennular peduncle article 1. Mandibular palp 2-articled.

Pereopods 3–5 ~1.5 as long as anterior pereopods. Maximum cl. 41 mm. Bathyal, abyssal (3436–4826 m). GOODS bathyal provinces: Cocos Plate, W Pacific. 3 species (Hanamura and Takeda 1996; Wicksten and Martin 2004).

Fig. 6.53.  Stylodactylidae. Carapace: a, Bathystylodactylus; b, Neostylodactylus; c; Parastylodactylus; d, Stylodactyloides; e, Stylodactylus. Antennular peduncle: f, Bathystylodactylus; g, Stylodactylus. Mandible: h, Bathystylodactylus. Maxilliped 2: i, Parastylodactylus. Pereopod 1: j, Neostylodactylus.

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Marine Decapod Crustacea

Neostylodactylus Hayashi & Miyake, 1968 Diagnosis. Carapace supraorbital tooth distinct. Stylocerite acute, extending beyond midlength of antennular peduncle article 1. Mandibular palp absent. Arthrobranchs at base of pereopods 1–4 in male only. Maximum cl. 4.4 mm. Subtidal–slope (9–522 m). Temperate Northern Pacific, Western and Central Indo-Pacific. 5 species (Cleva 1994; Komai 1997: addition of Cleva key).

Parastylodactylus Figueira, 1971 Diagnosis. Supraorbital tooth minute, or absent. Stylocerite acute, extending beyond midlength of antennular peduncle article 1. Mandibular palp absent. Maximum cl. 9.3 mm. Shelf, slope (106–700 m). Western and Central Indo-Pacific, Temperate Southern Africa. 8 species (Komai and Rajool Shanis 2011:key to species).

Stylodactyloides Cleva, 1990 Diagnosis. Supraorbital tooth distinct. Stylocerite rounded, extending nearly to midlength of antennular peduncle article 1. Mandibular palp uniarticulate at least in adult. Maximum cl. 18 mm. Slope (370–400 m). Central Indo-Pacific. 1 species (Cleva 1990).

Stylodactylus A. Milne-Edwards, 1881 Diagnosis. Supraorbital tooth distinct, or minute, or absent. Stylocerite acute, extending to or well beyond end of antennular peduncle article 1. Mandibular palp 2-articled. Maximum cl. 22 mm (Pl. 11k). Shelf, slope (122–1800 m). Tropical Atlantic, Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 16 species (Cleva 1990: key to 14 species). References Cleva R (1990) Crustacea Decapoda: Les genres et les espèces indo-ouest pacifiques de Stylodactylidae. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 6. Mémoires du Muséum National d’Histoire Naturelle, Paris 145, 71–136. Cleva R (1994) Some Australian Stylodactylidae (Crustacea: Decapoda), with descriptions of two new species. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 11, 53–64. Hanamura Y, Takeda M (1996) Establishment of a new genus Bathystylodactylus (Crustacea: Decapoda: Stylodactylidae), with description of a new species from Northwestern Pacific. Zoological Science 13, 929–934. doi:10.2108/zsj.13.929 Komai T (1997) A new species of the shrimp genus Neostylodactylus Hayashi and Miyake (Crustacea: Decapoda: Stylodactylidae) from Japan. Natural History Research 4, 125–133. Komai T, Rajool Shanis CP (2011) A new species of the genus Parastylodactylus Figueira, 1971 (Crustacea: Decapoda: Caridea: Stylodactylidae) from off Kollam, southwest coast of India. Zootaxa 3140, 60–68. doi:10.11646/zootaxa.3140.1.5

Fig. 6.54.  Anchialocarididae. Anchialocaris paulini Mejía-Ortíz, Yañez & López-Mejía, 2017. a, habitus; b, c, pereopods 1, 2. Wicksten MK, Martin JW (2004) A new species of caridean shrimp of the family Stylodactylidae from the eastern Pacific Ocean. Proceedings of the Biological Society of Washington 117, 377–384. Wicksten M, de Grave S, France S, Kelley C (2017) Presumed filter-feeding in a deep-sea benthic shrimp (Decapoda, Caridea, Stylodactylidae), with records of the deepest occurrence of carideans. ZooKeys 646, 17–23. doi:10.3897/zookeys.646.10969

Family incerta sedis A single species from anchialine caves in Mexico cannot be placed easily in any of the current superfamilies.

Anchialocarididae Mejía-Ortíz, Yañez & López-Mejía, 2017 Figure 6.54 Diagnosis. Rostrum short, triangular, without dorsal teeth. Carapace smooth. Eyestalks conical, without pigment. Mandible distinctly divided into broad incisor and slender conical molar, incisor, palp 2-articulated. Pereopods 1, 2 dissimilar, first shorter, slightly more slender than second, second with short propodus (Fig. 6.54b, c). Exopods present on maxilliped 3, pereopods 1, 2.

6 – Caridea – shrimps

Anchialocaris Mejía-Ortíz, Yañez & López-Mejía, 2017 Anchialocaris paulini Mejía-Ortíz, Yañez & López-Mejía, 2017 is found in the deeper parts of Cenote Chempita in Mexico, an anchialine environment. It differs from species of Agostocaris with which it co-occurs in having slender pereopods 1 and 2. Maximum cl. 14 mm. Marine anchialine caves. Tropical Atlantic (Mexico). 1 species (Mejía-Ortíz et al. 2017).

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References Mejía-Ortíz L, Yañez G, López-Mejía M (2017) Anchialocarididae, a new family of anchialine decapods and a new species of the genus Agostocaris from Cozumel Island, Mexico. Crustaceana 90, 381–398. doi:10.1163/15685403-00003657

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7. Astacidea – scampi and crayfish

The Astacidea Latreille, 1802, include a wide range of freshwater and marine crayfish, scampi and lobsters. They are differentiated from other decapods by the possession of three pairs of chelate or subchelate limbs of which the first is a strong pincer. Much of our detailed knowledge of the anatomy of Crustacea was gained in early research on astacideans and T.H. Huxley’s (1880) classic work, The crayfish: an introduction to the study of zoology, is based on the common European freshwater crayfish, Astacus astacus (Linnaeus, 1758). Astacidea comprise four superfamilies. Two are restricted to freshwater lakes, rivers and damp muddy environments – freshwater crayfish. Astacoidea Latreille, 1802, includes three families (Astacidae Latreille, 1802, Cambaridae Hobbs, 1942, and Cambaroididae Villabos, 1955) of northern hemisphere freshwater crayfishes or crawfish. Parastacoidea Huxley, 1879, of which there is a single family, Parastacidae Huxley, 1879, are the Southern Hemisphere freshwater crayfishes, restricted to Australia, South America and New Zealand. In Australia, a wide-ranging inland freshwater crayfish, Cherax destructor Clark, 1936, is known as the Yabby. Two superfamilies are marine and covered here. Each includes only a single living family, although other long extinct families are known from the fossil record (Ahyong 2006; Schweitzer et  al. 2010). Enoplometopoidea de Saint Laurent, 1988, are the dwarf reef lobsters, all in one genus,

Enoplometopus A. Milne-Edwards, 1862. Nephropoidea Dana, 1852, is more diverse, with 12 genera. Earlier keys to separate the families (Holthuis 1974, 1991; Poore 2004) are unfortunately unreliable. The key below relies on some of the characters used by de Saint Laurent (1988) who diagnosed Enoplometopoidea and Ahyong (2006) who explored the phylogeny of clawed lobsters. Molecular analyses support the monophyly of Astacidea, the four superfamilies and five families (Bracken-Grissom et al. 2014; Wolfe et al. 2019) although some earlier studies based on morphology had found Astacidea to be polyphyletic (e.g. Scholtz and Richter 1995). Wahle et al. (2012) reviewed the anatomy and biology of the marine astacideans. Diagnosis. Carapace subcylindrical, longer than wide; epistome short or elongated, not fused to basal antennular and antennal articles, lateral margins with narrow point of contact with lateral margin of carapace or full lateral margins with line of contact with carapace margins. Exoskeleton calcified. Thoracic sternites 1–8 fused or sternite 8 free. Pleon elongate, muscular; pleura not overlapping next most posterior pleuron, or pleuron 2 overlapping pleura 1 and 3. Antennular peduncle articles aligned linearly, main flagellum long, lash-like, aesthetascs more-or-less evenly distributed. Pereopods 1–3 chelate, first larger, claw-like. Pereopod 5 of similar length or shorter than more anterior pereopods. Uropods biramous.

Key to marine families of Astacidea 1.



Carapace with median spine row extending from proximal end of rostrum to carapace midlength; grooves absent or obsolete, postcervical groove incomplete, indicated dorsally only by shallow furrow. Pereopods 2, 3 subchelate (Fig. 7.1e). Telson with movable lateral and posterolateral spines movable spines (Fig. 7.1d). Pleonal pleura wide, round, overlapping adjacent pleura (Fig. 7.1c)���������������������������������� Enoplometopidae Carapace with or without median groove, without spine row (except Acanthacaris); grooves well developed, postcervical groove complete, deep, traversing lateral and dorsal surfaces. Pereopods 2, 3 chelate (Fig. 7.3s, t). Telson lateral and posterolateral spines, if present, fixed, immovable (Fig. 7.3u–A). Pleonal pleura narrow, scarcely or not overlapping adjacent pleura (Fig. 7.3i–l)������������������������������������������������������ Nephropidae

References Ahyong ST (2006) Phylogeny of the clawed lobsters (Crustacea: Decapoda: Homarida). Zootaxa 1109, 1–14. doi:10.11646/zootaxa.1109.1.1 Bracken-Grissom HD, Ahyong ST, Wilkinson RD, Feldmann RM, Schweitzer CE, et  al. (2014) The emergence of the lobsters: phylogenetic relationships, morphological evolution and divergence time comparisons of an ancient group (Decapoda: Achelata, Astacidea, Glypheidea, Polychelida). Systematic Biology 63, 457–479. doi:10.1093/sysbio/syu008

de Saint Laurent M (1988) Enoplometopoidea, nouvelle superfamille de Crustacés Décapodes Astacidea. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 307, 59–62. Holthuis LB (1974) Biological results of the University of Miami Deep-Sea Expeditions. 106. The lobsters of the superfamily Nephropidea of the Atlantic Ocean (Crustacea: Decapoda). Bulletin of Marine Science 24, 723–884. Holthuis LB (1991) FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue of species of interest to

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fisheries known to date. FAO Fisheries Synopsis. Vol. 125. Food and Agriculture Organization of the United Nations, Rome. Huxley TH (1880) The crayfish. An introduction to the study of zoology (International Scientific Series Vol. 28). Paul & Co., London. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Scholtz G, Richter S (1995) Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca). Zoological Journal of the Linnean Society 113, 289–328. doi:10.1006/zjls.1995.0011 Schweitzer CE, Feldmann RM, Garassino A, Karasawa H, Schweigert G (2010) Systematic list of fossil decapod crustacean species. Crustaceana Monographs 10, 1–222. doi:10.1163/ej.9789004178915.i-222 Wahle RA, Tshudy D, Cobb JS, Factor JR, Jaini M (2012) Chapter 70. Infraorder Astacidea Latreille, 1802 p.p.: the marine clawed lobsters. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea. Complementary to the volumes translated from the French of the Traité de Zoologie [founded by P.-P. Grassé]. (Eds) Schram FR, von Vaupel Klein JC. pp. 3–108. Brill, Leiden. Wolfe JM, Breinholt JW, Crandall KA, Lemmon AR, Lemmon EM, et al. (2019) A phylogenomic framework, evolutionary timeline and genomic resources for comparative studies of decapod crustaceans. Proceedings of the Royal Society B: Biological Sciences 286, 20190079. doi:10.1098/rspb.2019.0079

Enoplometopidae de Saint Laurent, 1988 dwarf reef lobsters Figure 7.1, Plate 13a–c Dwarf reef lobsters look like deep-water scampi and freshwater crayfish, differing most obviously in the tufts of long stiff, iridescent setae covering the body. The carapace of enoplometopids is largely smooth, with scant indication of grooves, whereas in nephropids the grooves are well marked. All enoplometopids are brightly coloured, usually with patterns over a background of orange or red (see photos in Chan and Ng 2008; Chan and Yu 1998; Holthuis 1983; Poupin 2002, 2003), and have become popular as aquarium pets. Indeed, three species, E. daumi Holthuis, 1983, E. debelius Holthuis, 1983, and E. crosnieri Chan, 1998, were first described as the result of discovery through the aquarium trade. Enoplometopids generally inhabit coral reefs

from shallow water to slope depths. Enoplometopus is the only genus but it has a complicated taxonomic history (Holthuis 1974; de Saint Laurent 1988). Two subgenera or genera have been suggested (Holthuis 1983) and rejected (Poupin 2003). Diagnosis. Carapace with median spine row extending from proximal end of rostrum to carapace midlength; without antennal groove; without hepatic groove (Fig. 7.1a, b); postercervical groove weakly indicated. Pleonal pleura wide, round, overlapping adjacent pleura (Fig. 7.1c). Telson with movable lateral and posterolateral spines (Fig. 7.1d). Pereopods 2, 3 subchelate (dactylus longer than fixed finger, fixed finger a short stump tipped with short movable spines; Fig. 7.1e). Body and pereopods covered with stiff, widely spaced, iridescent setae. Enoplometopus A. Milne-Edwards, 1862 Maximum cl. 55 mm (Pl. 13a–c). Intertidal–slope (0–300 m). Tropical Atlantic, Indo-West Pacific, Temperate Australasia. 12 species (Chan and Ng 2008: review and key to species; Poupin 2003: review, colour photos, key to 11 species). Photographs, diagnoses and links to the literature can be found at an interactive website (Poupin 2002). References Chan T-Y, Ng PKL (2008) Enoplometopus A. Milne-Edwards, 1862 (Crustacea: Decapoda: Nephropoidea) from the Philippines, with description of one new species and a revised key to the genus. Bulletin of Marine Science 83, 347–365. Chan T-Y, Yu H-P (1998) A new reef lobster of the genus Enoplometopus A. Milne Edwards, 1862 (Decapoda, Nephropoidea) from the western and southern Pacific. Zoosystema 20, 183–192. de Saint Laurent M (1988) Enoplometopoidea, nouvelle superfamille de Crustacés Décapodes Astacidea. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 307, 59–62. Holthuis LB (1974) Biological results of the University of Miami Deep-Sea Expeditions. 106. The lobsters of the superfamily Nephropidea of the Atlantic Ocean (Crustacea: Decapoda). Bulletin of Marine Science 24, 723–884.

Fig. 7.1.  Enoplometopidae, Enoplometopus. a, b, carapace; c; pleonites 1–6; d, telson, uropod; e, pereopod 2, end of propodus, dactylus.

7 – Astacidea – scampi and crayfish

Holthuis LB (1983) Notes on the genus Enoplometopus, with descriptions of a new subgenus and two new species (Crustacea, Decapoda, Axiidae). Zoologische Mededelingen 56, 281–298. Poupin J (2002) Tropical reef lobsters of the genus Enoplometopus: descriptions, illustrations, identification, and information retrieval, using the DELTA format Version: 30 May 2002. . Poupin J (2003) Reef lobsters Enoplometopus A. Milne Edwards, 1862 from French Polynesia, with a brief revision of the genus (Crustacea, Decapoda, Enoplometopidae). Zoosystema 25, 643–664.

Nephropidae Dana, 1852 Figures 7.2, 7.3, Plate 13d–h The best known nephropids are the Norwegian lobster or scampi, Nephrops norvegicus (Linnaeus, 1758), which is the basis of a fishery in Europe, and the Maine lobster, Homarus americanus H. Milne Edwards, 1837, which is also fished. Homarus americanus may attain an impressive weight of 12 kg, but most nephropids are of more modest size and are distributed in the tropical Indo-West Pacific. Until relatively recently, Thaumastochelidae, a small group with huge pectinate claws, was recognised as a separate family alongside the nephropids. The thaumastochelids are a monophyletic group but molecular evidence finds it nested

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within Nephropidae s.s. (Bracken-Grissom et al. 2014; Tshudy et al. 2009; Wolfe et al. 2019). For this reason, the family is no longer recognised. Relationships between the genera, including fossil genera, have been well investigated (Chang et  al. 2017; Rode and Babcock 2003; Tam and Kornfield 1998; Tshudy and Babcock 1997). Species were listed by Chan (2010) but others have been added since. Holthuis (1974, 1991) provided detailed and up-to-date treatments of the family. In the key below chelipeds refer only to pereopods 1. Diagnosis. Rostrum and carapace with or without median groove but without row of dorsal spines (except Acanthacaris). Carapace with antennal and hepatic grooves; postcervical groove well marked, complete. Pleonal pleura narrow, overlapping or not overlapping adjacent pleura. Telson with or without lateral or posterolateral spines. Pereopods 2, 3 chelate (dactylus as long as fixed finger). Implicit generic attributes. Rostrum dorsoventrally depressed with lateral and sometimes ventral teeth, without median dorsal teeth, sometimes without any teeth. Body never with spinules, sometimes with granules, or spinules only on carapace. Chelipeds with fingers less than twice as long as palm, teeth on cutting edge in same plane.

Key to genera of Nephropidae 1. – 2. – 3. – 4. – 5. – 6. –

Eyes fixed or reduced, without pigment (Fig. 7.3b, e, f). Telson margins unarmed (Fig. 7.3v, y, z). Chelipeds strongly unequal, strongly dissimilar; major with fingers more than 4 times as long as palm, cutting edges of fingers with many slender spines (Fig. 7.3r). Pleonal pleura broad, truncate (Fig. 7.3i)�����������������������������������2 Eyes movable, pigmented or not (Fig. 7.3a, c, d, g, h). Telson usually with posterolateral spines (Fig. 7.3u, w, A). Chelipeds equal or unequal, similar; fingers less than twice as long as palm (Figs 7.2, 7.3o–q). Pleonal pleura approximately triangular, pointed�����������������������������������������������������������������������������������������������������������������4 Uropodal exopod, distal article short, wide, half or more than half as wide as proximal article (Fig. 7.3v, y)�����������������3 Uropodal exopod, distal article minute, semicircular, narrower than one-quarter width of proximal article (Fig. 7.3z)����������������������������������������������������������������������������������������������������������������������������������������������������� Thaumastochelopsis Eyes slender, movable (Fig. 7.3b). Epistome with slender median and posterior portions, T-shaped; antenna surrounded by wide band of arthrodial membrane, articulating with anterolateral margins of epistome (Fig. 7.3m)������������������������������������������������������������������������������������������������������������������������������������������������������������������� Dinochelus Eyes short, stout, fixed (Fig. 7.3e). Epistome with broad median and posterior portions; antenna fitting tightly into anterolateral margins of epistome (Fig. 7.3n)������������������������������������������������������������������������ Thaumastocheles Rostrum laterally compressed, with dorsal and ventral teeth. Body uniformly and densely spinulose (Fig. 7.3a). Telson with 6–12 lateral spines (Fig. 7.3u)�������������������������������������������������������������������������������������� Acanthacaris Rostrum dorsoventrally depressed, with lateral teeth. Body ornamented but not uniformly spinulose. Telson with at most 3 small lateral spines�������������������������������������������������������������������������������������������������������������������������������5 Scaphocerite absent (Fig. 7.2a). Carapace without postorbital spine. Eyes small, unpigmented or with small cornea (Figs 7.2a, 7.3h)����������������������������������������������������������������������������������������������������������������������������������������������������������������6 Scaphocerite present (Fig. 7.2b). Carapace with postorbital spine. Eyes large, usually pigmented (Figs 7.2b, 7.3d, c)�������������������������������������������������������������������������������������������������������������������������������������������������������������������������8 Pleonal pleura broad, overlapping, ventral angles with short triangular tooth (Fig. 7.3l). Telson lateral margin often with 1 or 2 spines plus posterolateral spine (Fig. 7.3A)�������������������������������������������������������������������Thymops Pleonal pleura narrow, scarcely overlapping, ventral angles acute or square. Telson lateral margin unarmed or tuberculate except for posterolateral spine (Fig. 7.2a)������������������������������������������������������������������������������������������������������7

178

Marine Decapod Crustacea

Eye unpigmented (Fig. 7.2b). Body granular, setose. Pleonal pleura attenuated to acute spine (Fig. 7.3k)������� Nephropsis Eye pigmented. Body covered by conspicuous rounded pearly tubercles. Pleonal pleuron 2 rectangular, others angular (Fig. 7.3j)����������������������������������������������������������������������������������������������������������������������������������������Nephropides 8. Chelipeds unequal, major with crushing finger, minor with cutting fingers�������������������������������������������������������������������������9 – Chelipeds subequal, similar����������������������������������������������������������������������������������������������������������������������������������������������������������11 9. Cheliped palm with longitudinal grooves, ridges and rows of spines (Fig. 7.3q). Subdorsal carinae spinulate. Pleonites with transverse grooves, blunt carina separating tergites from pleura���������������������������������������������� Nephrops – Cheliped palm smooth. Subdorsal carinae without spinules. Pleonites smooth�����������������������������������������������������������������10 10. Body, chelipeds non-setose (Fig. 7.3p). Telson tapering to rounded posterior margin, smooth (Fig. 7.3x)��������� Homarus – Body, outer surfaces of chelipeds setose (Fig. 7.3o). Telson subparallel, bearing obsolescent spines and ridges (Fig. 7.3w)��������������������������������������������������������������������������������������������������������������������������������������������������������Homarinus 11. Antennal spine not carinate. Cornea unpigmented (Fig. 7.3g)����������������������������������������������������������������������������� Thymopides – Antennal spine carinate. Cornea pigmented (Figs 7.2a, 7.3d)������������������������������������������������������������������������������������������������ 12 12. Supraorbital spine followed by strong toothed ridge extending almost to postcervical groove. Posterior part of carapace with several longitudinal carinae (Fig. 7.2a)�������������������������������������������������������������������������������Metanephrops – Supraorbital spine followed by 1 spine. Posterior part of carapace granulate (Fig. 7.3c)����������������������������������� Eunephrops 7. –

Acanthacaris Bate, 1888

Homarinus Kornfield, Williams & Steneck, 1995

giant deep-sea lobster

Cape lobster

Diagnosis. Rostrum laterally compressed, with dorsal and ventral, without lateral teeth. Body densely covered by closely placed, sharp spinules. Telson with 6–12 lateral spines. Eyes reduced, unpigmented. Maximum cl. 212 mm (Pl. 13d). Slope–bathyal (293–2161 m). Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific. 2 species (Chang et al. 2015: colour photograph).

Diagnosis. Body, outer surfaces of chelipeds setose. Carapace with postorbital spine, subdorsal carinae without spinules. Pleonites smooth. Telson subparallel, bearing obsolescent spines and ridges. Eyes large, movable, with pigment. Scaphocerite present. Chelipeds unequal, major with crushing claw, palm longer than wide; minor with cutting fingers, palm smooth. Maximum cl. 32 mm. Intertidal–shelf (0–40 m). Temperate Southern Africa. 1 species (Kornfield et al. 1995).

Dinochelus Ahyong, Chan & Bouchet, 2010 Diagnosis. Pleonal pleura 2–5 broad, truncate, not ending in point. Telson unarmed. Eyes slender, slightly movable, without pigment. Epistome with slender median and posterior portions, T-shaped; antenna articulating with the anterolateral margins of epistome. Scaphocerite inner margin toothed. Chelipeds strongly unequal; larger one with fingers more than 4 times as long as palm, cutting edges of fingers with many slender spines. Uropodal exopod, distal article short, more than half as wide as proximal article, reaching anterior margin of proximal article. Maximum cl. 31 mm (Pl. 13e). Slope (247–249 m). Central Indo-Pacific. 1 species (Ahyong et al. 2010).

Eunephrops Smith, 1885 Diagnosis. Carapace supraorbital spine followed by 1 spine, with postorbital spine, antennal spine carinate, posterior part granulate. Eyes large, movable, with pigment. Scaphocerite present. Chelipeds subequal. Maximum cl. 90 mm. Shelf, slope (230–824 m). Tropical Atlantic (Caribbean). 4 species (Holthuis 1974; Manning 1997: key to species).

Fig. 7.2.  Nephropidae. a, Nephropsis serrata Macpherson, 1993; b, Metanephrops boschmai (Holthuis, 1964).

7 – Astacidea – scampi and crayfish

179

Fig. 7.3.  Nephropidae. Anterior carapace: a, Acanthacaris; b, Dinochelus; c, Eunephrops; d, Metanephrops; e, Thaumastocheles; f, Thaumastochelopsis; g, Thymopides; h, Thymops. Pleonites 1–6: i, Dinochelus; j, Nephropides; k, Nephropsis; l, Thymops. Epistome, base of antennae: m, Dinochelus; n, Thaumastocheles. Chelipeds: o, Homarinus; p, Homarus; q, Nephrops; r, Thaumastocheles. Pereopod 2, propodus, dactylus: s, Nephropsis; t, Thaumastocheles. Telson, uropod: u, Acanthacaris; v, Dinochelus; w, Homarinus; x, Homarus; y, Thaumastocheles; z, Thaumastochelopsis; A, Thymops.

Homarus Weber, 1795 American lobster, Maine lobster, European lobster Diagnosis. Body, chelipeds non-setose. Carapace with postorbital spine, subdorsal carinae without spinules. Pleonites smooth. Telson tapering to rounded posterior margin, smooth. Eyes large, movable, with pigment. Scaphocerite present. Chelipeds unequal, major with crushing claw, palm as wide as long; minor with cutting fingers, palm smooth. Maximum cl. 300 mm. Intertidal–shelf (0–480 m). Temperate Northern Atlantic. 2 species (Holthuis 1991).

Metanephrops Jenkins, 1972 scampi, langoustine Diagnosis. Carapace supraorbital spine followed by strong toothed ridge extending almost to postcervical groove, with postorbital spine,

antennal spine carinate, posterior part with several longitudinal carinae. Eyes large, movable, with pigment. Scaphocerite present. Chelipeds subequal. Maximum cl. 80 mm (Pl. 13f). Shelf, slope (50–994 m). Temperate Northern Pacific, Tropical Atlantic, Western and Central Indo-Pacific, Temperate South America, Temperate Southern Africa, Temperate Australasia. 18 species (Chan 1997: key to species, four species groups, distributions; Chan et al. 2009: origin and evolution). Species of Metanephrops are the basis of commercial fisheries, so are well studied (e.g. Robey et al. 2013; Verry et al. 2020).

Nephropides Manning, 1969 Diagnosis. Body covered by conspicuous rounded pearly tubercles. Carapace without postorbital spine. Pleonal pleura 2–5 narrow, scarcely overlapping, ventral margin truncate. Eyes small, with small pigmented cornea. Scaphocerite absent. Maximum cl. 60 mm. Slope (511–728 m). Tropical Atlantic (Caribbean). 1 species (Manning 1969).

180

Marine Decapod Crustacea

Nephrops Leach, 1814 Norway lobster Diagnosis. Carapace with postorbital spine, subdorsal carinae spinulate. Pleonites with transverse grooves, blunt carina separating tergites from pleura. Eyes large, movable, with pigment. Scaphocerite present. Chelipeds unequal, major with crushing claw, minor with cutting fingers, palm with longitudinal grooves, ridges and rows of spines. Maximum cl. 100 mm. Shelf, slope (20–800 m). Temperate Northern E Atlantic. 1 species (Holthuis 1991). The only species is of considerable commercial value throughout its range. Subpopulations have been identified (Angelini et al. 2020; Gallagher et al. 2019).

Nephropsis Wood-Mason, 1872 Diagnosis. Body granular, setose. Carapace without postorbital spine. Pleonal pleura 2–5 narrow, scarcely overlapping, attenuated to acute spine. Eyes small, unpigmented. Scaphocerite absent. Maximum cl. 71 mm (Pl. 13g). Shelf–bathyal (137–2900 m). Temperate Northern and Tropical Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate Australasia. 16 species (Chang et  al. 2020: 3 species resolved; Holthuis 1974: Atlantic species; Macpherson 1990: Indo-West Pacific species; key to 13 species).

Thaumastocheles Wood-Mason, 1874 Diagnosis. Pleonal pleura 2–5 broad, truncate, not ending in point. Telson unarmed. Eyes unpigmented, fixed. Epistome with broad median and posterior portions; antenna fitting tightly into anterolateral margins of epistome. Scaphocerite inner margin toothed. Chelipeds strongly unequal; larger one with fingers more than 4 times as long as palm, cutting edges of fingers with many slender spines. Uropodal exopod, distal article short, more than half as wide as proximal article, reaching anterior margin of proximal article. Maximum cl. 60 mm (Pl. 13h). Slope (250–1767 m, possibly as shallow as 50 m). Tropical W Atlantic, Central Indo-Pacific. 4 species (Chang et al. 2014).

Thaumastochelopsis Bruce, 1988 Diagnosis. Pleonal pleura 2–5 broad, truncate, not ending in point. Telson unarmed. Eyes slender, slightly movable, without pigment. Epistome with broad median and posterior portions; antenna fitting tightly into anterolateral margins of epistome. Scaphacerite inner margin toothed. Chelipeds strongly unequal; larger one with fingers more than 4 times as long as palm, cutting edges of fingers with many slender spines. Uropodal exopod, distal article minute, semicircular, narrower than one-quarter of proximal article. Maximum cl. 25 mm. Shelf, slope (199–425 m). Central Indo-Pacific. 2 species (Ahyong et al. 2007: key to 5 thaumastochelid species).

Thymopides Burukovsky & Averin, 1977 Diagnosis. Carapace with postorbital spine, antennal spine noncarinate. Pleonites with median carina. Eyes slender, slightly

small, movable, without pigment. Scaphocerite present. Chelipeds subequal. Maximum cl. 40 mm. Slope, bathyal (525–1220 and 3505 m). Southern Ocean. GOODS bathyal provinces: North Atlantic Ocean (hydrothermal vent). 2 species (Ledoyer 1974; Segonzac and Macpherson 2003).

Thymops Holthuis, 1974 Diagnosis. Body granular, setose. Carapace without postorbital spine. Pleonal pleura 2–5 broad, overlapping, ventral angles with short triangular tooth. Telson lateral margin often with 1 or 2 spines plus posterolateral spine. Eyes small, movable, pigmented. Scaphocerite absent. Maximum cl. 60 mm. Slope, bathyal (122–3040 m). Temperate South America. GOODS bathyal province: Subantarctic. 3 species. Ahyong et al. (2012) reviewed the three species of Thymops and Thymopsis Holthuis, 1974, concluding that only slight morphological differences separated them. All occur in the South Atlantic. Thymopsis is treated as a junior synonym here. References Ahyong ST, Chu KH, Chan T-Y (2007) Description of a new species of Thaumastochelopsis from the Coral Sea (Crustacea: Decapoda: Nephropoidea). Bulletin of Marine Science 80, 201–208. Ahyong ST, Chan T-Y, Bouchet P (2010) Mighty claws: a new genus and species of lobster from the Philippine deep sea (Crustacea, Decapoda, Nephropidae). Zoosystema 32, 525–535. doi:10.5252/z2010n3a11 Ahyong ST, Webber R, Chan T-Y (2012) Thymops takedai, a new species of deepwater lobster from the southwest Atlantic Ocean with additional records of ‘thymopine’ lobsters (Decapoda, Nephropidae). In: Komatsu H, Okuno J, Fukuoka K (Eds) Studies on Eumalacostraca: a homage to Masatsune Takeda. Crustaceana Monographs 17, 49–61. doi:10.1163/9789004202894_004 Angelini S, Martinelli M, Santojanni A, Colella S (2020) Biological evidence of the presence of different subpopulations of Norway lobster (Nephrops norvegicus) in the Adriatic Sea (Central Mediterranean Sea). Fisheries Research 221, 105365. doi:10.1016/j.fishres.2019.105365 Bracken-Grissom HD, Ahyong ST, Wilkinson RD, Feldmann RM, Schweitzer CE, et  al. (2014) The emergence of the lobsters: phylogenetic relationships, morphological evolution and divergence time comparisons of an ancient group (Decapoda: Achelata, Astacidea, Glypheidea, Polychelida). Systematic Biology 63, 457–479. doi:10.1093/sysbio/syu008 Chan T-Y (1997) Crustacea Decapoda: Palinuridae, Scyllaridae and Nephropidae collected in Indonesia by the KARUBAR cruise with an identification key for the species of Metanephrops In: Crosnier A, Bouchet P (Eds), Résultats des Campagnes MUSORSTOM, vol. 16. Mémoires du Muséum National d’Histoire Naturelle, Paris 172, 409–431. Chan T-Y (2010) Annotated checklist of the world’s marine lobsters (Crustacea: Decapoda: Astacidea, Glypheidea, Achelata, Polychelida). Raffles Bulletin of Zoology Supplement 23, 153–181. Chan T-Y, Ho KC, Li CP, Chu KH (2009) Origin and diversification of the clawed lobster genus Metanephrops (Crustacea: Decapoda: Nephropidae). Molecular Phylogenetics and Evolution 50, 411–422. doi:10.1016/j. ympev.2008.11.020 Chang S-C, Chan T-Y, Ahyong ST (2014) Two new species of the rare lobster genus Thaumastocheles Wood-Mason, 1874 (Reptantia: Nephropidae) discovered from recent deep-sea expeditions in the Indo-West Pacific. Journal of Crustacean Biology 34, 107–122. doi:10.1163/1937240X-00002201

7 – Astacidea – scampi and crayfish

Chang S-C, Wang T-W, Chan T-Y (2015) First record of the nephropid genus Acanthacaris Bate, 1888 (Crustacea: Decapoda: Nephropidae) from Taiwan. Zootaxa 4032, 595–599. doi:10.11646/zootaxa.4032.5.8 Chang S-C, Tshudy D, Sorhannus U, Ahyong ST, Chan T-Y (2017) Evolution of the thaumastocheliform lobsters (Crustacea, Decapoda, Nephropidae). Zoologica Scripta 46, 372–387. doi:10.1111/zsc.12205 Chang S-C, Chan T-Y, Kumar AB (2020) Deep-sea clawed lobster Nephropsis stewarti Wood-Mason, 1872 species complex in the IndoWest Pacific (Crustacea, Decapoda, Nephropidae), with description of a new species. ZooKeys 1008, 37–60. doi:10.3897/zookeys.1008.59966 Gallagher J, Finarelli JA, Jonasson JP, Carlsson J (2019) Mitochondrial D-loop DNA analyses of Norway lobster (Nephrops norvegicus) reveals genetic isolation between Atlantic and East Mediterranean populations. Journal of the Marine Biological Association of the United Kingdom 99, 933–940. doi:10.1017/S0025315418000929 Holthuis LB (1974) Biological results of the University of Miami Deep-Sea Expeditions. 106. The lobsters of the superfamily Nephropidea of the Atlantic Ocean (Crustacea: Decapoda). Bulletin of Marine Science 24, 723–884. Holthuis LB (1991) FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue of species of interest to fisheries known to date. FAO Fisheries Synopsis. Vol. 125. Food and Agriculture Organization of the United Nations, Rome. Kornfield I, Williams AB, Steneck RS (1995) Assignment of Homarus capensis (Herbst, 1792), the Cape Lobster of South Africa, to the new genus Homarinus (Decapoda: Nephropidae). Fishery Bulletin 93, 97–102. Ledoyer M (1974) Thymopides grobovi (Burukovsky & Averin, 1976) (Crustacea, Decapoda, Nephropidae) récolté aux îles Kerguelen au cours des campagnes du M.S. “Marion Dufresne” en 1974 et 1975. Tethys 9, 123–128. Macpherson E (1990) Crustacea Decapoda: on a collection of Nephropidae from the Indian Ocean and Western Pacific. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 6. Mémoires du Muséum National d’Histoire Naturelle, Paris 145, 289–328. Manning RB (1969) A new genus and species of lobster (Decapoda, Nephropidae) from the Caribbean Sea. Crustaceana 17, 303–309. doi:10.1163/156854069X00655

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Manning RB (1997) Eunephrops luckhursti, a new deep sea lobster from Bermuda (Crustacea: Decapoda: Nephropidae). Proceedings of the Biological Society of Washington 110, 256–262. Robey J, Fennessy ST, Everett BI, Santos J, Groeneveld JC (2013) Abundance and biology of the langoustine Metanephrops mozambicus (Nephropidae) on deep-water trawl grounds in eastern South Africa. Journal of Crustacean Biology 33, 760–771. doi:10.1163/1937240X-00002194 Rode AL, Babcock LE (2003) Phylogeny of fossil and extant freshwater crayfish and some closely related nephropid lobsters. Journal of Crustacean Biology 23, 418–435. doi:10.1163/20021975-99990351 Segonzac M, Macpherson E (2003) A new deep-sea lobster of the genus Thymopides (Crustacea: Decapoda: Nephropidae) collected near the hydorthermal vent Snake Pit, Mid-Atlantic Ridge. Cahiers de Biologie Marine 44, 361–367. Tam YK, Kornfield I (1998) Phylogenetic relationships of clawed lobster genera (Decapoda: Nephropidae) based on mitochondrial 16S rRNA gene sequences. Journal of Crustacean Biology 18, 138–146. doi:10.2307/1549528 Tshudy D, Babcock LE (1997) Morphology-based phylogenetic analysis of the clawed lobsters (family Nephropidae and the new family Chilenohphoberidae). Journal of Crustacean Biology 17, 253–263. doi:10.2307/1549275 Tshudy D, Robles R, Chan T-Y, Ho KC, Chu KH, et al. (2009) Phylogeny of marine clawed lobster families Nephropidae Dana, 1852, and Thaumastochelidae Bate, 1888, based on mitochondrial genes. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 357–367. CRC Press, Boca Raton. Verry AJF, Walton K, Tuck ID, Ritchie PA (2020) Genetic structure and recent population expansion in the commercially harvested deep-sea decapod, Metanephrops challengeri (Crustacea: Decapoda). New Zealand Journal of Marine and Freshwater Research 54, 251–270. doi:1 0.1080/00288330.2019.1707696 Wolfe JM, Breinholt JW, Crandall KA, Lemmon AR, Lemmon EM, et  al. (2019) A phylogenomic framework, evolutionary timeline and genomic resources for comparative studies of decapod crustaceans. Proceedings of the Royal Society B: Biological Sciences 286, 20190079. doi:10.1098/ rspb.2019.0079

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8. Glypheidea – lobsters

Glypheidean lobsters appeared in the Upper Permian, radiated extensively in the Triassic (Charbonnier et  al. 2013) and then rapidly declined. They were believed to be long extinct until living specimens, almost indistinguishable from some of their extinct relatives, were discovered in the Philippines in 1975 and recognised as ‘living fossils’ (Forest and de Saint Laurent 1975, 1976, 1981). Three decades later, a second species was found in New Caledonia and placed in a second genus (Boisselier-Dubayle et al. 2010; Forest 2006a; Richer de Forges 2006). For the century or so before the discovery of living species, and even decades afterwards, glypheideans were aligned with other achelate lobsters in the old grouping, Palinuridea, owing to the superficially similar structure of the first pereopods (Glaessner 1969; Holthuis 1991). Glypheideans share features with, but differ from, other lobsters (Achelata, Astacidea, Polychelida). Like Achelata, extant glypheideans do not have fully chelate first pereopods but this resemblance has proved to be superficial (Schram 2001). The subchelate first pereopod and simple pereopods 2 and 3 of modern glypheideans, both in the family Glypheidae, differ from the true chelae on pereopods 1–3 seen in Astacidea but Forest and de Saint Laurent (1989) and Forest (2006b) believed the two groups more closely related than to other reptant decapods. Confirmation of this affinity was found

in the observation that thoracic sternites 7 and 8 articulate, as in members of ‘Fractosternalia’, a group comprising some groups of Astacidea (Scholtz and Richter 1995), and similar alignment of the epistome (Schram and Ahyong 2002). This was confirmed further by analysis of molecular data and since then, the glypheideans have been accepted as representing a separate decapod infraorder (Ahyong and O’Meally 2004; Bracken-Grissom et al. 2014; Tan et al. 2018). Diagnosis. Carapace subcylindrical, longer than wide; epistome strongly elongate, not fused to basal antennular and antennal articles, lateral margins with long line of contact with carapace margins. Exoskeleton calcified. Thoracic sternite 8 free from thoracic sternites 1–7. Pleon elongate, muscular; pleura not overlapping next most posterior pleuron. Antennular peduncle articles aligned linearly, main flagellum long, lash-like, aesthetascs more-or-less evenly distributed. Pereopods 1–5 simple (in extant taxa), first larger. Pereopod 5 shorter than others. Uropods biramous.

Glypheidae Winkler, 1882 Figure 8.1 The family has the features of the infraorder. Forest (2006a) described more characters separating the two genera than are included here.

Fig. 8.1.  Glypheidae. a, Neoglyphea inopinata Forest & de Saint Laurent, 1989. Carapace, precervical region: b, Neoglyphea; c, Laurentaeglyphea. Pereopod 1. d, Laurentaeglyphea.

184

Marine Decapod Crustacea

Laurentaeglyphea Forest, 2006 Diagnosis. Carapace dorsally smooth or punctate; precervical surface either side of median groove with 3 strong smooth carinae (Fig. 8.1c). Pereopod 1 not reaching beyond eyes, propodus oval, twice as long as wide, subchelate (Fig. 8.1d). Maximum cl. 27 mm. Slope (367–536 m). Central Indo-Pacific (New Caledonia). 1 species (Forest 2006a; Richer de Forges 2006).

Neoglyphea Forest & de Saint Laurent, 1975 Diagnosis. Carapace dorsally densely covered with spines-tubercles; precervical surface either side of median groove with 3 lines of stronger spines (Fig. 8.1a, b). Pereopod 1 merus reaching beyond eyes, propodus 6 times as long as wide, linear (Fig. 8.1a). Maximum cl. 55 mm. Shelf (186–210 m). Central Indo-Pacific (Philippines, Indonesia). 1 species (Forest and de Saint Laurent 1975, 1981). References Ahyong ST, O’Meally D (2004) Phylogeny of the Decapoda Reptantia: resolution using three molecular loci and morphology. Raffles Bulletin of Zoology 52, 673–693. Boisselier-Dubayle M-C, Bonillo C, Cruaud C, Couloux A, Richer de Forges B, et al. (2010) The phylogenetic position of the ‘living fossils’ Neoglyphea and Laurentaeglyphea (Decapoda: Glypheidea). Comptes Rendus Biologies 333, 755–759. doi:10.1016/j.crvi.2010.08.007 Bracken-Grissom HD, Ahyong ST, Wilkinson RD, Feldmann RM, Schweitzer CE, et  al. (2014) The emergence of the lobsters: phylogenetic relationships, morphological evolution and divergence time comparisons of an ancient group (Decapoda: Achelata, Astacidea, Glypheidea, Polychelida). Systematic Biology 63, 457–479. doi:10.1093/sysbio/syu008 Charbonnier S, Garassino A, Schweigert G, Simpson M (2013) A worldwide review of fossil and extant glypheid and litogastrid lobsters (Crustacea, Decapoda, Glypheoidea). Mémoires du Muséum d’Histoire Naturelle, Marseille 28, 1–280. Forest J (2006a) Laurentaeglyphea, un nouveau genre pour la seconde espèce actuelle de Glyphéide récemment découverte (Crustacea Décapoda Glypheidae). Comptes Rendus Biologies 329, 841–846. doi:10.1016/j.crvi.2006.08.003 Forest J (2006b) The Recent glypheids and their relationship with their fossil relatives (Decapoda, Reptantia) Les glyphéides actuels et leur

relation avec les formes fossiles (Decapoda, Reptantia). Crustaceana 79, 769–793. doi:10.1163/156854006778008212 Forest J, de Saint Laurent M (1975) Présence dans la faune actuelle d’un représentant du groupe mésozoïque des Glyphéides: Neoglyphea inoponata gen. nov., sp. nov. (Crustacea Decapoda Glypheidae). Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 281, 155–158. Forest J, de Saint Laurent M (1976) Capture aux Philippines de nouveaux exemplaires de Neoglyphea inopinata (Crustacea Decapoda Glypheidae). Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 283, 935–938. Forest J, de Saint Laurent M (1981) La morphologie externe de Neoglyphea inopinata, espèce actuelle de Crustacé Décapode Glyphéide. In: Forest J (Ed.), Résultats des Campagnes MUSORSTOM. 1. Philippines (18–28 mars 1976). Mémoires du Muséum National d’Histoire Naturelle (Série A, Zoologie) 91, 51–84. Forest J, de Saint Laurent M (1989) Nouvelle contribution à la connaissance de Neoglyphea inopinata Forest & de Saint Laurent, à propos de la description de la femelle adulte. In: Forest J (Ed.), Résultats des Campagnes MUSORSTOM, vol. 5. Mémoires du Muséum National d’Histoire Naturelle, Paris 144, 75–92. Glaessner MF (1969) Decapoda. In Arthropoda 4. Part R. (Ed. Moore RC) pp. R399–R533. Geological Society of America/University of Kansas Press, Lawrence. Holthuis LB (1991) FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue of species of interest to fisheries known to date. FAO Fisheries Synopsis. Vol. 125. Food and Agriculture Organization of the United Nations, Rome. Richer de Forges B (2006) Découverte en mer du Corail d’une deuxième espèce de glyphéide (Crustacea, Decapoda, Glypheoidea). Zoosystema 28, 17–29. Scholtz G, Richter S (1995) Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca). Zoological Journal of the Linnean Society 113, 289–328. doi:10.1006/zjls.1995.0011 Schram FR (2001) Phylogeny of decapods: moving towards a consensus. Hydrobiologia 449, 1–20. doi:10.1023/A:1017543712119 Schram FR, Ahyong ST (2002) The higher affinities of Neoglyphea inopinata in particular and the Glypheoidea (Decapoda, Reptantia) in general. Crustaceana 75, 629–635. doi:10.1163/156854002760095651 Tan MH, Gan HM, Dally G, Horner SG, Rodríguez Moreno P, et al. (2018) More limbs on the tree: mitogenome characterisation and systematic position of ‘living fossil’ species Neoglyphea inopinata and Laurentaeglyphea neocaledonica (Decapoda : Glypheidea : Glypheidae). Invertebrate Systematics 32, 448–456. doi:10.1071/IS17050

9. Axiidea – sponge shrimps, ghost shrimps and mud lobsters

Axiidea comprise a diverse assortment of shrimp-like crustaceans. In axiideans the first two pairs of pereopods are always both chelate, the first being strong chelipeds and the second much smaller and similar in size to the simple pereopods 3 and 4. Pereopod 5 is often minutely chelate or subchelate. Thoracic sternite 8 to which it attaches is well separated from fused thoracic sternites 3–7 which more anterior limbs attach to (as in Gebiidea and Anomura). The families can be distinguished only on the basis of a combination of characters. Ghost shrimps, callianassids and similar families, tend to have an elongate pleon with weakly connected somites and weak pleura. Axiids and strahlaxiids are more robust and shrimp-like. Most are burrowers in soft sediments, or live under rocks or in crevices or commensally with sponges. For a detailed general review of biology see Dworschak et al. (2012) and family coverage below. Axiidea de Saint Laurent, 1979 is one of two infraorders until recently considered part of Thalassinidea Latreille, 1831. The other is Gebiidea (Chapter 10). The monophyly of Axiidea was proposed on morphological grounds (de Saint Laurent 1979; Sakai 2005b; Sakai and Sawada 2006) and confirmed by molecular studies (Robles et al. 2009; Tsang et al. 2008a; Tsang et al. 2008b). These authors discovered two clades of Axiidea, one comparing only Axiidae and the other the remaining families. The terminal families of the second clade were called ‘callianassoids’ by Robles et  al. (2020) and Poore et  al. (2019) who introduced many new genera and synonymised many others.

Poore (1994) provided keys to families and genera of what was then Thalassinidea. While some of these taxa are still valid the taxonomy has long since moved on. The most recent revisions of Axiidea (Sakai 2005a, 2011) treated the infrorder as either Callianassoidea or ‘Callianassida’. De Saint Laurent’s earlier name is now in general use (Dworschak et al. 2012; Poore et al. 2014). Sakai’s two papers introduced many new family and generic names, and synonymised others. Not all have been accepted (Dworschak 2007; Dworschak and Poore 2018; Poore 2008; Poore and Dworschak 2017). Our taxonomy relies on recent revisions (Poore 2015; Poore and Collins 2015) and the results of molecular and morphological revisions (Poore et  al. 2019; Robles et al. 2020). The infrorder comprises 11 families with 122 genera, most in Axiidae, Callianassidae and Callichiridae. Diagnosis. Carapace more or less cylindrical, longer than wide; epistome short, not fused to basal antennular and antennal articles, with narrow point of contact with lateral margin of carapace. Exoskeleton calcified, or chitinised (rarely). Thoracic sternite 8 free from thoracic sternites 1–7. Pleon elongate, muscular; pleonite 2 pleuron overlapping pleonites 1 and 3, or all pleura overlapping next most posterior pleuron. Antennular penduncle articles aligned linearly, main flagellum long, lash-like, aesthetascs more-or-less evenly distributed. Pereopods 1, 2 chelate, first a large cheliped. Pereopod 5 shorter than others, subchelate. Uropods biramous.

Key to families of Axiidea 1. –

2. – 3. –

Rostrum prominent, often with erect lateral spines, carapace with lateral gastric carinae originating from lateral margins of rostrum, often with submedian and median gastric carinae. Linea thalassinica absent. Eyestalks cylindrical (Figs 9.2–9.4, 9.24a–e)���������������������������������������������������������������������������������������������������������������������������2 Rostrum triangular, flat or reduced to short spine shorter than eyestalk, carapace without median and lateral gastric carinae (Figs 9.8, 9.14–9.16), or rostrum flat-unornamented, longer than eyestalk, carapace with lateral gastric carinae (Fig. 9.22). Linea thalassinica present over all or part of carapace length (Fig. 9.1, 9.15a), or absent (Fig. 9.22a), or short. Eyestalks contiguous, flat or cylindrical����������������������������������������������������������������3 Rostrum apex bifid, with lateral teeth (Fig. 9.24a, c). Pereopods 3 and 4 propodi without lateral spiniform setae (Fig. 9.24f)����������������������������������������������������������������������������������������������������������������������������������Strahlaxiidae Rostrum with acute or rounded apex (Figs 9.2–9.4). Pereopods 3 and 4 propodi with lateral spiniform setae (Fig. 9.6s–x)����������������������������������������������������������������������������������������������������������������������������������������������������������������������Axiidae Maxilla scaphognathite with long setae extending from posterior lobe into branchial chamber (Fig. 9.14c). Linea thalassinica complete, partial or absent (Fig. 9.22a)���������������������������������������������������������������������������������������������������4 Maxilla scaphognathite without long setae extending from posterior lobe into branchial chamber (Fig. 9.8q). Linea thalassinica complete over full carapace length (Figs 9.1b, 9.20a)��������������������������������������������������������5

186

4. – 5. – 6. – 7. – 8. – 9. – 10. –

Marine Decapod Crustacea

Posterior margin of carapace evenly curved, not interacting with anterolateral lobes on pleonite 1. Eyestalk flat, contiguous (Fig. 9.14a, b). Chelipeds flattened, asymmetrical (Fig. 9.14f–h)�������������������������������������Callianideidae Posterior margin of carapace with lateral lobes interacting with anterolateral lobes on pleonite 1 (Fig. 9.22a). Eyestalks cylindrical even if continuous (Fig. 9.22b–f). Chelipeds cylindrical, symmetrical (Fig. 9.22j, k)������������������������������������������������������������������������������������������������������������������������������������������������������������� Micheleidae Pleonite 1 with dorsal pair of lobes interacting with posterior margin of carapace (Fig. 9.23a). Female pleopod 1 with single broad expanded ramus (Fig. 9.23d)��������������������������������������������������������������������������Paracalliacidae Pleonite 1 with evenly curved dorsal margin, not interacting with posterior margin of carapace (Figs 9.1b, 9.8p, 9.15a, 9.20c). Female pleopod 1 with reduced ramus (Fig. 9.21t)��������������������������������������������������������������������������������6 Maxilliped 3 dactylus ovate, distally truncate, with dense distal field of setae (Figs 9.15c, 9.21a–f)����������������������������������7 Maxilliped 3 dactylus linear, with scattered groups of setae (Figs 9.1e, 9.9, 9.16m–q, 9.19u)�����������������������������������������������8 Uropodal exopod oval, without dorsal plate (Fig. 9.15b)����������������������������������������������������������������������������������� Callianopsidae Uropodal exopod with dorsal plate, indicated by secondary row of setae diverging on upper surface from anterior margin (Fig. 9.20d–g)������������������������������������������������������������������������������������������������������������������ Eucalliacidae Uropodal exopod without dorsal plate (Fig. 9.19c, d), or with distal margin interrupted by weak notch (Figs 9.1c, d, 9.19e, f)�������������������������������������������������������������������������������������������������������������������������������������������������������������������9 Uropodal exopod with dorsal plate, secondary row of setae on upper surface branching subdistally from anterior margin (Figs 9.13, 9.18a–j)����������������������������������������������������������������������������������������������������������������������������������������10 Pereopod 5 semichelate (fixed finger closing on proximal part of dactylus), dactylus a twisted plate longer than fixed finger (Fig. 9.19t). Minor cheliped merus with spine on lower margin (Fig. 9.19o–q)������������������������������������������������������������������������������������������������������������������������������������������������������� Ctenochelidae Pereopod 5 chelate (fixed finger closing complete length of dactylus, Fig. 9.1g). Minor cheliped merus lower margin smooth���������������������������������������������������������������������������������������������������������������������������������������� Anacalliacidae Male pleopod 2 absent or reduced. Female pleopod 2 rami styliform, endopod much longer than wide (Fig. 9.12f–h). Maxilliped 1 epipod truncate, without anterior lobe (Fig. 9.8r)�����������������������������������������Callianassidae Male and female pleopod 2 usually present (Fig. 9.18n–q), pleopod 2 rami narrower and with less setation than pleopods 3–5, endopod flattened and 2–5 times as long as wide (Fig. 9.18n–q). Maxilliped 1 epipod with acute anterior lobe lying obliquely along oblique suture of exopod (Fig. 9.16l)������������������������������������������������������������������������������������������������������������������������������������������������� Callichiridae

References de Saint Laurent M (1979) Vers une nouvelle classification des Crustacés Décapodes Reptantia. Bulletin de l’Office Nationale de Pêche de Tunisie 3, 15–31. Dworschak PC (2007) Book review. Sakai, K. 2005. Callianassoidea of the world (Decapoda, Thalassinidea). Crustaceana Monographs 4, i–vi, 285 pp., 44 textfigs. Koninklijke Brill, NV, Leiden, The Netherlands, ISBN 90 04 14211 8. Journal of Crustacean Biology 27, 158–160. doi:10.1651/S-2775.1 Dworschak PC, Poore GCB (2018) More cautionary tales: family, generic and species synonymies of recently published taxa of ghost and mud shrimps (Decapoda, Axiidea and Gebiidea). Zootaxa 4394, 61–76. doi:10.11646/zootaxa.4394.1.3 Dworschak PC, Felder DF, Tudge CC (2012) Chapter 69. Infraorders Axiidea de Saint Laurent, 1979 and Gebiidea de Saint Laurent, 1979 (formerly known collectively as Thalassinidea). In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea. Complementary to the volumes translated from the French of the Traité de Zoologie [founded by P.-P. Grassé]. (Eds Schram FR, von Vaupel Klein JC) pp. 109–219. Brill, Leiden. Poore GCB (1994) A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys to the families and genera. Memoirs of the Museum of Victoria 54, 79–120. doi:10.24199/j.mmv.1994.54.03

Poore GCB (2008) Book review “Sakai, K. 2006. Upogebiidae of the world (Decapoda, Thalassinidea). Crustaceana Monographs 6, i–ix, 185 pp., 23 textfigs. Koninklijke Brill, NV, Leiden, The Netherlands, ISBN-13: 978 90 04 15150 8, hardcover, Euro98/U.S.$132. Journal of Crustacean Biology 28, 422–423. Poore GCB (2015) Rediagnosis of Callianideidae and its genera (Crustacea: Decapoda: Axiidea), and description of a new species of Heardaxius Sakai, 2011. Zootaxa 3995, 229–240. doi:10.11646/zootaxa.3995.1.19 Poore GCB, Collins DJ (2015) Micheleidae (Crustacea: Decapoda: Axiidea): new family and generic synonymies, three new Australian species, and new records. Memoirs of Museum Victoria 73, 95–105. doi:10.24199/j.mmv.2015.73.08 Poore GCB, Dworschak PC (2017) Family, generic and species synonymies of recently published taxa of ghost shrimps (Decapoda, Axiidea, Eucalliacidae and Ctenochelidae): cautionary tales. Zootaxa 4294, 119–125. doi:10.11646/zootaxa.4294.1.6 Poore GCB, Ahyong ST, Bracken-Grissom HD, Chan T-Y, Chu KH, et al. (2014) On stabilising the names of the infraorders of thalassinidean shrimps, Axiidea de Saint Laurent, 1979 and Gebiidea de Saint Laurent, 1979 (Decapoda). Crustaceana 87, 1258–1272. doi:10.1163/15685403-00003354 Poore GCB, Dworschak PC, Robles R, Mantelatto FL, Felder DL (2019) A new classification of Callianassidae and related families (Crustacea: Decapoda: Axiidea) derived from a molecular phylogeny with

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

morphological support. Memoirs of Museum Victoria 78, 73–146. doi:10.24199/j.mmv.2019.78.05 Robles R, Tudge CC, Dworschak PD, Poore GCB, Felder DL (2009) Molecular phylogeny of the Thalassinidea based on nuclear and mitochondrial genes. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 309–326. CRC Press, Boca Raton. Robles R, Dworschak PC, Felder DL, Poore GCB, Mantelatto FL (2020) A molecular phylogeny of Callianassidae and related families (Crustacea: Decapoda: Axiidea) with morphological support. Invertebrate Systematics 34, 113–132. doi:10.1071/IS19021 Sakai K (2005a) Callianassoidea of the world (Decapoda: Thalassinidea). Crustaceana Monographs 4, 1–285. Sakai K (2005b) The diphyletic nature of the infraorder Thalassinidea (Decapoda, Pleocyemata) as derived from the morphology of the gastric mill. Crustaceana 77, 1117–1129. doi:10.1163/1568540042900268 Sakai K (2011) Axioidea of the world and a reconsideration of the Callianassoidea (Decapoda, Thalassinidea, Callianassida). Crustaceana Monographs 13, 1–616. doi:10.1163/9789047424185 Sakai K, Sawada T (2006) The taxa of the infraorders Astacidea, Thalassinidea, Palinura, and Anomura (Decapoda, Pleocyemata) classified by the form of the prepyloric ossicle. Crustaceana 78, 1353–1368. Tsang LM, Lin F-J, Chu KH, Chan T-Y (2008a) Phylogeny of Thalassinidea (Crustacea, Decapoda) inferred from three rDNA sequences: implications for morphological evolution and superfamily classification. Journal of Zoological Systematics and Evolutionary Research 46, 216–223. doi:10.1111/j.1439-0469.2008.00459.x Tsang LM, Ma KY, Ahyong ST, Chan T-Y, Chu KH (2008b) Phylogeny of Decapoda using two nuclear protein-coding genes: Origin and evolution of the Repantia. Molecular Phylogenetics and Evolution 48, 359–368. doi:10.1016/j.ympev.2008.04.009

Anacalliacidae Manning & Felder, 1991 Figure 9.1 The family is known only from the type material of its two species, both in Anacalliax (Poore et al. 2019). The family is diagnosed by the combination of a carinate rostrum and a uropodal exopod with a distal notch. The uropod exopod

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resembles that of species of Paragourretia (Ctenochelidae) but maxilliped 3 lacks an exopod (Fig. 9.1e), and the minor cheliped has fingers longer than the palm. They are assumed to be burrowers in soft sediments like callianassids and similar families. Diagnosis. Rostrum flat, triangular, shorter than eyestalks; carinate (Fig. 9.1a, b). Uropodal exopod with margin divided by notch (Fig. 9.1c, d). Pereopod 5 fingers symmetrical (Fig. 9.1g). Anacalliax de Saint Laurent, 1973 Maximum cl. 30 mm. Intertidal. Tropical Atlantic, Temperate South America (Argentina). 2 species (Manning and Felder 1991). References Manning RB, Felder DL (1991) Revision of the American Callianassidae (Crustacea: Decapoda: Thalassinidea). Proceedings of the Biological Society of Washington 104, 764–792. Poore GCB, Dworschak PC, Robles R, Mantelatto FL, Felder DL (2019) A new classification of Callianassidae and related families (Crustacea: Decapoda: Axiidea) derived from a molecular phylogeny with morphological support. Memoirs of Museum Victoria 78, 73–146. doi:10.24199/j.mmv.2019.78.05

Axiidae Huxley, 1879 sponge shrimps, burrowing lobsters Figures 9.2–9.7, Plates 14, 15 The Axiidae are most diverse in tropical and subtropical seas (Kensley 1996c, 2003; Kensley et  al. 2000; Poore 2018a; Poore and Collins 2009; Sakai 1992; Sakai and Apel 2002) but can be found in temperate environments (Poore

Fig. 9.1.  Anacalliacidae. Dorsal and lateral anterior carapace, eyestalks, antennules, antennae: a, Anacalliax argentinensis (Biffar, 1971). Telson, uropod: c, A. agassizi (Biffar, 1971); d, A. argentinensis. e, maxilliped 3; f, major cheliped; g, pereopod 5 fingers; h, male pleopod 1; i, female pleopod 2.

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and Collins 2009; Sakai 1987) and in the deep sea (Komai 2011; Komai et  al. 2010; Komai et  al. 2002; Poore 2020). Little is known of axiid biology but they are generally weak burrowers or live under stones or in cavities in coral rubble. One species, Axius serratus Stimpson, 1852, builds burrows up to 3 m deep (Pemberton et  al. 1976). Others such as species of Eiconaxius and Montanaxius reside in hexactinellid glass sponges in deep water (Dworschak 2016; Komai and Tsuchida 2012; Kou et al. 2020); species of Spongiaxius are associated with demosponges (Richer de Forges et  al. 2009) and sometimes with hexactinellids (Sakai 1986) The family was extensively reviewed by Sakai and de Saint Laurent (1989). Poore (1994) and Sakai (2011) provided keys to genera known then but the systematics of the family has substantially changed since (Dworschak 2016; Dworschak and Poore 2018; Poore 2017, 2018a, 2020). It is now represented by more than 200 species in 47 genera, many more than when these synthetic works were completed. The Axiidae now incorporates two families, Calocarididae Ortmann, 1891, of eight genera (Kensley 1989) and Eiconaxiidae Sakai, 2011, with one genus (Poore 2017). Calocaridids are monophyletic and, like Eiconaxius, are derived from within Axiidae s.s. according to molecular evidence (Robles et al. 2009; Tsang et al. 2008a; Tsang et al. 2008b). Diagnosis. Rostrum thick, bluntly triangular to prominent, with or without lateral dentition; median carina present as ridge or row of spines extending over gastric region;

posterior margin of carapace with lateral lobes interacting with anterolateral lobes on pleonite 1. Eyestalks usually cylindrical, with terminal subspherical cornea. Maxilla scaphognathite with 1 or few long setae on posterior lobe extending into branchial chamber. Pereopod 3 propodus with distal spiniform setae on lateral face; pereopod 5 simple. Female pleopod 2 rami similar to following pleopods. Implicit generic characters. Gonochoristic. Carapace smooth. Rostrum not especially depressed below level of carapace, about twice as long as eyestalks. Supraocular spines prominent; postcervical carina absent. Pleonites 2–5 smooth, tergum and pleura continuous; pleura 2–5 rounded, without angles. Eyestalk cylindrical, at most about twice as long as cornea diameter; cornea pigmented. Antenna article 2 distal spine directed anteriorly. Palm with few tufts of long setae. Pereopod 2 dactylus upper margin with few separate setae. Pereopods 3 and 4, dactyli narrow, tapering, often with row of spiniform setae on lateral face. Pereopod 5 dactylus tapering from expanded base. Male pleopod 1 absent. Male pleopod 2 endopod with digitiform appendix masculina and appendix interna. Pleopods 2–5 endopod with appendix interna. Uropodal endopod with straight anterior margin ending in tooth, anterior margin serrate or toothed, distal margin convex, smooth, continuous with posterior margin; exopod oval, with transverse suture, anterior margin serrate, article 2 wider than long, without armature. Telson dorsal surface with pairs of teeth.

Key to genera of Axiidae 1. – 2. – 3. – 4. – 5. – 6. –

Pleopod 2 endopod with dominating terminal appendix masculina and digitiform appendix interna. Usually hermaphroditic (gonopores on both coxae 3 and 5 (Fig. 9.5r), pleopods 1 and 2 not sexually dimorphic (Fig. 9.7a, b). Eyes without pigment (Figs 9.3e, f, r, 9.4a, j)��������������������������������������������������������������������������������2 Pleopods 2–5 endopods laminar, similar; pleopod 2 with appendices masculina and interna (if present in male) linear and attached mesially (Fig. 9.7c–j). Gonochoristic (sexes separate, gonopores on either coxae 3 [female] or coxae 5 [male]; pleopods 1 and 2 sexually dimorphic). Eyes usually pigmented����������������������������9 Uropodal exopod without transverse suture (Fig. 9.5k). Sexes separate but some individuals hermaphroditic��������������������������������������������������������������������������������������������������������������������������������������������������������Eucalastacus Uropodal exopod with complete or partial transverse suture (Fig. 9.5f). Hermaphroditic�������������������������������������������������3 Carapace with strong post-cervical carina (Fig. 9.3r)�����������������������������������������������������������������������������������������������������������������4 Carapace without post-cervical carina�����������������������������������������������������������������������������������������������������������������������������������������6 Eyes flattened, mesially contiguous (Fig. 9.3r)�������������������������������������������������������������������������������������������������������������� Calocaris Eyes rounded, not mesially contiguous����������������������������������������������������������������������������������������������������������������������������������������5 Carapace covered with granules. Submedian gastric carina absent (Fig. 9.4j)������������������������������������������������������ Lophaxius Carapace covered with stiff setae. Submedian gastric carina with 4 spines (Fig. 9.4i)�������������������������������������Lophaxiopsis Rostrum slightly lower than dorsal carapace (Fig. 9.3n). Pleopod 2 endopod with elongate-tapering appendix masculina bearing thumb-like basal appendix interna, with mesial row of fine setae (Fig. 9.7e)����������������7 Rostrum significantly depressed below dorsal carapace. Pleopod 2 endopod with broad, triangular appendix masculina bearing thumb-like basal appendix interna, with row of spiniform setae (Fig. 9.7b)�����������������8

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

189

Rostrum without lateral teeth (Fig. 9.3n)������������������������������������������������������������������������������������������������������������������� Calastacus Rostrum with 2 or 3 pairs of lateral teeth (Fig. 9.3o)����������������������������������������������������������������������������������������������� Calaxiopsis Major cheliped palm more than twice as long as high, fingers as long as upper margin (Fig. 9.6f). Scaphocerite much shorter than article 4 (Fig. 9.3f)��������������������������������������������������������������������������������������������� Ambiaxius – Major cheliped palm about as long as high, fingers shorter than upper margin (Fig. 9.6e). Scaphocerite almost as long as article 4 (Fig. 9.3e)������������������������������������������������������������������������������������������Ambiaxiopsis 9. Pleopods 3–5 endopod without appendix interna (Fig. 9.7i)���������������������������������������������������������������������������������������������������10 − Pleopods 3−5 endopod with appendix interna (Fig. 9.7h)�������������������������������������������������������������������������������������������������������14 10. Scaphocerite slender, reaching more than half length of antennal article 4 (Fig. 9.3m). Pleopod 2 with appendix interna (Fig. 9.7d). Pleurobranchs present above pereopods 2–4��������������������������������������������������Bouvieraxius – Scaphocerite plate-like, triangular or bifurcate, barely overlapping base of antennal article 4 (Fig. 9.4b). Pleopod 2 without appendix interna. Pleurobranchs absent above pereopods 2–4������������������������������������11 11. Chelipeds elongate, gaping, with erect tooth on fixed finger (Fig. 9.6n). Pleonal pleura 3–5 truncate, with posteroventral angle���������������������������������������������������������������������������������������������������������������������������������� Eutrichocheles – Chelipeds robust, fingers as long as or shorter than palm. Pleonal pleura 3–5 rounded�������������������������������������������������� 12 12. Submedian carinae present, smooth or tuberculate. Rostrum with 1 or more pairs of lateral teeth (Fig. 9.4r)�������������������������������������������������������������������������������������������������������������������������������������������������������������������� Paraxiopsis – Submedian carinae absent. Rostrum smooth or barely denticulate laterally�����������������������������������������������������������������������13 13. Supraocular teeth present. Scaphocerite with mesial tooth (Fig. 9.4w)�����������������������������������������������������������������Ralumcaris – Supraocular teeth absent (Fig. 9.4m). Scaphocerite triangular����������������������������������������������������������������������������Marianaxius 14. Pereopods 3–4 dactyli ovate, pereopod 5 dactylus spatulate, each with row of short closely-spaced spiniform setae on flexor margin, often with supplementary facial spiniform setae (Fig. 9.6v, y). Median gastric carina smooth, not extending posteriorly over broad triangular midgastric region (Fig. 9.3w, x). Cheliped palm massive, swollen, smooth (Fig. 9.2b)��������������������������������������������������������������������Eiconaxius – Pereopods 3–5 dactyli narrow, tapering, setose or with spiniform setae on flexor and/or lateral face (Fig. 9.6s–u, w, x, z). Median gastric carina smooth or dentate, extending to gastric region. Cheliped palm various������������������������������������������������������������������������������������������������������������������������������������������������������������������������������15 15. Pereopod 2 dactylus with dense field of fine setae (Fig. 9.6r) (1 exception). Chelipeds similar in size, carpus-palm as massive as carapace. Uropodal endopod with truncate distal margin usually defined by 2 spines (Fig. 9.5p)���������������������������������������������������������������������������������������������������������������������������������������������� Spongiaxius – Pereopod 2 dactylus without dense field of fine setae. Chelipeds often dissimilar in size, carpus-palm rarely massive. Uropodal endopod distoposterior margin usually convex���������������������������������������������������������������������16 16. Uropodal exopod without transverse suture (Fig. 9.5g)�����������������������������������������������������������������������������������������������������������17 − Uropodal exopod with transverse suture (Fig. 9.5a–f, h–j, l–o)����������������������������������������������������������������������������������������������19 17. Eyestalks flat, colourless, callianassoid-like (Fig. 9.3g). Chelipeds subequal, similar (Fig. 9.6h)���������� Anophthalmaxius – Eyestalks cylindrical, cornea pigmented. Chelipeds unequal, minor cheliped elongate (Fig. 9.2c)����������������������������������18 18. Rostrum significantly depressed below near vertical anteromedial margin (Fig. 9.4x)������������������������������������� Scytoleptus – Rostrum level with or slightly below gastric region of carapace; gastric region horizontal or gently sloped (Fig. 9.2c)������������������������������������������������������������������������������������������������������������������������������������ Parascytoleptus 19. Uropodal exopod anterior margin convex, with 4–12 prominent teeth; endopod anterior margin with prominent proximal shoulder. Telson shorter than uropodal endopod. Rostrum not longer than eyestalk��������������� 20 – Uropodal exopod anterior margin convex or straight, smooth or denticulate; endopod oval or with straight anterior margin. Telson as long as uropodal endopod. Rostrum usually longer than eyestalk��������������������21 20. Scaphocerite deep, curved, longer than antenna article 4, overreaching expanded spine on article 2 (Fig. 9.3b). Uropodal exopod article 2 longer than wide, with distal spine (Fig. 9.5b)������������������������������� Alienaxiopsis – Scaphocerite horn-shaped, much shorter than antenna article 4, as long as short spine on article 2 (Fig. 9.3c). Uropodal exopod article 2 about as wide as long, with row of 5 short even teeth (Fig. 9.5c)������������������������������Allaxiopsis 21. Pleonal pleura 1–5 attenuated ventrally (Fig. 9.2d)������������������������������������������������������������������������������������������������������������������ 22 7. – 8.

190

Marine Decapod Crustacea

– Pleonal pleura 1–5 rounded or truncate ventrally (e.g. Fig. 9.2a, c)�������������������������������������������������������������������������������������� 23 22. Cheliped carpus and propodus upper margin without spines (Fig. 9.2d). Uropodal endopod oval (Fig. 9.5o). Male pleopod 2 without appendix masculina���������������������������������������������������������������������������������� Pilbaraxius – Cheliped carpus and propodus upper margin with erect spines (Fig. 9.6j). Uropodal endopod with straight anterior margin, with distal tooth (Fig. 9.5i). Male pleopod 2 with appendix masculina���������������������������������Calaxius 23. Carapace without submedian carinae (Figs 9.3u, 9.4g, p, u)�������������������������������������������������������������������������������������������������� 24 – Carapace with submedian carinae (e.g. Fig. 9.3a, d, i)������������������������������������������������������������������������������������������������������������� 28 24. Carapace lateral carinae convex, with 4 or 5 teeth including supraocular tooth (almost appearing as ring of teeth) (Fig. 9.3u). Telson strongly tapering from near base to rounded posterior half (Fig. 9.5j)������������ Coralaxius – Carapace lateral carinae diverging posteriorly, without teeth (Fig. 9.4g, p, u). Telson tapering to rounded apex (Fig. 9.5m)������������������������������������������������������������������������������������������������������������������������������������������������������������������������ 25 25. Rostrum level with dorsal carapace (Fig. 9.4f)������������������������������������������������������������������������������������������������������� Levantocaris – Rostrum level depressed below dorsal carapace (Fig. 9.4o)���������������������������������������������������������������������������������������������������� 26 26. Carapace median carina with 2 teeth (Fig. 9.4p)��������������������������������������������������������������������������������������������������� Montanaxius – Carapace median carina smooth������������������������������������������������������������������������������������������������������������������������������������������������ 27 27. Rostrum broadly triangular, with rim of erect-oblique teeth. Lateral carina absent. Cornea flattened, not pigmented (Fig. 9.4s)����������������������������������������������������������������������������������������������������������������������������������������������������� Paraxius – Rostrum acutely triangular, margin with obsolete teeth, running on to lateral carina. Cornea globular, pigmented (Fig. 9.4u)�������������������������������������������������������������������������������������������������������������������������������������������������� Planaxius 28. Rostrum one-third carapace length, styliform, with obsolete lateral teeth. Eyestalk at least 4 times cornea diameter������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 29 – Rostrum less than one-third carapace length, triangular, with obsolete or prominent lateral teeth. Eyestalk rarely more than twice cornea diameter������������������������������������������������������������������������������������������������������������������������������ 30 29. Eyestalk one-third length of rostrum (Fig. 9.3h). Chelipeds symmetrical, flat, oval, carpus–dactylus with erect spines on upper and lower margins (Fig. 9.6g)��������������������������������������������������������������������������������������� Australocaris – Eyestalk as long as rostrum (Fig. 9.4q). Cheliped asymmetrical, linear, carpus–dactylus with short teeth or oblique spines on upper margin (Fig. 9.6q)��������������������������������������������������������������������������������������������������Oxyrhynchaxius 30. Rostrum strongly depressed below median gastric carina. Gastric carina with only supraocular spine. Median carina with small anterior median tooth above steeply sloping ridge (Fig. 9.3v)������������������������ Dorphinaxius – Rostrum level with or slightly below median gastric carina. Gastric carina more or less dentate. Median carina usually with denticles or teeth������������������������������������������������������������������������������������������������������������������������������������31 31. Carapace lateral gastric carina unarmed (Fig. 9.3j, k, 9.4c, e)�������������������������������������������������������������������������������������������������32 – Carapace lateral gastric carina with denticles or teeth (e.g. Fig. 9.3l, t, 9.4k)����������������������������������������������������������������������� 36 32. Cheliped palm with 2 tufts of long plumose setae proximally on upper margin (Fig. 9.6i). Eyestalk as long as rostrum (Fig. 9.3j)�������������������������������������������������������������������������������������������������������������������������������������������������Axiorygma – Cheliped palm without tufts of long plumose setae proximally on upper margin. Eyestalk shorter than rostrum������������� 33 33. Carapace median gastric carina smooth (Fig. 9.3k, 9.4e)�������������������������������������������������������������������������������������������������������� 34 – Carapace median gastric carina denticulate����������������������������������������������������������������������������������������������������������������������������� 36 34. Carapace submedian carina short, reaching one-third length of gastric region. Median carina unarmed. Rostrum apex rounded (Fig. 9.3k)����������������������������������������������������������������������������������������������������������������������������������� Axius – Carapace submedian carina short, reaching two-thirds length of gastric region. Median carina with 2 teeth. Rostrum apex acute (Fig. 9.4e)���������������������������������������������������������������������������������������������������������������Leonardsaxius 35. Telson without dorsal teeth. Uropodal rami each without row of teeth on ribs (Fig. 9.5l). Scaphocerite straight (Fig. 9.4c)��������������������������������������������������������������������������������������������������������������������������������������������������� Formosaxius – Telson with 4 pairs of dorsal teeth. Uropodal rami each with row of teeth on ribs (Fig. 9.5h). Scaphocerite short, comma-shaped (Fig. 9.3l)�������������������������������������������������������������������������������������������������������������������������������Boasaxius 36. Cheliped usually flattened, upper margin of propodus, often carpus–dactylus denticulate, dentate or spinose. Lateral gastric carina usually with 1 prominent tooth followed by 1 or more smaller teeth. Telson anterolateral lobe with posteriorly directed tooth���������������������������������������������������������������������������������������������������37

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

– 37. – 38. – 39. – 40. – 41. – 42. – 43. –

191

Cheliped cylindrical, upper margin of carpus–dactylus smooth (denticulate in some Calocarides). Lateral gastric carina with several similar small teeth. Telson anterolateral lobe without posteriorly directed tooth������������� 42 Scaphocerite with basal mesial tooth (Fig. 9.4d)���������������������������������������������������������������������������������������������������� Guyanacaris Scaphocerite without basal mesial tooth (Fig. 9.4k, t)������������������������������������������������������������������������������������������������������������� 38 Carapace supraocular spine absent; median gastric carina with 1 prominent tooth; lateral gastric carina with 1 prominent tooth. Eyestalk flat, cornea weakly pigmented (Fig. 9.3s, t)��������������������������������������������Carolinaxius Carapace supraocular spine present; median gastric carina denticulate; median gastric carina with 1 prominent tooth, usually followed by other teeth. Eyestalk cylindrical, cornea fully pigmented������������������������������ 39 Carapace lateral gastric carina with single prominent tooth set about one-third way along gastric region, well back from supraocular tooth; supraocular spine prominent (Figs 9.2a, 9.3a, d, 9.4k, t)��������������������������������������� 40 Carapace lateral gastric carina comprising several similar small teeth; supraocular spine undifferentiated from gastric teeth (Figs 9.3i, q, 9.4h, n, v)����������������������������������������������������������������������������������������������������������������������������� 43 Carapace lateral gastric carina with first tooth set about one-third to half way along gastric region from supraocular tooth, smaller posterior teeth. Antennal article 2 distal spine directed anteriorly (Fig. 9.4k, t)�������������� 41 Carapace lateral gastric carina with first prominent tooth set just behind anterior margin of carapace and followed by 1 or more teeth converging posteriorly. Antennal article 2 distal spine directed anteriorly (Fig. 9.3a, d)������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 42 Cheliped carpus–propodus upper margin with row of denticles, teeth or spines; dactylus unarmed (Fig. 9.6p)����������������������������������������������������������������������������������������������������������������������������������������������������������������������Manaxius Cheliped merus–dactylus or carpus–dactylus upper margin with teeth or spines��������������������������������������� Pillsburyaxius Carapace postcervical carina absent (Fig. 9.3a). Cheliped merus-dactylus upper margin with row of spines (Fig. 9.6a, b). Uropodal exopod oval (Fig. 9.5a)������������������������������������������������������������������������������������������������� Acanthaxius Carapace postcervical carina present (Fig. 9.3d). Cheliped carpus–dactylus upper margins with short teeth and tubercles (Fig. 9.6d). Uropodal exopod with straight posterior margin, longitudinal ridge ending as spine adjacent to article 2 (Fig. 9.5e)������������������������������������������������������������������������������������������������������������������Amakusaxius Scaphocerite short, comma-shaped (Fig. 9.4h)������������������������������������������������������������������������������������������������������������������������� 44 Scaphocerite as long as article 4, straight���������������������������������������������������������������������������������������������������������������������������������� 45

Fig. 9.2.  Axiidae. a, Acanthaxius garawa Poore & Collins, 2009; b, Eiconaxius mallacoota Poore & Collins, 2009 (major chela only); c, Parascytoleptus papua Poore & Collins, 2010 (with minor chela only); d, Pilbaraxius kariyarra Poore & Collins, 2009.

192

Marine Decapod Crustacea

Fig. 9.3.  Axiidae. Anterior carapace, eyestalk, antennule, antenna: a, Acanthaxius; b, Alienaxiopsis; c, Allaxiopsis; d, Amakusaxius; e, Ambiaxiopsis; f, Ambiaxius; g, Anophthalmaxius; h, Australocaris; i, Axiopsis; j, Axiorygma; k, Axius; l, Boasaxius; m, Bouvieraxius; n, Calastacus; o, Calaxiopsis; p, Calaxius; q, Calocarides; r, Calocaris; s, t, Carolinaxius; u, Coralaxius; v, Dorphinaxius; w, x, Eiconaxius.

44. Major cheliped palm swollen, 1.4 times as long as wide, with scattered simple setae (Fig. 9.6c). Carapace submedian gastric carina double, rows of spinules��������������������������������������������������������������������������������������������������Allaxius – Major cheliped palm subrectangular, 1.1 times as long as wide, with clusters of plumose setae (Fig. 9.6o). Carapace submedian gastric carina single row of blunt teeth (Fig. 9.4h)�����������������������������������������������������������Litoraxius 45. Pleonites 2–5 with 5 longitudinal dorsal ridges and lateral carina above pleura (Fig. 9.5q). Eyestalk ~4 times cornea diameter (Fig. 9.4v)��������������������������������������������������������������������������������������������������������������� Pseudoaxiopsis – Pleonites 2–5 with smooth dorsally, with or without lateral carina above pleura. Eyestalk about twice cornea diameter������������������������������������������������������������������������������������������������������������������������������������������������������������������������ 46 46. Scaphocerite without basal mesial tooth. Carapace submedian gastric carina straight (Fig. 9.3q). Cheliped usually with denticles along upper margin and faces (Fig. 9.6k)����������������������������������������������������� Calocarides – Scaphocerite with (rarely without) basal mesial tooth. Carapace submedian gastric carina converging anteriorly or recurved, often with intermediate tubercles (Figs 9.3i, 9.4n). Cheliped smooth or with scales along upper margin������������������������������������������������������������������������������������������������������������������������������������������������������������������ 47

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

193

Fig. 9.4.  Axiidae. Anterior carapace, eyestalk, antennule, antenna: a, Eucalastacus; b, Eutrichocheles; c, Formosaxius; d, Guyanacaris; e, Leonardsaxius; f, g, Levantocaris; h, Litoraxius; i, Lophaxiopsis; j, Lophaxius; k, Manaxius; l, m, Marianaxius; n, Michelaxiopsis; o, p, Montanaxius; q, Oxyrhynchaxius; r, Paraxiopsis; s, Paraxius; t, Pillsburyaxius; u, Planaxius; v, Pseudoaxiopsis; w, Ralumcaris; x, Scytoleptus; y, z, Spongiaxius.

47. –

Carapace lateral gastric carina unarmed; submedian gastric carina a hair-pin shaped double ridge of tubercles; median gastric carina with few spines and/or beads (Fig. 9.4n). Male pleopod 2 without appendix masculina (Fig. 9.7g)�������������������������������������������������������������������������������������������������������������������������Michelaxiopsis Carapace lateral gastric carina with several teeth; submedian gastric carina of numerous teeth; median gastric carina with several teeth (Fig. 9.3i). Male pleopod 2 with appendix masculina (Fig. 9.7c)�������������������Axiopsis Acanthaxius Sakai & de Saint Laurent, 1989

Diagnosis. Rostrum shorter than eyestalks or about as long as eyestalks, narrowly triangular or triangular, with prominent,

spaced lateral spines. Carapace smooth, or covered with granules; median carina denticulate; submedian gastric carina comprising row of carinate teeth; lateral gastric carina with first prominent tooth set just behind anterior margin of carapace and followed by

194

Marine Decapod Crustacea

Fig. 9.5.  Axiidae. Telson, uropod: a, Acanthaxius; b, Alienaxiopsis; c, Allaxiopsis; d, Allaxius; e, Amakusaxius; f, Ambiaxius; g, Anophthalmaxius; h, Boasaxius; i, Calaxius; j, Coralaxius; k, Eucalastacus; l, Formosaxius; m, Levantocaris; n, Manaxius; o, Pilbaraxius; p, Spongiaxius. Pleon, telson, uropod: q, Pseudoaxiopsis. Thoracic sternum, left pereopods 3–5 coxae: r, Ambiaxius.

1 or more teeth converging posteriorly. Pleonites 2–5 smooth, tergum and pleura continuous, or with longitudinal groove above pleura; pleura 2–5 truncate, with posteroventral angle. Antenna article 2 distal spine directed anteromesially; scaphocerite simple. Major cheliped elongate, merus-dactylus upper margin with row of spines, palm densely setose; fingers longer than palm. Minor cheliped elongate, fingers 2–3 times palm length. Maximum cl. 21 mm (Pl. 14a). Shelf, slope (49–450 m). Temperate Northern W Pacific, Central Indo-Pacific. 9 species (Poore 2020: discussion of genus; Sakai 2011: key to 17 species, some now in other genera).

Alienaxiopsis Sakai, 2011 Diagnosis. Rostrum about as long as eyestalks, broadly triangular (as long as wide at base), with 1 or more pairs of lateral teeth. Carapace median carina with 1 tooth and broad triangular plate posteriorly; submedian gastric carina indefinite, with 2 blunt teeth; lateral gastric carina indefinite, with 1 or 2 teeth. Scaphocerite deep, curved, longer than antenna article 4, overreaching expanded spine on article 2. Major cheliped cylindrical, upper margins smooth; fingers shorter than palm; fixed finger blade with subdistal blunt step. Minor cheliped half width of major, fingers little longer than palm. Uropodal endopod subrectangular, anterior margin with prominent proximal shoulder, 1 or 2

strong teeth at midpoint, distal margin truncate, with 2 anterior teeth and 2 laterally; exopod strongly convex anteriorly, straight posteriorly, anterior margin with 4 strong teeth, article 2 longer than wide, oval, with 1 strong distal spine. Maximum cl. 4.7 mm (Pl. 14b). Subtidal (3–20 m). Central Indo-Pacific. 1 species (Poore 2018a: species synonymy; Sakai 2011).

Allaxiopsis Sakai, 2011 Diagnosis. Rostrum about as long as eyestalks, rectangular, with trifid apex, with 1 pair of lateral teeth. Carapace median carina indefinite, with 2–4 blunt teeth; submedian gastric carina indefinite, with irregular widely-spaced blunt teeth; lateral gastric carina indefinite, with 1 or 2 teeth. Major cheliped cylindrical, propodus upper margin with erect spines; fingers shorter than palm. Minor cheliped half width of major. Uropodal endopod subrectangular, anterior margin with prominent proximal shoulder, posterodistal margin convex, with row of 8–9 short teeth; exopod strongly convex anteriorly, straight posteriorly, anterior margin with 7–10 strong teeth, article 2 about as wide as long, with row of 5 short even teeth. Maximum cl. 9.3 mm (Pl. 14c). Subtidal (3–23 m). Central Indo-Pacific. 2 species (Poore 2018a: illustrations; Sakai 2011).

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195

Fig. 9.6.  Axiidae. Pereopod 1. a, b, Acanthaxius (major, minor); c, Allaxius; d, Amakusaxius; e, Ambiaxiopsis; f, Ambiaxius; g, Australocaris; h, Anophthalmaxius; i, Axiorygma; j, Calaxius; k, Calocarides; l, m, Eiconaxius; (major, minor); n, Eutrichocheles; o, Litoraxius; p, Manaxius; q, Oxyrhynchaxius. Pereopod 2: r, Spongiaxius. Pereopod 3 propodus, dactylus: s, Calaxius; t, Pillsburyaxius. Pereopod 4 dactylus: u, Bouvieraxius; v, Eiconaxius; w, Manaxius; x, Spongiaxius. Pereopod 5 dactylus: y, Eiconaxius; z, Manaxius.

Allaxius Sakai & de Saint Laurent, 1989

Amakusaxius Sakai, 2011

Diagnosis. Rostrum about as long as eyestalks, triangular, with 2 or 3 pairs of lateral teeth. Carapace median carina denticulate or smooth; submedian gastric carina double, rows of spinules; lateral gastric carina with several small teeth. Scaphocerite short, comma-shaped in lateral view. Major cheliped palm swollen, upper margins smooth; fingers shorter than palm. Minor cheliped similar to but half as wide as major cheliped. Uropodal endopod anterior margin smooth, posterodistal margin convex, with row of 8–9 short teeth; exopod article 2 wider than long, with few distal spines. Maximum cl. 13 mm. Subtidal, shelf. Western Indo-Pacific. 1 species (Sakai 2011: rediagnosis, confined to one species).

Diagnosis. Rostrum shorter than eyestalks, triangular, with 1 or more pairs of lateral teeth. Carapace median carina denticulate; submedian gastric carina comprising row of carinate teeth; lateral gastric carina with first prominent tooth set just behind anterior margin of carapace and followed by 1 or more teeth converging posteriorly; postcervical carina present over whole postcervical length. Antenna article 2 distal spine directed anteromesially; scaphocerite simple. Major cheliped elongate, carpus–dactylus upper margins with short teeth and tubercles, palm densely setose; fingers longer than palm. Minor cheliped elongate, carpusdactylus upper margin with row of spines, fingers twice palm length. Uropodal exopod with straight posterior margin,

196

Marine Decapod Crustacea

Fig. 9.7.  Axiidae. Male pleopod 1: a, Ambiaxius. Male pleopod 2: b, Ambiaxius; c, Axiopsis; d, Bouvieraxius; e, Calastacus; f, Eiconaxius. Female pleopod 2: g, Michelaxiopsis. Pleopod 3: h, Axiopsis; i, Bouvieraxius; j, Calocaris. k, Maxilla.

longitudinal ridge ending as spine adjacent to article 2. Maximum cl. 15.6 mm. Shelf (20–40 m). Temperate Northern W Pacific, Central IndoPacific. 1 species (Sakai 2011).

Ambiaxiopsis Komai, 2011 Diagnosis. Hermaphroditic. Rostrum significantly depressed below level of carapace, 3 or more times as long as short eyestalks, styliform. Carapace median carina indistinct; submedian gastric carina absent; lateral gastric carina unarmed. Eyestalk short, flattened, about as long as wide, fused to carapace; cornea unpigmented. Scaphocerite simple, straight. Major cheliped palm ovoid, upper margins smooth; fixed finger blade with erect tooth near base. Minor cheliped palm slightly narrower and more dilating than major. Pleopod 1 article 2 subtriangular, one-third length of first, folded obliquely, posterior part distally lobed, anterior part smaller, narrower, with distal margin oblique, with short appendix interna. Male pleopod 2 endopod with broad, triangular appendix masculina bearing thumb-like basal appendix interna, with row of spiniform setae. Uropodal endopod oval. Maximum cl. 6.6 mm. Slope (247–269 m). Temperate Northern W Pacific. 1 species (Komai 2011).

Ambiaxius Sakai & de Saint Laurent, 1989 Diagnosis. Hermaphroditic. Rostrum significantly depressed below level of carapace, 2–3 or more times as long as short eyestalks, narrowly triangular, tapering quickly to single elongate

spine (rarely with lateral pair of spines) or styliform. Carapace median carina indistinct; submedian gastric carina absent; lateral gastric carina unarmed. Eyestalk short, flattened, about as long as wide, fused to carapace; cornea unpigmented. Scaphocerite simple, much shorter than article 4. Major cheliped palm dilating, twice as long as wide, merus, propodus upper margins with subdistal tooth; fingers as long as palm. Minor cheliped similar to major cheliped. Pleopod 1 article 2 subtriangular, one-third length of first, folded obliquely, posterior part distally lobed, anterior part smaller, narrower, with distal margin oblique, with short appendix interna. Male pleopod 2 endopod with broad, triangular appendix masculina bearing thumb-like basal appendix interna, with row of spiniform setae. Uropodal endopod oval; exopod with transverse or partial suture. Maximum cl. 17 mm. Slope (400–1275 m). Temperate Northern W Pacific, Tropical W Atlantic, Western and Central Indo-Pacific. 9 species (Komai et  al. 2010: composition of genus; Poore 2020: composition of genus, 4 species; Sakai and de Saint Laurent 1989).

Anophthalmaxius De Man, 1905 Diagnosis. Carapace supraocular spines absent; median carina smooth; submedian gastric carina absent; lateral gastric carina unarmed. Eyestalk flattened, about as long as wide, distally emarginate, fused to carapace; cornea unpigmented. Scaphocerite. Major cheliped cylindrical, upper margins smooth; fingers shorter than palm; fixed finger blade with erect tooth near base. Uropodal endopod oval; exopod without transverse suture. Maximum cl. 6 mm. Slope (1788 m). Central Indo-Pacific. 1 species (Sakai 2011: rediagnosis; Sakai and de Saint Laurent 1989: rediagnosis).

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

Australocaris Poore & Collins, 2009 Diagnosis. Rostrum one-third carapace length, narrowly triangular, barely denticulate laterally. Carapace supraocular spines indistinguishable from others on rostrum; median and submedian gastric carinae smooth; lateral gastric carina unarmed. Eyestalk cylindrical, elongate, ~3 times as long as cornea diameter; cornea unpigmented. Scaphocerite elongate, with basal mesial spine and 4 spines on lower margin. Major cheliped propodus laterally flattened, broad, carpus-dactylus upper and lower margins carinate, with strong spines; fingers longer than palm. Minor cheliped similar to major cheliped. Male pleopod 1 of 1 simple article. Male pleopod 2 endopod with appendix interna, without appendix masculina. Uropodal endopod anterior margin smooth, distal margin continuous with anterior and posterior margins. Maximum cl. 27 mm. Slope (397–423 m). Temperate SW Australia. 1 species (Poore and Collins 2009).

Axiopsis Borradaile, 1903 Diagnosis. Rostrum triangular, with prominent, spaced lateral spines. Carapace supraocular spines indistinguishable from others on rostrum; median carina with spines or teeth; submedian gastric carina spinose, converging anteriorly, often with spinules between median and submedian carinae; lateral gastric carina with several small teeth. Scaphocerite simple (rare), with basal mesial spine. Major cheliped cylindrical, upper margins smooth; fingers shorter than palm. Minor cheliped half width of major, fingers shorter than palm. Maximum cl. 22 mm. Intertidal–slope (0–275 m). Cosmopolitan except polar seas. 8 species  – species differentiation is difficult and some species are apparently widespread (Kensley 2003; Ngoc-Ho 2005; Poore 2018a; Sakai 2011). Molecular evidence suggests that some nominal species treated as synonyms of A. serratifrons (A. Milne-Edwards, 1873) may need to be revived.

Axiorygma Kensley & Simmons, 1988 Diagnosis. Rostrum about as long as eyestalks, narrowly triangular, barely denticulate laterally. Carapace supraocular spines indistinguishable from others on rostrum; median and submedian gastric carinae smooth; lateral gastric carina unarmed. Eyestalk cylindrical, elongate, ~4 times as long as cornea diameter. Scaphocerite simple. Major cheliped cylindrical, upper margins smooth, palm with 2 tufts of long plumose setae proximally on upper margin; fingers shorter than palm. Minor cheliped similar to but slightly narrower than major cheliped. Male pleopod 2 endopod with appendix interna, without appendix masculina. Maximum cl. 7.5 mm. Shelf (52–59 m). Tropical Atlantic (Gulf of Mexico). 1 species (Kensley and Simmons 1988).

Axius Leach, 1815 Diagnosis. Rostrum triangular, barely denticulate laterally. Carapace supraocular spines indistinguishable from others on rostrum; median carina smooth; submedian gastric carina smooth (short, obsolete); lateral gastric carina unarmed. Scaphocerite

197

simple. Major cheliped cylindrical, upper margins smooth; fingers shorter than palm. Minor cheliped similar to but slightly narrower than major cheliped. Male pleopod 1 article 2 oval, with distal lobe-like appendix interna. Uropodal endopod anterior margin smooth; exopod with transverse or partial suture. Maximum cl. 32 mm. Subtidal–slope (18–260 m). Temperate Northern Atlantic. 3 species (Kensley 2001: W Atlantic species; Ngoc-Ho 2003: E Atlantic, Mediterranean species, ecology).

Boasaxius Sakai, 2011 Diagnosis. Rostrum triangular, with smooth median carina, with 2 or 3 pairs of lateral teeth. Carapace supraocular spines indistinguishable from others on rostrum; median carina denticulate; submedian gastric carina comprising row of carinate teeth; lateral gastric carina unarmed. Scaphocerite short, comma-shaped in lateral view. Major cheliped robust, propodus upper margin tuberculate, with subdistal tooth; fingers as long as palm. Minor cheliped half as wide as major cheliped; palm upper margin with 5 erect spines. Male pleopod 1 articles 1 and 2 fused, distally with fingerlike projection. Uropodal endopod anterior margin with tooth at midpoint, 2 distally; upper face with row of teeth on rib; exopod anterior margin with 4 teeth, upper face with row of teeth on rib, article 2 wider than long, with 2 distal spines. Maximum cl. 80 mm. Subtidal–slope (5–300 m). Temperate Northern Pacific, Central Indo-Pacific. 1 species (Marin 2015: redescription; Sakai 2011: diagnosis).

Bouvieraxius Sakai & de Saint Laurent, 1989 Diagnosis. Hermaphroditic. Rostrum narrowly triangular, with 1–4 pairs of lateral teeth. Carapace median carina smooth; submedian gastric carina with 3–5 well spaced teeth; lateral gastric carina unarmed. Scaphocerite simple. Major cheliped cylindrical, propodus upper margin tuberculate; fingers shorter than palm. Minor cheliped similar to but slightly narrower than major cheliped. Male pleopod 1 articles 1 and 2 fused, apex bilobed, mesial margin with linear row of hooks forming appendix interna. Pleopods 3–5 endopod without appendix interna. Maximum cl. 13 mm (Pl. 14d). Intertidal–slope (0–424 m). Tropical W Atlantic, Indo-West Pacific. 4 species (Poore and Collins 2009: rediagnosis).

Calastacus Faxon, 1893 Diagnosis. Hermaphroditic. Rostrum 2–3 or more times as long as short eyestalks, styliform. Carapace median carina indistinct; submedian gastric carina absent; lateral gastric carina unarmed except for prominent supraocular. Eyestalk short, flattened, about as long as wide, fused to carapace; cornea unpigmented. Scaphocerite simple. Major cheliped palm dilating, merus, propodus upper margins with subdistal tooth, sometimes 0–3 on propodus; fingers as long as palm. Minor cheliped similar to but slightly narrower than major cheliped. Pleopod 1 article 2 subtriangular, one-third length of first, folded obliquely, posterior part distally lobed, anterior part smaller, narrower, with distal margin oblique, with short appendix interna.

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Marine Decapod Crustacea

Male pleopod 2 endopod with elongate-tapering appendix masculina bearing thumb-like basal appendix interna, with mesial row of fine setae. Uropodal endopod oval. Maximum cl. 7 mm. Slope (350–1400 m). Temperate Northern and Tropical A ­ tlantic, Central Indo-Pacific, Tropical Eastern Pacific, Temperate SW ­Australia. 11 species (Ngoc-Ho 2003: rediagnosis; Poore 2020: species description; Sakai 2011: rediagnosis, key to 8 species).

Calaxiopsis Sakai & de Saint Laurent, 1989 Diagnosis. Hermaphroditic. Rostrum 3 or more times as long as short eyestalks, with 2 or 3 pairs of lateral teeth. Carapace median carina smooth; submedian gastric carina converging mesially on to medial carina; lateral gastric carina unarmed except for prominent supraocular. Eyestalk short, flattened, about as long as wide, fused to carapace; cornea unpigmented. Scaphocerite simple. Major cheliped palm dilating, merus, propodus upper margins with subdistal tooth; fingers longer than palm. Minor cheliped similar to but slightly narrower than major cheliped. Pleopod 1 article 2 subtriangular, one-third length of first, folded obliquely, posterior part distally lobed, anterior part smaller, narrower, with distal margin oblique, with short appendix interna. Male pleopod 2 endopod with elongate-tapering appendix masculina bearing thumb-like basal appendix interna, with mesial row of fine setae. Uropodal endopod oval; exopod with transverse or partial suture. Maximum cl. 9 mm. Shelf, slope (80–470 m). Temperate Northern W Pacific, Tropical E Atlantic. 2 species (Komai 2000; Sakai and de Saint Laurent 1989). Nipponcalaxiopsis Sakai, 2011 is indistinguishable from Calaxiopsis and it is treated as a synonym here. Ezaxius de Matos-Pita & Ramil, 2015 is a junior synonym and E. ferachevali de Matos-Pita & Ramil, 2015 is a probable junior synonym of C. serrata Sakai & de Saint Laurent, 1989.

Calaxius Sakai & de Saint Laurent, 1989 Diagnosis. Carapace smooth or tuberculate. Rostrum 2–3 or more times as long as short eyestalks, narrowly triangular, with 1 or more pairs of lateral teeth. Carapace median carina denticulate; submedian gastric carina comprising row of carinate teeth, or with 2 teeth; lateral gastric carina with single prominent tooth set well back (about third way along gastric region) from supraocular tooth; postcervical carina absent, or present as rounded ridge over short posterior section. Pleonites 2–5 with longitudinal groove above pleura; pleura 2–5 attenuated to point. Scaphocerite simple. Major cheliped elongate, carpus-propodus upper margin with row of denticles, teeth or spines, palm covered with numerous tufts of long setae; fingers longer than palm; fixed finger blade with 2–4 erect triangular teeth, diminishing distally. Minor cheliped similar to but slightly narrower than major cheliped. Maximum cl. 26 mm (Pl. 14e, f). Shelf, slope (82–514 m). Tropical W Atlantic, Central IndoPacific. 4 species (Poore 2020: rediagnosis).

Calocarides Wollebaek, 1909 Diagnosis. Rostrum triangular, with prominent, spaced lateral spines. Carapace supraocular spines indistinguishable from others on rostrum; median carina with 1 or more teeth; submedian gastric

carina comprising row of carinate teeth; lateral gastric carina with 1 or few well spaced teeth (in addition to anterior defining supraocular tooth); postcervical carina absent or present. Scaphocerite simple. Major cheliped cylindrical, upper margins denticulate or dentate; fingers shorter than palm; fixed finger blade with bladelike tooth near base. Minor cheliped half width of major, fingers shorter than palm. Uropodal endopod anterior margin smooth or serrate or toothed, distal margin convex, smooth, continuous with posterior margin or distal margin convex, continuous with anterior and posterior margins. Maximum cl. 27 mm. Shelf–bathyal (50–2036 m). Widespread. 16 species (Kensley 1996d; Ngoc-Ho 2003). Although the genus is variable, Balssaxius Sakai, 2011 is treated as a junior synonym.

Calocaris Bell, 1846 Diagnosis. Hermaphroditic. Rostrum 3 or more times as long as short eyestalks, triangular, with prominent, spaced lateral spines. Carapace median carina smooth; submedian gastric carina absent; lateral gastric carina with 1 or few well-spaced teeth (in addition to anterior defining supraocular tooth); postcervical carina present over whole postcervical length. Eyestalk short, flattened, about as long as wide, fused to carapace; cornea unpigmented. Scaphocerite simple, much shorter than article 4. Major cheliped palm dilating, merus, propodus upper margins with subdistal tooth; fingers 3 times as long as palm, gaping; fixed finger blade with proximal triangular tooth. Minor cheliped similar to major cheliped. Pleopod 1 with article 2 as long as article 1, triangular, with appendix interna represented by small hooks. Male pleopod 2 endopod with axial elongate-tapering appendix masculina bearing free, elongate basal appendix interna, setose distally. Uropodal endopod oval. Maximum cl. 18 mm. Shelf, slope (84–1272 m). Temperate Northern and Tropical Atlantic, Temperate Southern Africa, Temperate Australasia. 6 species (Kensley 1996b: description; Ngoc-Ho 2003: rediagnosis, redescription, biology; Sakai 2011: rediagnosis, key to 4 species). Calocarisopsis Sakai, 2011 is treated as a junior synonym.

Carolinaxius Kou, Poore & Li, 2021 Diagnosis. Rostrum one-quarter carapace length, narrowly triangular, with few small lateral teeth. Carapace supraocular spines absent; median carina with 1 tooth; submedian gastric carina converging posteriorly, with 4 or 5 teeth; lateral gastric carina with 1 prominent tooth. Eyestalk reaching one-third length of rostrum, with acute distomesial dorsal tooth; cornea weakly pigmented. Scaphocerite simple. Male pleopod 1 article 2 as long as article 1, triangular, tapering to acute hooked apex bearing subapical seta, mesial margin with blunt appendix interna bearing field of hooks and prominent acute triangular process. Maximum cl. 6 mm. Slope (796–1510 m). Central Indo-Pacific. 1 species (Kou et al. 2021).

Coralaxius Kensley & Gore, 1981 Diagnosis. Rostrum shorter than eyestalks, simple, spike-like, with 1 or more pairs of lateral teeth or not dentate laterally.

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

Carapace median carina with spines or teeth, or smooth; submedian gastric carina absent; lateral gastric carina convex, with 4 or 5 teeth including supraocular tooth (almost appearing as ring of teeth). Pleura 2–5 truncate, with posteroventral angle. Scaphocerite scale-like, spinose. Major cheliped cylindrical, carpus-dactylus upper and lower margins with row of blunt teeth; fingers shorter than palm. Minor cheliped propodus narrower than major, fixed finger with 2 erect spines on cutting edge. Male pleopod 1 article 2 oval, with distal lobe-like appendix interna. Male pleopod 2 endopod with digitiform appendix interna, with bilobed broad appendix masculina. Maximum cl. 4 mm. Subtidal–slope (5–230 m). Tropical W Atlantic, Central IndoPacific, Tropical Eastern Pacific. 3 species (Kensley 1994; Kensley and Gore 1981). Meinertaxius Sakai, 2011 is treated as a junior synonym.

Dorphinaxius Sakai & de Saint Laurent, 1989 Diagnosis. Rostrum significantly depressed below level of carapace, little longer than eyestalks, broadly based triangle, barely denticulate laterally. Carapace median carina with spines or teeth; submedian gastric carina smooth; lateral gastric carina unarmed. Scaphocerite short, comma-shaped in lateral view. Major cheliped propodus cylindrical, swollen, upper margins smooth; fingers shorter than palm; fixed finger blade with tooth near midpoint. Minor cheliped propodus narrower than major, compact. Maximum cl. 21 mm. Intertidal, subtidal (0–8 m). Temperate Australasia. 1 species (Poore and Collins 2009: rediagnosis, redescription).

Eiconaxius Bate, 1888 Diagnosis. Rostrum triangular, concave dorsally, triangular, produced acutely beyond and below dorsolateral margins, not or barely denticulate laterally. Carapace supraocular spines absent; median carina not extending posteriorly over broad triangular midgastric region; submedian gastric carina smooth, curved to meet median carina at base of rostrum; lateral gastric carina unarmed. Cornea usually unpigmented. Scaphocerite simple. Major cheliped carpus-palm swollen, as massive as carapace; fingers shorter than palm, or with blade-like tooth near base. Minor cheliped massive, with tapering fixed finger, curved dactylus longer than palm. Pereopods 3 and 4, dactyli broad, ovate, each with single row of spiniform setae along flexor margin, few supplementary lateral spiniform setae. Pereopod 5 dactylus spatulate, distal-flexor margin oblique, with row of spiniform setae, with or without facial spiniform setae. Male pleopod 1 of 1 simple article, or absent. Uropodal exopod without transverse suture. Maximum cl. 10 mm (Pl. 14g, h). Shelf–bathyal (140–2019 m; associated with demosponges and hexactinellid sponges). Tropical W Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate Australasia. 37 species (Poore 2017: synonymy; Poore 2018b: Caribbean species; Poore 2020: rediagnosis, IndoWest Pacific species; Poore and Dworschak 2018).

199

Eucalastacus Sakai, 1992 Diagnosis. Hermaphroditic. Rostrum 3 or more times as long as short eyestalks, styliform, with prominent, spaced lateral spines. Carapace median and submedian gastric carinae smooth; lateral gastric carina unarmed. Eyestalk short, flattened, about as long as wide, fused to carapace; cornea unpigmented. Scaphocerite simple. Major cheliped palm dilating; fingers twice as long as palm, gaping. Minor cheliped similar to major cheliped. Pleopod 1 with article 2 as long as article 1, triangular, with appendix interna represented by small hooks. Male pleopod 2 endopod with axial elongate-tapering appendix masculina bearing free, elongate basal appendix interna, setose distally. Uropodal endopod oval; exopod without transverse suture. Maximum cl. 11.7 mm. Shelf, slope (290–341 m). Temperate Australasia (New ­Zealand). 1 species (Sakai 1992).

Eutrichocheles Wood Mason, 1876 Diagnosis. Hermaphroditic. Rostrum significantly depressed below level of carapace, narrowly triangular, barely denticulate laterally. Carapace median carina with spines or teeth; submedian gastric carina comprising row of carinate teeth; lateral gastric carina unarmed or with few well spaced teeth (in addition to anterior defining supraocular tooth). Scaphocerite with small spine on mesial margin. Major cheliped elongate, propodus upper margin tuberculate, with distal tooth; fingers longer than palm; fixed finger blade with erect tooth near base. Minor cheliped similar to major cheliped, fixed finger with smaller tooth. Male pleopod 1 of 1 simple article. Pleopods 2–5 endopod of both sexes without appendix interna. Maximum cl. 30 mm. Subtidal, shelf (6–30 m). Western and Central Indo-Pacific. 2 species (Ngoc-Ho et al. 2005: rediagnosis). Paratrichocheles Sakai, 2011 is treated as a junior synonym.

Formosaxius Komai, Lin & Chan, 2010 Diagnosis. Rostrum little longer than eyestalks, triangular, with 4 pairs of lateral teeth. Carapace cupraocular spines indistinguishable from others on rostrum; median carina denticulate; submedian gastric carina comprising row of carinate teeth; lateral gastric carina unarmed. Scaphocerite simple. Major chela upper margins smooth; fixed finger blade with row of corneous setae. Maximum cl. 12.6 mm. Shelf, slope (249–360 m). Central Indo-Pacific (Taiwan). 1 species (Komai et al. 2010).

Guyanacaris Sakai, 2011 Diagnosis. Rostrum about as long as eyestalks, styliform, barely denticulate laterally. Carapace supraocular spines indistinguishable from others on rostrum; median carina denticulate; submedian gastric carina comprising row of carinate teeth; lateral gastric carina with several small teeth; postcervical carina present over whole postcervical length. Scaphocerite straight, with basal mesial spine. Major cheliped elongate, carpus-propodus or carpus-dactylus upper margin with row of teeth, palm densely setose;

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fingers as long or longer than palm. Minor cheliped similar to but slightly narrower than major cheliped. Male pleopod 1 of 1 simple article. Maximum cl. 30 mm. Shelf, slope (41–260 m). Temperate Northern Pacific, IndoWest Pacific, Tropical Eastern Pacific. 5 species (Boesch and Smalley 1972: species description; Dworschak and Poore 2018; Sakai 2011: diagnosis; Squires 1979: species description). Bruceaxius Sakai, 2011 is treated as a junior synonym – its diagnosis is indistinguishable.

Leonardsaxius Sakai, 2011 Diagnosis. Rostrum triangular, with prominent, spaced lateral spines. Carapace supraocular spines indistinguishable from others on rostrum; median carina with 2 teeth; submedian gastric carina smooth; lateral gastric carina unarmed. Scaphocerite simple. Major cheliped cylindrical, merus, propodus upper margins with subdistal tooth; fingers shorter than palm; fixed finger blade with tooth near midpoint. Minor cheliped half width of major, fingers little longer than palm. Male pleopod 1 article 2, triangular, with minute appendix interna. Maximum cl. 30 mm. Subtidal, shelf (2–31 m). Temperate Northern Pacific, Temperate SE Australia. 3 species (Marin 2015: redescription; Poore and Griffin 1979; Sakai 2011: diagnosis).

Levantocaris Galil & Clark, 1993 Diagnosis. Hermaphroditic. Rostrum narrowly triangular, with 1–4 pairs of lateral teeth. Carapace supraocular spines indistinguishable from others on rostrum; median carina smooth; submedian gastric carina absent; lateral gastric carina unarmed. Cornea weakly pigmented. Scaphocerite simple. Major cheliped cylindrical, upper margins smooth; fingers as long as palm. Minor cheliped similar to major cheliped. Male pleopod 1 article 2, triangular, with minute appendix interna. Maximum cl. 11 mm. Slope (1400 m). Temperate Northern Atlantic (E Mediterranean). 1 species (Galil and Clark 1993).

Carapace covered with short, stiff, yellowish, forwardly-directed setae; median carina smooth; submedian gastric carina with 4 teeth; lateral gastric carina with 1 prominent tooth; postcervical carina present over whole postcervical length. Pleonites 2–5 with median carina becoming obsolete posteriorly. Eyestalk short, with obtuse apex; cornea unpigmented. Scaphocerite simple, much shorter than article 4. Major cheliped palm dilating, upper margins denticulate; fingers longer than palm, gaping; fixed finger blade with tooth near middle. Minor cheliped similar to but slightly narrower than major cheliped. Uropodal endopod oval. Maximum cl. 8 mm. Slope (1733 m). Western Indo-Pacific. 1 species (Sakai 2011).

Lophaxius Kensley, 1989 Diagnosis. Hermaphroditic. Rostrum 3 or more times as long as short eyestalks, triangular, with prominent, spaced lateral spines. Carapace covered with granules; median carina with 1 tooth; submedian gastric carina absent; lateral gastric carina with 1 prominent tooth; postcervical carina present over whole postcervical length. Eyestalk short, with obtuse apex; cornea unpigmented. Scaphocerite simple, much shorter than article 4. Major cheliped palm dilating, upper margins denticulate; fingers longer than palm, gaping. Minor cheliped similar to but slightly narrower than major cheliped. Pleopod 1 with article 2 shorter than article 1, wedge-shaped, distally lobate, with small appendix interna lobe. Male pleopod 2 endopod with axial elongate-tapering appendix masculina bearing free, elongate basal appendix interna, setose distally. Maximum cl. 19.5 mm. Shelf, slope (250–1733 m). Temperate Northern Pacific, Western Indo-Pacific. 3 species (Kensley 1989). Calocarisopsis Sakai, 2011 was said to differ from Lophaxius on the supposed absence of an appendix masculina, an observation contradicted by illustrations. We treat the genus as a junior synonym.

Manaxius Kensley, 2003

Diagnosis. Rostrum about as long as eyestalks, triangular, with 1 pair of lateral teeth. Carapace median carina denticulate; submedian gastric carina with ~5 tubercles; lateral gastric carina with 1 or few well spaced teeth (in addition to anterior defining supraocular tooth). Scaphocerite short, comma-shaped in lateral view. Major cheliped cylindrical, upper margins smooth, palm with clusters of plumose setae; fingers shorter than palm. Minor cheliped one-third width of major, fingers as long as palm. Male pleopod 1 article 2, triangular, with minute appendix interna. Maximum cl. 15.7 mm. Intertidal, subtidal (0–10 m). Temperate Northern W Pacific. 1 species (Komai and Tachikawa 2007).

Diagnosis. Rostrum 1–2 times as long as eyestalks, triangular, with 1 or more pairs of lateral teeth. Carapace median carina denticulate; submedian gastric carina comprising 2 or row of carinate teeth; lateral gastric carina with first tooth set about one-third to half way along gastric region from supraocular tooth. Pleonites 2–5 with longitudinal groove above pleura. Scaphocerite simple. Major cheliped elongate, carpus-propodus upper margin with row of denticles, teeth or spines, palm covered with numerous tufts of long setae. Minor cheliped similar to but slightly narrower than major cheliped. Telson anterolateral lobe with posteriorly directed tooth. Maximum cl. 10.4 mm (Pl. 15a). Shelf, slope (49–638 m). Tropical Atlantic, Temperate Northern W Pacific, Indo-West Pacific, Tropical Eastern Pacific. 18 species. The taxonomic history of Manaxius is especially confusing but has recently been resolved (Komai 2020; Komai and Anker 2020; Poore 2020).

Lophaxiopsis Sakai, 2011

Marianaxius Kensley, 1996

Diagnosis. Hermaphroditic. Rostrum 3 or more times as long as short eyestalks, narrowly triangular, with 2 or 3 pairs of lateral teeth.

Diagnosis. Hermaphroditic. Rostrum significantly depressed below level of carapace, broadly triangular, not dentate laterally.

Litoraxius Komai & Tachikawa, 2007

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

Carapace supraocular spines indistinguishable from others on rostrum; median carina indistinct; submedian gastric carina absent; lateral gastric carina unarmed. Scaphocerite simple. Major cheliped propodus cylindrical, swollen, upper margins smooth; fingers shorter than palm; fixed finger blade with blade-like tooth near base. Minor cheliped similar to major cheliped. Male pleopod 1 of 1 simple article. Pleopods 2–5 endopod of both sexes without appendix interna. Maximum cl. 6 mm. Subtidal (1–15 m). Central Indo-Pacific (Guam). 1 species (Kensley 1996c).

Michelaxiopsis Poore & Collins, 2009 Diagnosis. Rostrum triangular, with prominent, spaced lateral spines. Carapace supraocular spines indistinguishable from others on rostrum; median carina denticulate; submedian gastric carina a hair-pin shaped double row of tubercles; lateral gastric carina unarmed. Scaphocerite straight, with basal mesial spine. Major cheliped cylindrical, upper margins smooth; fingers shorter than palm; fixed finger blade with blade-like tooth near base. Minor cheliped half width of major. Male pleopod 2 endopod with appendix interna, without appendix masculina. Maximum cl. 25 mm. Intertidal, subtidal (0–6 m). Temperate S Australia. 2 species (Poore and Collins 2009).

Montanaxius Dworschak, 2016 Diagnosis. Rostrum significantly depressed below level of carapace, broadly triangular (as long as wide at base), with 4 pairs of lateral teeth. Carapace supraocular spines indistinguishable from others on rostrum; median carina with 2 teeth; submedian gastric carina absent; lateral gastric carina unarmed. Scaphocerite straight, with basal mesial spine. Major cheliped palm ovoid, propodus upper margin carinate; fingers shorter than palm; fixed finger blade with tooth near middle. Minor cheliped similar to major cheliped, fingers as long as palm. Male pleopod 1 article 2 half as long as article 1, triangular, with distomesial field of hooks and distolateral spine. Maximum cl. 7.6 mm (Pl. 15b). Slope (299–1135 m; associated with hexactinellid sponges). Western Indo-Pacific. 2 species (Dworschak 2016; Poore 2020).

Oxyrhynchaxius Parisi, 1917 Diagnosis. Rostrum one-third carapace length, styliform, barely denticulate laterally. Carapace covered with scales or spines; supraocular spines minute; median carina smooth; submedian gastric carina comprising row of carinate teeth; lateral gastric carina with 1 prominent tooth. Eyestalk cylindrical, elongate, as long as rostrum. Scaphocerite simple. Major cheliped elongate, carpus– dactylus upper margins with short teeth or oblique spines, palm covered with numerous tufts of long setae; fingers shorter than palm. Minor cheliped similar to but slightly narrower than major cheliped. Male pleopod 1 of 1 simple article. Maximum cl. 25.5 mm. Shelf, slope (40–600 m). Temperate Northern W Pacific, Central Indo-Pacific. 3 species (Lin 2006; Lin et al. 2000: rediagnosis, second species).

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Parascytoleptus Sakai & de Saint Laurent, 1989 Diagnosis. Rostrum significantly depressed below level of carapace, about as long as eyestalks, broadly triangular (as long as wide at base), with 1 or more pairs of lateral teeth. Carapace supraocular spines indistinguishable from others on rostrum; median carina denticulate; submedian gastric carina absent; lateral gastric carina unarmed. Scaphocerite simple. Major cheliped cylindrical, upper margins smooth; fingers shorter than palm; fixed finger blade with blade-like tooth near base. Minor cheliped elongate, half width of major. Male pleopod 1 of 1 simple article, or absent. Male pleopod 2 endopod with appendix interna, without appendix masculina. Uropodal exopod without transverse suture. Maximum cl. 7 mm (Pl. 15c). Subtidal, shelf (0.5–59 m). Indo-West Pacific. 2 species (Poore and Collins 2010: rediagnosis, second species).

Paraxiopsis De Man, 1905 Diagnosis. Gonochoristic or hermaphroditic. Rostrum significantly depressed below level of carapace, narrowly triangular, with 1 or more pairs of lateral teeth or barely denticulate. Carapace median carina with spines or teeth; submedian gastric carina smooth or comprising row of carinate teeth; lateral gastric carina unarmed, or with 1 or few well spaced teeth (in addition to anterior defining supraocular tooth). Scaphocerite with small spine on mesial margin. Major cheliped cylindrical, propodus upper margin tuberculate; fingers shorter than palm; fixed finger blade with blade-like tooth near base. Minor cheliped similar to major cheliped, fixed finger with smaller tooth. Male pleopod 1 of 1 simple article. Pleopods 2–5 endopod of both sexes without appendix interna. Maximum cl. 12 mm (Pl. 15d). Intertidal–shelf (0–140 m). Tropical W Atlantic, Indo-West Pacific. 18 species (Dworschak 2020: burrows; Felder 2021: key to 9 W Atlantic species; Kensley 1996a: rediagnosis, key to 9 species; Ngoc-Ho et al. 2005: rediagnosis).

Paraxius Bate, 1888 Diagnosis. Rostrum significantly depressed below level of carapace, broadly triangular, with lateral rows of contiguous erect teeth (usually 4 or more pairs). Carapace supraocular spines indistinguishable from others on rostrum; median carina smooth; submedian gastric carina and lateral gastric carina absent. Eyestalk flattened distally; cornea unpigmented. Scaphocerite short, comma-shaped in lateral view. Major cheliped palm dilating, upper margins smooth; fingers as long as palm; fixed finger blade denticulate. Minor cheliped similar to major cheliped. Male pleopod 1 of 1 simple article. Male pleopod 2 endopod with appendix interna, without appendix masculina. Maximum cl. 12.2 mm (Pl. 15e). Slope (300–1957 m). Central Indo-Pacific. 1 species (Poore 2020: figures; Sakai and de Saint Laurent 1989: rediagnosis).

Pilbaraxius Poore & Collins, 2009 Diagnosis. Rostrum narrowly triangular, with 1 pair of lateral teeth. Carapace tuberculate; median carina smooth (pore

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tubercle only); submedian gastric carina with 1 tooth; lateral gastric carina with single prominent tooth set well back (about third way along gastric region) from supraocular tooth. Pleura 2–5 attenuated to point. Scaphocerite simple. Major cheliped palm dilating, propodus upper margin tuberculate, palm covered with numerous tufts of long setae; fingers longer than palm; fixed finger blade with erect tooth near base. Minor cheliped similar to but slightly narrower than major cheliped. Male pleopod 2 endopod with appendix interna, without appendix masculina. Maximum cl. 7 mm. Slope (400 m). Central Indo-Pacific (NW Australia). 1 species (Poore and Collins 2009).

Pillsburyaxius Sakai, 2011 Diagnosis. Rostrum narrowly triangular, with 1 or more pairs of lateral teeth. Carapace median carina denticulate; submedian gastric carina converging posteriorly, with 4 or 5 teeth; lateral gastric carina with single prominent tooth set well back (about third way along gastric region) from supraocular tooth. Scaphocerite simple. Major cheliped elongate, carpus–dactylus upper margins with short teeth or oblique spines, palm covered with numerous tufts of long setae; fingers longer than palm. Minor cheliped similar to but half as wide as major cheliped. Male pleopod 1 of 1 simple article. Maximum cl. 28 mm. Shelf, slope (135–438 m). Central Indo-Pacific. 5 species (Poore 2020: rediagnosis).

Planaxius Komai & Tachikawa, 2008 Diagnosis. Rostrum significantly depressed below level of carapace, narrowly triangular, barely denticulate laterally. Carapace median carina smooth; submedian gastric carina absent; lateral gastric carina unarmed. Scaphocerite short, comma-shaped in lateral view. Major cheliped propodus cylindrical, swollen, upper margins smooth; fingers shorter than palm; fixed finger blade smooth. Minor cheliped half width of major, fingers little longer than palm. Male pleopod 1 articles 1 and 2 fused, distal part expanded, bifid apex, mesial margin with row of hooks forming appendix interna. Maximum cl. 4.4 mm. Shelf (47–114 m). Temperate Northern W Pacific, Central Indo-Pacific. 1 species (Komai and Tachikawa 2008).

Pseudoaxiopsis Sakai, 2011 Diagnosis. Rostrum triangular, with prominent, spaced lateral spines. Carapace covered with scales or spines; supraocular spines indistinguishable from others on rostrum; median carina with spines or teeth; submedian gastric carina comprising row of carinate teeth; lateral gastric carina with 1 or few well spaced teeth (in addition to anterior defining supraocular tooth). Pleonites 2–5 with 5 longitudinal ridges and lateral carina above pleura. Eyestalk cylindrical, elongate, ~4 times as long as cornea diameter. Scaphocerite straight, with basal mesial spine. Major cheliped cylindrical, upper margins smooth; fingers longer than palm; fixed finger blade denticulate. Minor cheliped similar to major

cheliped. Male pleopod 1 an obsolete button. Maximum cl. 18 mm. Shelf (25 m). Central Indo-Pacific (NE Australia). 1 species (Sakai 2011: diagnosis, redescription).

Ralumcaris Sakai, 2011 Diagnosis. Rostrum significantly depressed below level of carapace, broadly triangular (as long as wide at base), with or without pair of lateral teeth. Carapace median carina smooth; submedian gastric carina absent; lateral gastric carina unarmed. Scaphocerite with small spine on mesial margin. Major cheliped propodus cylindrical, swollen, upper margins smooth; fingers shorter than palm; fixed finger blade smooth. Minor cheliped similar to major cheliped. Pleopods 2–5 endopod of both sexes without appendix interna. Maximum cl. 6 mm (Pl. 15f). Intertidal–shelf (1–36 m). Central Indo-Pacific. 1 species (Poore 2018a: illustration; Sakai 2011).

Scytoleptus Gerstaecker, 1856 Diagnosis. Rostrum significantly depressed below near vertical anteromedial margin, broadly triangular (as long as wide at base), not dentate laterally. Carapace supraocular spines indistinguishable from others on rostrum; median carina with spines or teeth; submedian gastric carina smooth; lateral gastric carina unarmed. Pleura 2–5 truncate, with posteroventral angle. Scaphocerite simple. Major cheliped palm swollen, upper margins smooth; fingers shorter than palm; fixed finger blade with blade-like tooth near base. Minor cheliped elongate, half width of major, fingers half length of palm. Male pleopod 1 of 1 simple article. Uropodal exopod without transverse suture. Maximum cl. 26 mm. Intertidal–shelf (0–36 m). Western and Central Indo-Pacific. 2 species (Sakai 2011: rediagnosis, second species).

Spongiaxius Sakai & de Saint Laurent, 1989 Diagnosis. Rostrum narrowly triangular or linguiform, barely denticulate laterally or with erect marginal teeth. Carapace supraocular spines indistinguishable from others on rostrum; median carina with spines or teeth; submedian gastric carina smooth, or comprising row of carinate teeth; lateral gastric carina unarmed, or denticulate or with 1 or few well spaced teeth (in addition to anterior defining supraocular tooth. Cornea fully or weakly pigmented. Scaphocerite simple, curved down. Major cheliped carpus-palm swollen, as massive as carapace; fingers sometimes massive. Minor cheliped massive, with tapering fixed finger, deep dactylus longer than palm. Pereopod 2 dactylus upper margin with dense field of fine setae (S. mananalem excepted). Male pleopod 1 of 1 simple article, or article 2, triangular, with minute appendix interna, or absent, or an obsolete button. Uropodal endopod with distal angles, distal margin truncate or oblique, with posterodistal angle often defined by spine, exceeding anterodistal angle. Maximum cl. 24  mm (Pl. 15g).

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

Shelf, slope (60–690 m; associated with demosponges and hexactinellid sponges). Central Indo-Pacific, Temperate Australasia. 15 species (Poore 2020: rediagnosis, Indo-West Pacific species; Poore and Collins 2010: rediagnosis). References Boesch DF, Smalley AE (1972) A new axiid (Decapoda, Thalassinidea) from the Northern Gulf of Mexico and tropical Atlantic. Bulletin of Marine Science 22, 45–52. de Matos-Pita SS, Ramil F (2015) Additions to thalassinidean fauna (Crustacea: Decapoda) off Mauritania (NW Africa) with the description of a new genus and a new species. Zootaxa 4020, 571–587. doi:10.11646/zootaxa.4020.3.9 Dworschak PC (2016) A new genus and species of axiid shrimp (Crustacea, Decapoda) from a southwestern Indian Ocean seamount. European Journal of Taxonomy 229, 1–11. doi:10.5852/ejt.2016.229 Dworschak PC (2020) A new species of Paraxiopsis de Man, 1905 (Decapoda: Axiidea: Axiidae) from Safaga (Red Sea) with notes on its burrows. Annalen des Naturhistorischen Museums in Wien, B 122, 159–167. Dworschak PC, Poore GCB (2018) More cautionary tales: family, generic and species synonymies of recently published taxa of ghost and mud shrimps (Decapoda, Axiidea and Gebiidea). Zootaxa 4394, 61–76. doi:10.11646/zootaxa.4394.1.3 Felder DL (2021) Description of Paraxiopsis kensleyi n. sp., a new axiid lobster from the Gulf of Mexico (Crustacea: Decapoda: Axiidea). Zootaxa 4965, 129–141. doi:10.11646/zootaxa.4965.1.6 Galil BS, Clark PF (1993) A new genus and species of axiid (Decapoda, Thalassinidea) from the Levantine Basin of the Mediterranean. Crustaceana 64, 48–55. doi:10.1163/156854093X00054 Kensley B (1989) New genera in the thalassinidean families Calocarididae and Axiidae (Crustacea: Decapoda). Proceedings of the Biological Society of Washington 102, 960–967. Kensley B (1994) The genus Coralaxius redefined, with descriptions of two new species (Crustacea: Decapoda: Axiidae). Journal of Natural History 28, 813–828. doi:10.1080/00222939400770421 Kensley B (1996a) The genus Paraxiopsis De Man, with descriptions of new species from the Western Atlantic (Crustacea: Decapoda: Axiidae). Bulletin of Marine Science 58, 709–729. Kensley B (1996b) New species of Calocarididae from the Caribbean Sea and Gulf of Mexico (Crustacea: Decapoda: Thalassinidea). Bulletin of Marine Science 59, 158–168. Kensley B (1996c) New thalassinidean shrimp from the Pacific Ocean (Crustacea: Decapoda: Axiidae and Calocarididae). Bulletin of Marine Science 59, 469–489. Kensley B (1996d) Systematics and distribution of the genus Calocarides (Crustacea: Decapoda: Axiidae). Proceedings of the Biological Society of Washington 109, 53–69. Kensley B (2001) Two sympatric species of Axius from the north-west Atlantic (Decapoda, Thalassinidea, Axiidae). Crustaceana 74, 951–962. doi:10.1163/15685400152682692 Kensley B (2003) Axioid shrimps from Guam (Crustacea, Decapoda, Thalassinidea). Micronesica 35–36, 359–384. Kensley B, Gore RH (1981) Coralaxius abelei, new genus and new species (Crustacea: Decapoda: Thalassinidea: Axiidae): a coral-inhabiting shrimp from the Florida Keys and the western Caribbean Sea. Proceedings of the Biological Society of Washington 93, 1277–1294. Kensley B, Simmons GM (1988) Axiorygma nethertoni, a new genus and species of thalassinidean shrimp from Florida (Decapoda: Axiidae). Journal of Crustacean Biology 8, 657–667. doi:10.2307/1548701

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Pemberton SG, Risk MJ, Buckley DE (1976) Supershrimp: deep bioturbation in the Strait of Canso, Nova Scotia. Science 192, 790–791. doi:10.1126/science.192.4241.790 Poore GCB (1994) A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys to the families and genera. Memoirs of the Museum of Victoria 54, 79–120. doi:10.24199/j.mmv.1994.54.03 Poore GCB (2017) Synonymy and problematic species of Eiconaxius Spence Bate, 1888, with descriptions of new species (Crustacea: Decapoda: Axiidea: Axiidae). Zootaxa 4231, 364–376. doi:10.11646/ zootaxa.4231.3.4 Poore GCB (2018a) Burrowing lobsters mostly from shallow coastal environments in Papua New Guinea (Crustacea: Axiidea: Axiidae, Micheleidae). Memoirs of Museum Victoria 77, 1–14. doi:10.24199/j. mmv.2018.77.01 Poore GCB (2018b) Caribbean species of Eiconaxius (Decapoda: Axiidea: Axiidae). Zootaxa 4524, 139–146. doi:10.11646/zootaxa.4524.1.11 Poore GCB (2020) Axiid and micheleid lobsters from Indo-West Pacific deep-sea environments (Crustacea: Decapoda: Axiidea: Axiidae, Micheleidae). In: Corbari L, Chan T-Y, Ahyong ST (Eds) Deep-sea crustaceans from Papua New Guinea. Tropical Deep-Sea Benthos vol. 31. Mémoires du Muséum National d’Histoire Naturelle, Paris 213, 259–367. Poore GCB, Collins DJ (2009) Australian Axiidae (Crustacea: Decapoda: Axiidea). Memoirs of Museum Victoria 66, 221–287. doi:10.24199/j. mmv.2009.66.20 Poore GCB, Collins DJ (2010) Parascytoleptus and Spongiaxius redefined with two new species from the southwestern Pacific (Decapoda: Axiidea: Axiidae). In Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume (Eds Fransen, C, De Grave S, Ng P). Crustaceana Monographs 14, 611–625. Poore GCB, Dworschak PC (2018) The Eiconaxius cristagalli species complex (Decapoda, Axiidea, Axiidae). Memoirs of Museum Victoria 77, 105–120. doi:10.24199/j.mmv.2018.77.06 Poore GCB, Griffin DJG (1979) The Thalassinidea (Crustacea: Decapoda) of Australia. Records of the Australian Museum 32, 217–321. doi:10.38 53/j.0067-1975.32.1979.457 Richer de Forges B, Tan SH, Bouchet P, Ng PKL, Chan T-Y, et  al. (2009) Panglao 2005 – Survey of the deep-water benthic fauna of the Bohol Sea and adjacent waters. Raffles Bulletin of Zoology Supplement 20, 21–38. Robles R, Tudge CC, Dworschak PD, Poore GCB, Felder DL (2009) Molecular phylogeny of the Thalassinidea based on nuclear and mitochondrial genes. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 309–326. CRC Press, Boca Raton. Sakai K (1986) Axiopsis brucei sp. nov., a new sponge-inhabiting axiid (Crustacea: Decapoda: Thalassinidea), from north-west Australia. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 3, 11–20. Sakai K (1987) Two new Thalassinidea (Crustacea: Decapoda) from Japan, with the biogeographical distribution of the Japanese Thalassinidea. Bulletin of Marine Science 41, 296–308. Sakai K (1992) Axiid collections of the Zoological Museum, Copenhagen, with the description of one new genus and six new species (Axiidae, Thalassinidea, Crustacea). Zoologica Scripta 21, 157–180. doi:10.1111/j.​ 1463-6409.​1992.tb00318.x Sakai K (2011) Axioidea of the world and a reconsideration of the Callianassoidea (Decapoda, Thalassinidea, Callianassida). Crustaceana Monographs 13, 1–616. doi:10.1163/9789047424185 Sakai K, Apel M (2002) Thalassinidea (Crustacea: Decapoda) from Socotra Archipelago, Yemen, with a new species of Lepidophthalmus. Fauna of Arabia 19, 273–288. Sakai K, de Saint Laurent M (1989) A check list of Axiidae (Decapoda, Crustacea, Thalassinidea, Anomula), with remarks and in addition

descriptions of one new subfamily, eleven new genera and two new species. Naturalists, Publications of Tokushima Biological Laboratory, Shikoku University 3, 1–104. Squires JH (1979) Axiopsis caespitosa (Thalassinidea, Axiidae), a new species from the Pacific coast of Colombia. Canadian Journal of Zoology 57, 1584–1591. doi:10.1139/z79-207 Tsang LM, Lin F-J, Chu KH, Chan T-Y (2008a) Phylogeny of Thalassinidea (Crustacea, Decapoda) inferred from three rDNA sequences: implications for morphological evolution and superfamily classification. Journal of Zoological Systematics and Evolutionary Research 46, 216–223. doi:10.1111/j.1439-0469.2008.00459.x Tsang LM, Ma KY, Ahyong ST, Chan T-Y, Chu KH (2008b) Phylogeny of Decapoda using two nuclear protein-coding genes: Origin and evolution of the Repantia. Molecular Phylogenetics and Evolution 48, 359–368. doi:10.1016/j.ympev.2008.04.009

Callianassidae Dana, 1852 ghost shrimps Figures 9.8–9.13, Plate 16a–g Callianassids are elongate, very soft-bodied shrimps, white or pale pink, with a pleon longer than the carapace. They are common, mostly in the tropics where they build permanent burrows in soft sediments of the shelf or reef lagoons. Burrow structure and feeding behaviour are well studied (Berkenbusch and Rowden 1998; Bird et al. 1999; Bird and Poore 1999; Dworschak 1998; Nickell and Atkinson 1995; Stapleton et  al. 2001; Tamaki et  al. 2018; Tamaki and Ueno 1998; de Vaugelas and de Saint Laurent 1984; Witbaard and Duineveld 1989). Some are filter feeders but others pull seagrass down into the burrows to harvest as food (Stamhuis et  al. 1998). There have been several studies of their larval development (e.g. Nates et al. 1997). The largest callianassids are up to 15 cm long and in south-east Asia are sold as food. In Australia, ghost shrimps are captured for fish bait using a ‘yabby pump’. The taxonomy of Callianassidae sensu lato has been fluid over the last several decades. Several family and generic names have been introduced, notably summarised by Sakai (2005, 2011). A more recent classification based on molecular and morphological phylograms (Poore et  al. 2019; Robles et  al. 2020) differs considerably from Sakai’s arrangements and is followed here. Poore et al. (2019) provided more extensive diagnoses, keys, references and lists of species by genus. Diagnosis. Rostrum flat, triangular, shorter than eyestalks (Fig. 9.8b, f), or spike-like (Fig. 9.8c, e, h, j); anterior branchiostegal lobe sclerotised, merging smoothly with anterodorsal branchiostegal angle (Fig.  9.8e, g, i, l). Scaphocerite rudimentary (Fig. 9.8b–m). Maxilliped 1 epipod truncate, without anterior lobe (Fig.  9.8r). Maxilliped 3 propodus longer than wide, not prominently lobed on lower margin, or about as wide as long (Fig. 9.9). Female pleopod 2 rami styliform, endopod much longer than broad, or absent (Fig. 9.12f–h). Uropodal exopod with elevated dorsal plate (Fig. 9.13).

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

Implicit generic attributes. Gonochoristic. Rostrum triangular, not reaching cornea. Antennular peduncle about as long or little longer than antennal peduncle. Maxilliped 3 merus mesiodistally convex, extending only slightly beyond articulation of carpus, without distal spine on mesial margin; dactylus tapering, with scattered setae over upper margin, dense brush of short setae distally on lower margin.

205

Major cheliped carpus proximal and lower margins evenly convex. Minor cheliped slender, considerably narrower than major cheliped; carpus upper margin longer than propodus. Uropodal endopod ovoid; exopod distal margin poorly differentiated from anterior margin, about as long as wide. Telson longer than wide, tapering evenly from near base.

Key to genera of Callianassidae 1. – 2. – 3. –

Antennular peduncle length ~4 times width of both eyestalks, twice as long as antennal peduncle, with 2 dense rows of adjacent long setae on lower surface (Fig. 9.8o). Maxilliped 3 merus produced as massive triangular lobe alongside palp (Fig. 9.9k)������������������������������������������������������������������������������������������������������������������� Trypaea Antennular peduncle length at most 3 times width of both eyestalks, with at most bands of scattered long setae on lower surface (Fig. 9.8a). Maxilliped 3 merus distally convex at most����������������������������������������������������������������2 Small triangular sclerite present on branchiostegite at anterior end of linea thalassinica (Fig. 9.8a)��������Aqaballianassa Without triangular sclerite on branchiostegite at anterior end of linea thalassinica (Fig. 9.8g, i, l)�����������������������������������3 Telson tapering, posterior margin with medial spine in deep notch (Fig. 9.13g, o)���������������������������������������������������������������4 Telson posterior margin convex, truncate or at most with shallow medial concavity, sometimes with small medial spine���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������5

Fig. 9.8.  Callianassidae. Anterior carapace, eyestalks, antennules, antennae: a, Aqaballianassa; b, Arenallianassa; c, Caviallianassa; d, e, Coriollianassa; f, g, Filhollianassa; h, Jocullianassa; i, Notiax j, Praedatrypaea; k, Pugnatrypaea; l, m, Rayllianassa; n, Rudisullianassa; o, Trypaea. Carapace, pleonites 1, 2: p, Arenallianassa. Maxilla: q, Arenallianassa. Maxilliped 1: r, Arenallianassa. Mandible, mesial and lower views: s, Callianassa; t, Rayllianassa.

206

4.



5. – 6. – 7. – 8. – 9.

– 10.

Marine Decapod Crustacea

Maxilliped 3 merus almost rectangular, distally truncate with squarish angle between distal and lower margins (Fig. 9.9d). Pereopod 3 propodus linear, without lobe on lower margin (Fig. 9.12a). Uropodal exopod distal margin with anterodistal angle obtuse. Telson with pair of broad posterior lobes, widely excavate at midpoint (Fig. 9.13g)�������������������������������������������������������������������������������������������������������������������Lipkecallianassa Maxilliped 3 merus distally oblique with obtuse angle between distal and lower margins (Fig. 9.9h). Pereopod 3 propodus oval, lower margin slightly convex, leading to narrow, distinctly rounded proximal lobe. Uropodal exopod distal margin clearly differentiated from anterior margin, anterodistal corner right-angled. Telson tapering over distal third to pair of posterior lobes separated by deep notch (Fig. 9.13o)�������������������������������Pugnatrypaea Antennular peduncle exceeded by antennal peduncular article 5 (Fig. 9.8d, j)���������������������������������������������������������������������6 Antennular peduncle as long or longer than antennal peduncle (Fig. 9.8b, c, h, m)�����������������������������������������������������������10 Maxilliped 3 merus with spine on distal free margin (Fig. 9.9g). Proximolateral lobe of telson prominent, defined posteriorly by clear unchitinised region (Fig. 9.13n)�����������������������������������������������������������������������Praedatrypaea Maxilliped 3 merus without spine on distal free margin. Proximolateral lobe of telson prominent or not, indefinitely defined posteriorly������������������������������������������������������������������������������������������������������������������������������������������������7 Major cheliped carpus articulating by means of a short neck with merus, merus with bifid proximal spine (Fig. 9.10c). Scaphocerite bifid (Fig. 9.8d)��������������������������������������������������������������������������������������������� Coriollianassa Major cheliped carpus without neck, lower margin evenly curved, merus lower margin simple, with simple spine or blade. Scaphocerite simple������������������������������������������������������������������������������������������������������������������������������������������8 Major cheliped upper margins of ischium and merus, lower margins of merus and carpus beaded, with dense row of fine setae, propodus with deep notch at base of fingers (Fig. 9.10d, e)������������������������������������������ Biffarius Major cheliped upper margins of ischium and merus, lower margins of merus and carpus smooth, with few well spaced setae, propodus without deep notch at base of fingers�������������������������������������������������������������������������������������9 Rostrum acute, as long as eyestalk (Fig. 9.8j). Cornea diffusely defined. Maxilliped 3 longer than wide at ischium-merus suture (Fig. 9.9c). Major and minor chelipeds similar, with saw-tooth blade on lower margin of merus, upper margin concave (Fig. 9.10a). Pleonite with sublateral ventral sharp ridge, flared posteriorly (Fig. 9.13d, e)��������������������������������������������������������������������������������������������������������������������������������������������Cheramus Rostrum obtusely triangular, not reaching cornea (Fig. 9.8n). Cornea well defined. Maxilliped 3 wider at ischium-merus suture than long (Fig. 9.9m). Major and minor chelipeds dissimilar, merus unarmed or with small teeth (Fig. 9.11h, m). Pleonite without sublateral ventral sharp ridge�����������������������������������Rudisullianassa Rostrum acute, reaching beyond cornea (Fig. 9.8h, j)��������������������������������������������������������������������������������������������������������������11

Fig. 9.9.  Callianassidae. Maxilliped 3: a, Callianassa; b, Caviallianassa; c, Cheramoides; d, Lipkecallianassa; e, Necallianassa; f, Neotrypaea; g, Praedatrypaea; h, Pugnatrypaea; i, Scallasis; j, Spinicallianassa; k, Trypaea; l, Arenallianassa; m, Biffarius.

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207

Fig. 9.10.  Callianassidae. Male major cheliped: a, Cheramus; b, Fragillianassa; c, Coriollianassa; d, e, Biffarius; f, Arenallianassa; g, Caviallianassa; h, Callianassa; i, Cheramoides; j, Filhollianassa; k, Gilvossius. Minor cheliped: l, Cheramoides; m, Neotrypaea.

– 11.

Rostrum obsolete or triangular, not reaching cornea (Fig. 9.8b–g, i, m, n)��������������������������������������������������������������������������15 Pereopod 3 propodus bean-shaped, lower margin concave, with broadly rounded proximal lobe (Fig. 9.12d). Major cheliped merus with simple perpendicular proximal spine (Fig. 9.11a)�����������������������Jocullianassa – Pereopod 3 propodus oval or with straight lower margin (Fig. 9.12b, c). Major cheliped merus smooth (Fig. 9.10i), or with blade (Fig. 9.10j), or with spine (Fig. 9.11i, j)�������������������������������������������������������������������������������������� 12 12. Maxilliped 3 almost rectangular, distally truncate with squarish angle between distal and lower margins (Fig. 9.9c). Major cheliped merus without prominent hooked blade (unknown in Poti) (Fig. 9.10i)���������������������������13 – Maxilliped 3 distally convex, extending beyond articulation of carpus (Fig. 9.9j), or distally oblique with obtuse angle between distal and lower margins (Fig. 9.9i). Major cheliped merus with 1 or more teeth or spines�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������14 13. Uropodal exopod at least twice as long as wide (Fig. 9.13d). Pleopod 2 absent in male�����������������������������������Cheramoides – Uropodal exopod about as long as wide (Fig. 9.13l). Pleopod 2 present in male (Fig. 9.12h)������������������������������������������Poti 14. Maxilliped 3 merus wider at ischium-merus suture than long, distally convex, extending only slightly beyond articulation of carpus (Fig. 9.9j). Male major cheliped merus with oblique spine about third or half-way along lower margin, propodus swollen, longer than carpus (Fig. 9.11j). Pereopod 3 propodus subrectangular (Fig. 9.12c)������������������������������������������������������������������������������������������������������������������������������ Spinicallianassa

208

Marine Decapod Crustacea

Fig. 9.11.  Callianassidae. Male major cheliped: a, Jocullianassa; b, Notiax; c, Necallianassa; d, Neotrypaea; e, f, Paratrypaea; g, Rayllianassa; h, Rudisullianassa; i, Scallasis; j, Spinicallianassa; k, Tastrypaea. Minor cheliped: l, Jocullianassa; m, Rudisullianassa.



Maxilliped 3 merus longer than wide at ischium-merus suture, distally oblique with obtuse angle between distal and lower margins (Fig. 9.9i). Male major cheliped merus with 1 or 2 proximal similar teeth and distal denticles on lower margin, propodus narrow, shorter than carpus (Fig. 9.11i). Pereopod 3 propodus oval (Fig. 9.12b)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Scallasis 15. Major cheliped with dorsodistolateral propodus and lateral dactylus with dense brush of setae (Fig. 9.11e, f). Uropodal endopod with facial distal transverse row of short spiniform setae (Fig. 9.13m)������������������ Paratrypaea – Major cheliped with dorsodistolateral propodus and lateral dactylus sparsely setose. Uropodal endopod sometimes with facial spiniform setae but not as obvious transverse row����������������������������������������������������������������������16 16. Anterior carapace strongly curved down at ~45° in lateral view, rostrum blunt (Fig. 9.8f, g)���������������������� Filhollianassa – Anterior carapace moderately or not depressed in lateral view (Fig. 9.8i)����������������������������������������������������������������������������17 17. Antennular peduncle exceeding antennal peduncle by about half length of article 3 (Fig. 9.8m). Major and minor chelipeds similar, merus without prominent hook or spine on lower margin, carpus and propodus

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

209

ovoid in cross-section, upper and lower margins not carinate, carpus of both shorter than upper margin of propodus (Fig. 9.11g). Mandibular molar dominating, calcified, swollen projection without sharp edge, incisor without teeth (Fig. 9.8t)������������������������������������������������������������������������������������������������������������������� Rayllianassa – Antennular peduncle about as long or little longer than antennal peduncle (Fig. 9.8b, c). Chelipeds dissimilar, major cheliped merus with prominent hook or spine on lower margin, carpus and propodus flattened, upper and especially lower margins carinate, submarginal mesial face slightly concave (Fig. 9.10b, g). Minor cheliped carpus longer than upper margin of propodus (Fig. 9.10m). Mandibular molar with simple sharp margin, incisor toothed (Fig. 9.8s)������������������������������������������������������������������������������������������������������������������ 18 18. Maxilliped 3 dactylus with dense brush of long setae over most of upper-distal margin, few setae along lower margin (Fig. 9.9b)�����������������������������������������������������������������������������������������������������������������������������������������������������������19 – Maxilliped 3 dactylus with scattered setae over upper margin, dense brush of short setae distally on lower margin (Fig. 9.9a)��������������������������������������������������������������������������������������������������������������������������������������������������������������������� 20 19. Male and female major cheliped merus with prominent truncate hook, serrate along lower margin, excavate laterally at base, with deep notch at base of fingers (Fig. 9.10b). Pleopod 1 present in male, pleopod 2 absent in male. Uropodal endopod without facial spiniform setae������������������������������������������� Fragillianassa – Male major cheliped merus with 3 oblique similar short proximal spines on lower margin, beaded beyond (Fig. 9.10g), female with simple hook. Pleopods 1 and 2 present in male (Fig. 9.12e). Uropodal endopod with at least 1 facial spiniform seta (Fig. 9.13b, c)������������������������������������������������������������������������������������������Caviallianassa 20. Maxilliped 3 merus longer than width at ischium-merus suture, distally oblique with obtuse angle between distal and lower margins (see above) (Fig. 9.9a). Male major cheliped merus with simple proximal hook on lower margin (Fig. 9.10h)����������������������������������������������������������������������������������������������������������������������������������� Callianassa – Maxilliped 3 merus wider at ischium-merus suture than long, distally convex, extending beyond articulation of carpus (Fig. 9.9e, f). Male major cheliped merus with prominent complex truncate hook (Figs 9.10f, k, 9.11c)��������������������������������������������������������������������������������������������������������������������������������������������������������������������21 21. Uropodal endopod anterior margin with distal spine (Fig. 9.13h). Telson usually with 1 or 2 pairs of lateral spines���������������������������������������������������������������������������������������������������������������������������������������������������������������������� Necallianassa – Uropodal endopod anterior margin and telson lateral margin without spines (Fig. 9.13f, i, j)����������������������������������������� 22 22. Telson wider than long, posterior margin semicircular (Fig. 9.13f) or subtruncate. Uropodal endopod without facial spiniform setae����������������������������������������������������������������������������������������������������������������������������������� Gilvossius – Telson as wide as long or longer than wide, posterior margin truncate, slightly concave or slightly convex between posterolateral angles. Uropodal endopod with short distal transverse row of facial spiniform setae (Fig. 9.13i–k, p)���������������������������������������������������������������������������������������������������������������������������������������������������������������� 23

Fig. 9.12.  Callianassidae. Pereopod 3: a, Lipkecallianassa; b, Scallasis; c, Spinicallianassa; d, Jocullianassa. Male pleopod 1: e, Caviallianassa. Female pleopod 2: f, Callianassa; g, Caviallianassa; h, Poti.

210

Marine Decapod Crustacea

23. Male major cheliped merus with dentate blade at midpoint (Fig. 9.11k). Uropodal exopod posterodistal margin with row of c. 6–8 long blade-like setae proximal to long setae on distal margin (Fig. 9.13c)���������������������������������������������������������������������������������������������������������������������������������������������������� Tastrypaea – Male major cheliped merus with prominent basal truncate tooth (Figs 9.7f, 9.8b, d). Uropodal exopod distal and posterior margins densely setose, without blade-like setae��������������������������������������������������������������������������������������� 24 24. Rostrum acute, with ventral broad swelling (Fig. 9.8i). Maxilliped 3 crista dentata absent or of few proximal denticles. Male major cheliped with distal margin of propodus bearing tooth at base of dactylus (Fig. 9.11b). Uropodal endopod distal margin well differentiated from anterior margin, distal margin truncate. Telson longer than wide, tapering from base (Fig. 9.13k)��������������������������������������������������������������������������Notiax – Rostrum flat against eyestalks (Fig. 9.8b). Maxilliped 3 crista dentata consisting of a row of denticles (Fig. 9.9f). Male major cheliped with distal margin of propodus having deep notch at base of fingers

Fig. 9.13.  Callianassidae. Pleonite 6, telson, uropod: a, Arenallianassa; b, c, Caviallianassa (uropod in detail); d, e, Cheramoides (with pleonite 6 lateral); f, Gilvossius; g, Lipkecallianassa; h, Necallianassa; i, j, Neotrypaea; k, Notiax; l, Poti; m, Paratrypaea; n, Praedatrypaea; o, Pugnatrypaea; p, Tastrypaea.

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

211

(Fig. 9.11d). Uropodal exopod distal margin well or poorly differentiated from anterior margin. Telson wider than or about as wide as long�������������������������������������������������������������������������������������������������������������������������������������� 25 25. Maxilliped 3 merus distally convex, extending only slightly beyond articulation of carpus (Fig. 9.9l). Telson wider than long, posterior margin as wide as base, weakly convex between rounded posterolateral angles (Fig. 9.13a)����������������������������������������������������������������������������������������������������������������������������������������������� Arenallianassa – Maxilliped 3 merus expanded distally as rounded lobe beyond articulation with ischium (Fig. 9.9f). Telson at least as wide as long, usually wider than long, posterior margin narrower than base, often with median spine (Fig. 9.9i, j)���������������������������������������������������������������������������������������������������������������������������������������� Neotrypaea Aqaballianassa Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Hermaphrodite. Triangular sclerite separate from branchiostegite at anterior end of linea thalassinica. Pleonite 6 with sublateral ventral sharp ridge, flared posteriorly. Major cheliped merus of male with oblique spine about third or half way along lower margin; dactylus with dense setae along upper margin. Minor cheliped two-thirds width of major cheliped. Uropodal exopod distal margin clearly differentiated from anterior margin, anterodistal corner right-angled. Maximum cl. 7 mm. Subtidal, shelf (1–73 m). Western and Central Indo-Pacific. 8 species (Poore et al. 2019).

Arenallianassa Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Major cheliped merus of male with prominent truncate hook, serrate along lower margin, excavate laterally at base. Uropodal exopod distal margin clearly differentiated from anterior margin, anterodistal corner right-angled. Telson lateral margins convex. Maximum cl. 14 mm. Intertidal–subtidal (0–19 m). Temperate Australasia. 1 species (Poore et al. 2019). The type species is widespread in sandy seagrass environment in southern Australia (Bird 2000; Bird and Poore 1999; Butler and Bird 2008; Poore 1975).

Biffarius Manning & Felder, 1991 Diagnosis. Antennular peduncle exceeded by antennal peduncular article 5. Major cheliped merus of male with prominent truncate hook, serrate along lower margin, excavate laterally at base; upper margins of ischium and merus, lower margins of merus and carpus beaded, with dense row of long fine setae. Uropodal exopod distal margin clearly differentiated from anterior margin, anterodistal corner right-angled. Maximum cl. 10 mm (Pl. 16a). Subtidal. Tropical Atlantic, Temperate Australasia. 4 species (Hernáez et al. 2020: key to species; Poore et al. 2019). Two species are from the western Atlantic (Biffar 1971) and the third is extremely common in muddy sediments around south-eastern Australia (Poore 1975).

Callianassa Leach, 1814 Diagnosis. Antennular peduncle exceeded by antennal peduncular article 5. Maxilliped 3 merus distally oblique with obtuse angle between distal and mesial margins. Major cheliped merus of male

with simple proximal hook on lower margin. Uropodal endopod ovoid, usually longer than wide, anterior margin straight or slightly convex, posterodistal margin evenly convex or asymmetrical, at least as wide as long, distal margin truncate-convex, at right angles to straight anterior margin; exopod distal margin clearly differentiated from anterior margin, anterodistal corner right-angled. Maximum cl. 13 mm. Subtidal–shelf. Temperate Northern and Tropical Atlantic, Temperate Southern Africa. 6 species (Poore et al. 2019: restricted diagnosis). Callianassa has been a catch-all genus while the taxonomy of the family remained confusing but Poore et  al. (2019) have confined the genus to a few species from the Mediterranean, eastern Atlantic and southern Africa. Callianassa subterranea (Montagu, 1808) is a well studied burrower feeding on sediment (Stamhuis et al. 1997; Witbaard and Duineveld 1989).

Caviallianassa Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Antennular peduncle exceeding antennal peduncle by about half length of article 3. Maxilliped 3 dactylus ovate, with dense brush of long setae over most of upper-distal margin, few setae along lower margin. Major cheliped merus of male with 1–3 oblique similar short proximal spines on lower margin, beaded beyond. Minor cheliped two-thirds width of major cheliped. Telson about as wide as long, tapering from anterolateral lobe; proximolateral lobe undefined. Maximum cl. 4 mm. Shelf. Temperate Northern W Pacific, Central Indo-Pacific. 2 species (Komai and Fujiwara 2012: illustrations of Cheramus cavifrons; Poore et al. 2019).

Cheramoides Sakai, 2011 Diagnosis. Rostrum acute, as long as eyestalks. Pleonite 6 with sublateral ventral sharp ridge, flared posteriorly. Antennular peduncle exceeded by antennal peduncular article 5. Maxilliped 3 merus almost rectangular, distally truncate with squarish angle between distal and mesial margins. Minor cheliped slender, narrower than major cheliped, with attenuated curved dactylus, longer than palm. Uropodal exopod distal margin clearly differentiated from anterior margin, anterodistal corner right-angled, twice as long as wide. Telson posterior margin slightly concave, with medial spine. Maximum cl. 5 mm. Shelf. Tropical W Atlantic, Central Indo-Pacific. 3 species (Biffar 1971: illustrations of C. marginata (Rathbun, 1901); Poore et al. 2019).

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Marine Decapod Crustacea

Cheramus Bate, 1888 Diagnosis. Rostrum acute, as long as eyestalks. Pleonite 6 with sublateral ventral sharp ridge, flared posteriorly. Antennular peduncle exceeded by antennal peduncular article 5. Maxilliped 3 merus almost rectangular, distally truncate with squarish angle between distal and mesial margins. Minor and major chelipeds similar, both attenuated, with swollen palms. Major cheliped merus of male with serrate blade over lower margin, upper margin concave. Minor cheliped carpus upper margin not longer than propodus. Uropodal exopod distal margin clearly differentiated from anterior margin, anterodistal corner right-angled, almost as wide as long. Maximum cl. 6 mm. Slope. Tropical W Atlantic. 1 species (Biffar 1973: description of only species; Poore et al. 2019).

Coriollianassa Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Rostrum acute, as long as eyestalks, with ventral keel. Antennular peduncle exceeded by antennal peduncular article 5. Maxilliped 3 merus distally oblique with obtuse angle between distal and mesial margins. Major cheliped merus of male with proximal curved spine on lower margin; carpus articulating my means of a short neck with merus. Uropodal exopod 1.5–1.8 times as long as wide. Maximum cl. 10 mm. Shelf, slope. Western and Central Indo-Pacific. 3 species (Ngoc-Ho 2014: illustrations of C. coriolisae; Poore et al. 2019).

Filhollianassa Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Anterior carapace strongly curved down over anterior quarter. Antennular peduncle exceeded by antennal peduncular article 5. Major cheliped merus of male with prominent truncate hook, serrate along lower margin, excavate laterally at base. Uropodal endopod asymmetrical, at least as wide as long, distal margin truncate-convex, at right angles to straight anterior margin. Telson lateral margins convex. Maximum cl. 26 mm. Intertidal–subtidal. Temperate Australasia. 2 species. (­Berkenbusch and Rowden 1998, 2000; Poore 2004; Poore et al. 2019).

Fragillianassa Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Maxilliped 3 dactylus ovate, with dense brush of long setae over most of upper-distal margin, few setae along lower margin. Major cheliped merus of male with prominent truncate hook, serrate along lower margin, excavate laterally at base. Maximum cl. 6 mm (Pl. 16b). Intertidal–subtidal. Tropical W Atlantic. 3 species (Biffar 1970: illustrations as Callianassa fragilis Biffar, 1970; Pachelle and Tavares 2020; Poore et al. 2019).

Gilvossius Manning & Felder, 1992 Diagnosis. Major cheliped merus of male with prominent truncate hook, serrate along lower margin, excavate laterally at base. Telson

wider than long, subsemicircular; posterior margin rounded over posterior half or more. Maximum cl. 16 mm. Subtidal–shelf. Temperate Northern and Tropical Atlantic, Temperate Southern Africa. 9 species (Dworschak 2002; Felder and Robles 2020b; Manning and Felder 1992; Ngoc-Ho 2003; Poore et al. 2019).

Jocullianassa Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Rostrum acute, as long as eyestalks. Maxilliped 3 merus tapering, not mesially produced. Major cheliped merus of male with proximal perpendicular spine on lower margin. Pereopod 3 propodus bean-shaped, lower margin concave, leading to broadly rounded proximal lobe. Uropodal exopod distal margin clearly differentiated from anterior margin, anterodistal corner right-angled, 1.5–1.8 times as long as wide. Maximum cl. 5 mm. Subtidal. Indo-West Pacific. 5 species (Komai and Tachikawa 2008: illustrations of J. joculatrix (De Man, 1905); Poore et  al. 2019).

Lipkecallianassa Sakai, 2002 Diagnosis. Rostrum acute, as long as eyestalks. Maxilliped 3 merus almost rectangular, distally truncate with squarish angle between distal and mesial margins, with acute distomesial angle. Uropodal exopod 1.5 times as long as wide. Telson posterior margin with pair of broad posterolateral lobes, widely excavate at midpoint, with mesial spine. Maximum cl. 3 mm. Slope. Central Indo-Pacific. 1 species (Poore et al. 2019; Sakai 2002).

Necallianassa Heard & Manning, 1998 Diagnosis. Major cheliped merus of male with prominent truncate hook, serrate along lower margin, excavate laterally at base. Uropodal endopod asymmetrical, at least as wide as long, distal margin truncate-convex, at right angles to straight anterior margin, anterior margin with distal spine. Maximum cl. 12 mm. Subtidal–shelf. Temperate Northern W Atlantic. 3 species (Ngoc-Ho 2003; Poore et al. 2019).

Neotrypaea Manning & Felder, 1991 Diagnosis. Maxilliped 3 merus expanded distomesially as rounded lobe beyond articulation with ischium. Major cheliped merus of male with prominent truncate hook, serrate along lower margin, excavate laterally at base. Uropodal endopod asymmetrical, at least as wide as long, distal margin truncate-convex, at right angles to straight anterior margin; exopod distal margin clearly differentiated from anterior margin, anterodistal corner rightangled. Telson lateral margins convex; posterior margin slightly concave. Maximum cl. 21 mm (Pl. 16c). Estuarine, marine, intertidal–shelf. Tropical W Atlantic, Temperate Northern Pacific, Central Indo-Pacific, Tropical Eastern Pacific, Temperate South America. 16 species (Poore et  al.

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

2019). Japanese species have been treated as a separate genus, Nihonotrypaea, but this grades into Neotrypaea (Lin et al. 2007). The ecology of the Japanese species has been well studied (Tamaki et al. 2018; Tamaki et al. 1999).

Notiax Manning & Felder, 1991 Diagnosis. Rostrum acute, reaching middle of cornea, with ventral broad swelling. Major cheliped merus of male with prominent truncate hook, serrate along lower margin, excavate laterally at base. Uropodal endopod asymmetrical, at least as wide as long, distal margin truncate-convex, at right angles to straight anterior margin. Maximum cl. 10 mm. Subtidal. Temperate South America. 2 species (Poore et  al. 2019; Thatje 2000).

Paratrypaea Komai & Tachikawa, 2008 Diagnosis. Rostrum triangular, not reaching cornea or acute, as long as eyestalks. Major cheliped merus of male with blade dentate over lower margin, widest proximally; dorsodistolateral propodus and lateral dactylus with dense brush of setae. Uropodal exopod distal margin poorly differentiated from anterior margin, anterodistal corner rounded or clearly differentiated from anterior margin, anterodistal corner right-angled. Maximum cl. 7 mm (Pl. 16d, e). Intertidal–subtidal. Indo-West Pacific. 2 species (Dworschak 2012; Komai and Fujita 2014; Poore et al. 2019).

Poti Rodrigues & Manning, 1992 Diagnosis. Rostrum acute, as long as eyestalks. Antennular peduncle exceeded by antennal peduncular article 5. Maxilliped 3 merus almost rectangular, distally truncate with squarish angle between distal and mesial margins. Uropodal exopod distal margin clearly differentiated from anterior margin, anterodistal corner right-angled, 1.8 times as long as wide. Maximum cl. 4 mm. Shelf. Tropical W Atlantic. 1 species (Rodrigues and Manning 1992).

Praedatrypaea Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Rostrum acute, as long as eyestalks, with ventral keel. Pleonite 6 with sublateral ventral sharp ridge, flared posteriorly. Antennular peduncle exceeded by antennal peduncular article 5. Maxilliped 3 merus with spine on distal free margin. Major cheliped merus of male with blade dentate over lower margin, widest proximally. Uropodal exopod distal margin clearly differentiated from anterior margin, anterodistal corner right-angled, 1.5–1.8 times as long as wide. Telson proximolateral lobe prominent, defined posteriorly by clear unchitinised region; posterior margin truncate or slightly convex between posterolateral angles or slightly concave, with medial spine. Maximum cl. 5 mm. Shelf. Temperate Northern W Atlantic, Central Indo-Pacific. 4 species (Ngoc-Ho 1994: illustrations of 2 included species as Callianassa, C. propinqua De Man, 1905, C. praedatrix De Man, 1905; Poore et al. 2019).

213

Pugnatrypaea Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Rostrum acute, as long as eyestalks. Maxilliped 3 merus distally oblique with obtuse angle between distal and mesial margins. Major cheliped merus of male with simple proximal hook on lower margin. Uropodal exopod distal margin clearly differentiated from anterior margin, anterodistal corner rightangled, 1.5–1.8 times as long as wide. Telson posterior margin tapering over posterior third to pair of lobes separated by deep notch, with medial spine. Maximum cl. 4 mm (Pl. 16f). Slope. Western and Central Indo-Pacific. 8 species (Felder and Robles 2020a; De Man 1928: figures of type species; Poore et al. 2019; Liu 2022: key to species).

Rayllianassa Sakai & Tachikawa, 2008 Diagnosis. Hermaphrodite. Antennular peduncle exceeding antennal peduncle by about half length of article 3. Minor cheliped about two-thirds width of major cheliped, both swollen; carpus upper margin as long or shorter than propodus. Telson lateral margins convex; posterior margin slightly concave, sometimes with medial spine. Maximum cl. 7 mm. Subtidal. Indo-West Pacific. 5 species (Komai and Tachikawa 2008; Poore et al. 2019).

Rudisullianassa Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Hermaphrodite. Antennular peduncle exceeded by antennal peduncular article 5. Maxilliped 3 dactylus ovate, with dense brush of long setae over most of upper-distal margin, few setae along lower margin. Minor cheliped about two-thirds width of major cheliped, both swollen; carpus upper margin as long or shorter than propodus. Telson about as wide as long, tapering from anterolateral lobe; posterior margin slightly concave, sometimes with medial spine. Maximum cl. 3 mm. Subtidal. Temperate Northern W Pacific. 1 species (Komai et  al. 2014a: illustrations as Rayllianassa rudisulcus; Poore et  al. 2019).

Scallasis Bate, 1888 Diagnosis. Rostrum acute, as long as eyestalks. Pleonite 6 with sublateral ventral sharp ridge, flared posteriorly. Maxilliped 3 merus distally oblique with obtuse angle between distal and mesial margins. Major cheliped merus of male with 2 or 3 proximal similar teeth and distal denticles on lower margin. Uropodal endopod anterior margin unarmed or with spine at midpoint; exopod about as long as wide or ~1.5–1.8 times as long as wide. Telson lateral margins convex; posterior margin truncate or slightly convex between posterolateral angles or slightly concave, sometimes with medial spine; transverse ridge with fine setae and spiniform setae. Maximum cl. 5 mm (Pl. 16g). Subtidal–shelf. Temperate Northern W Pacific, Western and Central Indo-Pacific. 8 species (Komai et  al. 2020; Poore et  al. 2019).

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Marine Decapod Crustacea

Spinicallianassa Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Hermaphrodite. Rostrum acute, as long as eyestalks. Minor cheliped two-thirds width of major cheliped. Major cheliped merus of male with oblique spine about third or half way along lower margin; dactylus with dense setae along upper margin. Uropodal exopod distal margin clearly differentiated from anterior margin, anterodistal corner right-angled, 1.5–1.8 times as long as wide. Telson about as wide as long, tapering from anterolateral lobe; posterior margin slightly concave; transverse ridge sometimes with spiniform setae. Maximum cl. 7 mm. Subtidal–shelf. Temperate Northern W Pacific, Central IndoPacific. 4 species (Komai et  al. 2014b: illustrations as Cheramus spinicauda; Poore et al. 2019).

Tastrypaea Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Minor cheliped two-thirds width of major cheliped. Major cheliped merus of male with 1 or 2 small teeth just before midpoint. Uropodal endopod asymmetrical, at least as wide as long, distal margin truncate-convex, at right angles to straight anterior margin; exopod distal margin clearly differentiated from anterior margin, anterodistal corner right-angled. Telson lateral margins convex. Maximum cl. 5 mm. Subtidal. Temperate Australasia. 1 species (Poore et  al. 2019; Sakai 1999: as Callianassa poorei).

Trypaea Dana, 1852 Diagnosis. Antennular peduncle twice as long as antennal peduncle. Maxilliped 3 merus grossly expanded distomesially beyond articulation with ischium. Major cheliped merus of male with prominent truncate hook, serrate along lower margin, excavate laterally at base. Telson lateral margins convex; posterior margin slightly concave. Maximum cl. 21 mm. Intertidal–subtidal. Temperate Australasia. 1 species (Poore et al. 2019). The genus has been used to include many species but is now restricted to one species. Trypaea australiensis Dana, 1852 is abundant in south-eastern Australia where it is the basis of a recreational bait fishery (Contessa and Bird 2004). References Berkenbusch K, Rowden AA (1998) Population dynamics of the burrowing ghost shrimp Callianassa filholi on an intertidal sandflat in New Zealand (Decapoda: Thalassinidea). Ophelia 49, 55–69. doi:10.1080/00 785326.1998.10409373 Berkenbusch K, Rowden AA (2000) Latitudinal variation in the reproductive biology of the burrowing ghost shrimp Callianassa filholi (Decapopda, Thalassinidea). Marine Biology 136, 497–504. doi:10.1007/s002270050709 Biffar TA (1970) Three new species of callianassid shrimp (Decapoda, Thalassinidea) from the western Atlantic. Proceedings of the Biological Society of Washington 83, 35–50. Biffar TA (1971) The genus Callianassa (Crustacea, Decapoda, Thalassinidea) in south Florida, with keys to the Western Atlantic species. Bulletin of Marine Science 21, 637–715.

Biffar TA (1973) The taxonomic status of Callianassa occidentalis Bate, 1888 and C. batei Borradaile, 1903 (Decapoda, Callianassidae). Crustaceana 24, 224–230. doi:10.1163/156854073X00399 Bird FL (2000) Physicochemical and microbial properties of burrows of the deposit-feeding thalassinidean ghost shrimp Biffarius arenosus (Decapoda: Callianassidae). Estuarine, Coastal and Shelf Science 51, 279–291. doi:10.1006/ecss.2000.0676 Bird FL, Poore GCB (1999) Functional burrow morphology of Biffarius arenosus (Decapoda: Callianassidae) from southern Australia. Marine Biology 134, 77–87. doi:10.1007/s002270050526 Bird FL, Ford PW, Hancock GJ (1999) Effect of burrowing macrobenthos on the flux of dissolved substances across the water-sediment interface. Marine and Freshwater Research 50, 523–532. doi:10.1071/MF98059 Butler SN, Bird FL (2008) Temporal changes in burrow structure of the thalassinidean ghost shrimps Trypaea australiensis and Biffarius arenosus. Journal of Natural History 42, 2041–2062. doi:10.1080/00222930802254656 Contessa L, Bird FL (2004) The impact of bait-pumping on populations of the ghost shrimp Trypaea australiensis Dana (Decapoda: Callianassidae) and the sediment environment. Journal of Experimental Marine Biology and Ecology 304, 75–97. doi:10.1016/j.jembe.2003.11.021 De Man JG (1928) The Decapoda of the Siboga-Expedition. Part 7. The Thalassinidae and Callianassidae collected by the Siboga-Expedition with some remarks on the Laomediidae. Siboga-Expéditie 39a6, 1–187. de Vaugelas J, de Saint Laurent M (1984) Premières données sur l’écologie de Callichirus laurae de Saint Laurent sp. nov. (Crustacé Décapode Callianassidae): son action bioturbatrice sur les formations sédimentaires du golfe d’Aqaba (Mer Rouge). Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 6, 147–152. Dworschak PC (1998) The role of tegumental glands in burrow construction by two Mediterranean callianassid shrimps. Senckenbergiana Maritima 28, 143–149. doi:10.1007/BF03043145 Dworschak PC (2002) The burrows of Callianassa candida (Olivi, 1792) and C. whitei Sakai, 1999 (Crustacea: Decapoda: Thalassinidea). In The Vienna school of marine biology. A tribute to Jörg Ott. (Eds Bright M, Dworschak PC, Stachowitshch M) pp. 63–71. Facultas Universitätsverlag, Wien. Dworschak PC (2012) On the identities of Callianassa bouvieri Nobili, 1904, C. maldivensis Borradaile, 1904, and C. gravieri Nobili, 1905 (Crustacea: Decapoda: Callianassidae): a morphometric approach. Zootaxa 3149, 39–56. doi:10.11646/zootaxa.3149.1.2 Felder DL, Robles R (2020a) A new mud shrimp of the genus Pugnatrypaea from outer continental shelf waters of the northern Gulf of Mexico, commonly associated with hydrocarbon seeps (Crustacea: Decapoda: Callianassidae). Zootaxa 4732, 545–555. doi:10.11646/zootaxa.4732.4.3 Felder DL, Robles R (2020b) Two new burrowing mud shrimps of the genus Gilvossius from the Gulf of Mexico (Crustacea: Decapoda: Callianassidae). Nauplius 28, e2020018. doi:10.1590/2358-2936e2020018 Hernáez P, Miranda MS, Tavares M (2020) A new species of Biffarius Manning & Felder, 1991 (Decapoda: Axiidea: Callianassidae) from the intertidal coast of northeastern Brazil. Zootaxa 4759, 575–583. doi:10.11646/zootaxa.4759.4.8 Komai T, Fujita Y (2014) New record of a callianassid ghost shrimp Paratrypaea maldivensis (Borradaile, 1904) (Crustacea: Decapoda: Axiidea) from subtidal flats in Okinawa-jima Island, Ryukyu Islands, Japan. Fauna Ryukyuana 8, 1–7. Komai T, Fujiwara Y (2012) New records of callianassid ghost shrimp (Crustacea: Decapoda: Axiidea) from reducing environments in Kyushu, southwestern Japan. Zootaxa 3271, 55–67. doi:10.11646/ zootaxa.3271.1.5 Komai T, Tachikawa H (2008) Thalassinidean shrimps (Crustacea: Decapoda) from the Ogasawara Islands, Japan. Natural History Research 10, 19–52.

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Komai T, Fujita Y, Maenosono T (2014a) Additional record of Rayllianassa amboinensis (de Man, 1888) from Japan, and description of a new species from Okinawa Island, Ryukyu Islands (Crustacea: Decapoda: Axiidea: Callianassidae). Zootaxa 3835, 549–563. doi:10.11646/zootaxa.​ 3835.4.6 Komai T, Maenosono T, Fujita Y (2014b) Two new species of ghost shrimp assigned to the genus Cheramus Spence Bate, 1888 (Crustacea: Decapoda: Axiidea: Callianassidae) from the Ryukyu Islands, Japan. Zootaxa 3895, 503–524. doi:10.11646/zootaxa.3895.4.3 Komai T, Poore GCB, Fujita Y (2020) Redescription of the poorly known ghost shrimp species Scallasis amboinae Bate, 1888, review of the genus and description of a new species from the Ryukyu Islands, Japan (Decapoda: Axiidea: Callianassidae). Zootaxa 4766, 401–420. doi:10.11646/zootaxa.4766.3.1 Lin F-J, Komai T, Chan T-Y (2007) A new species of callianassid shrimp (Crustacea: Decapoda: Thalassinidea) from deep-water hydrothermal vents off Taiwan. Proceedings of the Biological Society of Washington 120, 143–158. doi:10.2988/0006-324X(2007)120[143:ANSOCS]2.0.CO;2 Liu W (2022) A new ghost shrimp of the genus Pugnatrypaea (Decapoda, Axiidea, Callianassidae) from the South China Sea. ZooKeys 1085, 183–192. doi:10.3897/zookeys.1085.79278 Manning RB, Felder DL (1992) Gilvossius, a new genus of callianassid shrimp from the eastern United States (Crustacea: Decapoda: Thalassinidea). Bulletin of Marine Science 49, 558–561. Nates SF, Felder DL, Lemaitre R (1997) Comparative larval development in two species of the burrowing ghost shrimp genus Lepidophthalmus (Decapoda: Callianassidae). Journal of Crustacean Biology 17, 497–519. doi:10.2307/1549444 Ngoc-Ho N (1994) Some Callianassidae and Upogebidae from Australia with description of four new species (Crustacea: Decapoda: Thalassinidea). Memoirs of the Museum of Victoria 54, 51–78. doi:10.24199/j. mmv.1994.54.02 Ngoc-Ho N (2003) European and Mediterranean Thalassinidea (Crustacea, Decapoda). Zoosystema 25, 439–555. Ngoc-Ho N (2014) Six species of Axiidea and Gebiidea from the IndoWest Pacific (Crustacea, Decapoda). Zoosystema 36, 545–561. doi:10.5252/z2014n3a1 Nickell LA, Atkinson RJA (1995) Functional morphology of burrows and trophic modes of three thalassinidean shrimp species, and a new approach to the classification of thalassinidean burrow morphology. Marine Ecology Progress Series 128, 181–197. doi:10.3354/meps128181 Pachelle PPG, Tavares M (2020) Axiidean ghost shrimps (Decapoda: Axiidae, Callianassidae, Callichiridae, Micheleidae) of the Trindade and Martin Vaz Archipelago, Vitória-Trindade Seamounts Chain and Abrolhos, off southeastern Brazil. Zootaxa 4758, 103–126. doi:10.11646/​ zootaxa.4758.1.4 Poore GCB (1975) Systematics and distribution of Callianassa (Crustacea, Decapoda, Macrura) from Port Phillip Bay, Australia, with descriptions of two new species. Pacific Science 29, 197–209. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Poore GCB, Dworschak PC, Robles R, Mantelatto FL, Felder DL (2019) A new classification of Callianassidae and related families (Crustacea: Decapoda: Axiidea) derived from a molecular phylogeny with morphological support. Memoirs of Museum Victoria 78, 73–146. doi:10.24199/j.mmv.2019.78.05 Robles R, Dworschak PC, Felder DL, Poore GCB, Mantelatto FL (2020) A molecular phylogeny of Callianassidae and related families (Crustacea: Decapoda: Axiidea) with morphological support. Invertebrate Systematics 34, 113–132. doi:10.1071/IS19021

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Rodrigues SdA, Manning RB (1992) Poti gaucho, a new genus and species of ghost shrimp from southern Brazil (Crustacea: Decapoda: Callianassidae). Bulletin of Marine Science 51, 9–13. Sakai K (1999) A new species, Callianassa poorei, sp. nov. (Decapoda: Crustacea: Callianassidae) from Tasmania. Journal of the Marine Biological Association of the United Kingdom 79, 373–374. doi:10.1017/ S0025315498000460 Sakai K (2002) Callianassidae (Decapoda, Thalassinidea) in the Andaman Sea, Thailand. Phuket Marine Biological Center Special Publication 23, 461–532. Sakai K (2005) Callianassoidea of the world (Decapoda: Thalassinidea). Crustaceana Monographs 4, 1–285. Sakai K (2011) Axioidea of the world and a reconsideration of the Callianassoidea (Decapoda, Thalassinidea, Callianassida). Crustaceana Monographs 13, 1–616. doi:10.1163/9789047424185 Stamhuis EJ, Schreurs CE, Videler JJ (1997) Burrow architecture and turbative activity of the thalassinid shrimp Callianassa subterranea from the central North Sea. Marine Ecology Progress Series 151, 155–163. doi:10.3354/meps151155 Stamhuis EJ, Dauwe B, Videler JJ (1998) How to bite the dust: morphology, motion pattern and function of the feeding appendages of the deposit-feeding thalassinid shrimp Callianassa subterranea. Marine Biology 132, 43–58. doi:10.1007/s002270050370 Stapleton KL, Long M, Bird FL (2001) Comparative feeding ecology of two spatially coexisting species of ghost shrimp; Biffarius arenosus and Trypaea australiensis (Decapoda: Callianassidae). Ophelia 55, 141–150. doi:10.1080/00785236.2001.10409481 Tamaki A, Ueno H (1998) Burrow morphology of two callianassid shrimps, Callianassa japonica Ortmann, 1891 and Callianassa sp. (=Callianassa japonica: De Man, 1928)(Decapoda: Thalassinidea). Crustacean Research 27, 28–39. doi:10.18353/crustacea.27.0_28 Tamaki A, Itoh J-I, Kubo K (1999) Distributions of thee species of Nihonotrypaea (Decapoda: Thalassinidea: Callianassidae) in intertidal habitats along an estuary it open-sea gradient in western Kyushu, Japan. Crustacean Research 28, 37–51. doi:10.18353/crustacea.28.0_37 Tamaki A, Itoh J-I, Hongo Y, Takeuchi S, Takikawa T (2018) Normal delayed establishment of a semilunar brooding and larval release cycle in the course of the reproductive season of the ghost shrimp population on a warm temperate intertidal sandflat. Journal of Shellfish Research 37, 529–570. doi:10.2983/035.037.0309 Thatje S (2000) Notiax santarita, a new species of the Callianassidae (Decapoda, Thalassinidea) from the Beagle Channel, southernmost America. Crustaceana 73, 289–299. doi:10.1163/156854000504381 Witbaard R, Duineveld GCA (1989) Some aspects of the biology and ecology of the burrowing shrimp Callianassa subterranea (Montagu) (Thalassinidea) from the southern North Sea. Sarsia 74, 209–219. doi: 10.1080/00364827.1989.10413430

Callianideidae Kossman, 1880 ghost shrimps Figure 9.14, Plate 16h, i Callianideid ghost shrimps resemble callianassids and similar families in having a short triangular rostrum, flat eyestalks and no interacting lobes between the carapace and pleon but differ in having a long seta attached to the posterior lobe of the scaphognathite (epipod of the maxilla) (Poore 2015; Poore et al. 2019). Micheleids also have this long seta but in members of that family the carapace and pleonite interact by

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means of a pair of lobes and the eyestalks are cylindrical. ­Callianideids are assumed to be burrowers in soft sediments but species of Thomassinia have a unique toothbrush-like maxilliped 3 that suggests feeding on hard surfaces.

Diagnosis. Posterior margin of carapace without lateral lobes, pleonite 1 without anterolateral lobes, weakly chitinised. Maxilla scaphognathite with long seta on posterior lobe extending into branchial chamber.

Key to genera of Callianideidae 1. – 2. – 3.

Pleopods 2–5 with marginal filaments (Fig. 9.14l.k). Chelipeds strongly dissimilar, minor cheliped cylindrical (Fig. 9.14f, g)����������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Pleopods 2–5 without marginal filaments. Chelipeds similar or nearly so, both with flattened articles (Fig. 9.14h)������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Pleopodal marginal filaments branching (Fig. 9.14k) or not. Male pleopod 1 of 1 article, distally setose, with thumb-like appendix interna (Fig. 9.14i)����������������������������������������������������������������������������������������������������� Callianidea Pleopodal marginal filaments simple (Fig. 9.14l). Male pleopod 1 of 1 or 2 indistinct articles, second triangular, appendix interna present as mesial row of hooks (Fig. 9.14j)������������������������������������������������� Paracallianidea Maxilliped 3 with toothbrush of stiff setae on ischium (Fig. 9.14e). Uropodal endopod with obliquetransverse row of spiniform setae (Fig. 9.14p)���������������������������������������������������������������������������������������������������Thomassinia

Fig. 9.14.  Callianideidae. Carapace, antennules, antennae, eyes: a, Crosniera; b, Heardaxius. Maxilla: c, Callianidea. Maxilliped 3: d, Crosniera; e, Thomassinia. Major, minor chelipeds: f, g, Callianidea. Cheliped: h, Heardaxius. Male pleopod 1: i, Callianidea. j, Paracallianidea. Pleopods 2–5, marginal filaments: k, Callianidea typa; l, Paracallianidea. Male pleopod 2: m, Heardaxius. Telson, uropod: n, Heardaxius; o, Mictaxius; p, Thomassinia.

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

– 4. – 5. –

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Maxilliped 3 with only tapering setae on ischium. Uropodal endopod without oblique-transverse row of spiniform setae (Fig. 9.14n, o)����������������������������������������������������������������������������������������������������������������������������������������������������4 Rostrum spike-like or sharply triangular (Fig. 9.14a). Maxillipedal 3 exopod as long as ischium (Fig. 9.14d)���������������������������������������������������������������������������������������������������������������������������������������������������������������������Crosniera Rostrum obsolete (Fig. 9.14b). Maxillipedal 3 exopod vestigial or absent�������������������������������������������������������������������������������5 Gonochoristic. Pleopod 2 with appendix interna scarcely exceeding fused appendix masculina. Uropodal exopod with dorsal plate or notch (Fig. 9.14o)�������������������������������������������������������������������������������������������������������� Mictaxius Hermaphroditic. Male pleopod 2 with appendix interna well exceeding fused appendix masculina. Uropodal exopod evenly ovate (Fig. 9.14n)���������������������������������������������������������������������������������������������������������� Heardaxius Callianidea H. Milne Edwards, 1837

Diagnosis. Linea thalassinica absent. Maxilliped 3 exopod reaching end of ischium. Male pleopod 1 present, appendix interna thumb-like, slightly projecting. Male pleopod 2 without appendix masculina, with small digitiform appendix interna. Pleopods 2–5 rami with tubular segmented marginal filaments, branching or not. Maximum cl. 14 mm (Pl. 16h, i). Intertidal, subtidal. Temperate Northern W Pacific, Indo-West Pacific. 2 species (Dworschak 2014).

Crosniera Kensley & Heard, 1991 Diagnosis. Rostrum spine-like or triangular. Linea thalassinica incomplete. Maxilliped 3 exopod reaching end of ischium. Male pleopod 1 of 1 simple article, or 2-articled, with distal triangular appendix interna. Male pleopod 2 with elongate appendix masculina with long-terminal setae, appendix interna digitiform, attached mesially. Shelf, slope. Tropical W Atlantic, Central Indo-Pacific. 5 species (Heard and King 2007; Poore 2015).

Heardaxius Sakai, 2011 Diagnosis. Hermaphroditic. Linea thalassinica present, reaching posterior margin or almost so. Maxilliped 3 exopod vestigial. Male pleopod 1 2-articled, with distal triangular appendix interna. Male pleopod 2 with elongate appendix masculina with long-terminal setae, appendix interna a fused thumb-like extension. Subtidal (coral sands). Central and Eastern Indo-Pacific. 2 species (Poore 2015).

Mictaxius Kensley & Heard, 1991 Diagnosis. Linea thalassinica present, reaching posterior margin or almost so. Maxilliped 3 exopod vestigial. Male pleopod 1 absent. Male pleopod 2 with appendices masculina and interna totally fused, with terminal stiff setae and apical hooks. Uropodal exopod with dorsal plate. Subtidal, shelf. Tropical W Atlantic, Eastern Indo-Pacific. 2 species (Poore 2015).

Paracallianidea Sakai, 1992 Diagnosis. Linea thalassinica incomplete. Maxilliped 3 exopod reaching end of ischium. Male pleopod 1 of 1 or 2 weakly

separated articles, distal article triangular, appendix interna represented by mesial row of hooks. Male pleopod 2 with elongate appendix masculina with long-terminal setae, appendix interna digitiform, attached mesially. Pleopods 2–5 rami with simple segmented tubular marginal filaments. Intertidal, subtidal. Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Poore 2015).

Thomassinia de Saint Laurent, 1979 Diagnosis. Linea thalassinica present, reaching posterior margin or almost so. Maxilliped 3 with brush of stiff non-tapering setae on ischium and merus; exopod absent. Male pleopod 1 absent. Male pleopod 2 without appendix masculina, with elongate appendix interna. Subtidal. Indo-West Pacific. 3 species (Poore 1997). References Dworschak PC (2014) The Axiidea (Crustacea, Decapoda) of Cocos (Keeling) and Christmas Islands, with description of a new species of Eucalliax Manning & Felder, 1991. Raffles Bulletin of Zoology Supplement 30, 230–245. Heard RW, King RA (2007) Crosniera wennerae, a new species of thalassinidean (Crustacea: Decapoda: Thomassiniidae) from the South Atlantic Bight. Zootaxa 1644, 41–49. doi:10.11646/zootaxa.1644.1.2 Poore GCB (1997) A review of the thalassinidean families Callianideidae Kossmann, Micheleidae Sakai, and Thomassiniidae de Saint Laurent (Crustacea: Decapoda) with descriptions of fifteen new species. Zoosystema 19, 345–420. Poore GCB (2015) Rediagnosis of Callianideidae and its genera (Crustacea: Decapoda: Axiidea), and description of a new species of Heardaxius Sakai, 2011. Zootaxa 3995, 229–240. doi:10.11646/zootaxa.3995.1.19 Poore GCB, Dworschak PC, Robles R, Mantelatto FL, Felder DL (2019) A new classification of Callianassidae and related families (Crustacea: Decapoda: Axiidea) derived from a molecular phylogeny with morphological support. Memoirs of Museum Victoria 78, 73–146. doi:10.24199/j.mmv.2019.78.05

Callianopsidae Manning & Felder, 1991 ghost shrimps Figure 9.15 Callianopsids comprise a small family of burrowing ghost shrimps that shares with Eucalliacidae a dilating, truncate dactylus on maxilliped 3, with dense field of setae on its distal margin. They differ in lacking a dorsal plate on the

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uropodal exopod, and having spines or a small proximal tooth on the lower margin of the merus of the cheliped (Poore et al. 2019). They are assumed to be burrowers in soft sediments but unlike most callianassids and similar families they are found at lower shelf or slope depths. Two

species in two of the three genera are associated with chemoautotrophic communities. Diagnosis. Maxilliped 3 dactylus broad, with dense field of setae on truncate distal margin (Fig.  9.15e). Uropodal exopod without elevated dorsal plate (Fig. 9.15b, c).

Key to genera of Callianopsidae 1. – 2. –

Pleonite 6 with prominent lateral projections (Fig. 9.15b). Carapace with longitudinal carina running from rostrum. Epipods absent���������������������������������������������������������������������������������������������������������������������������������������� Callianopsis Pleonite 6 without prominent lateral projections. Carapace without longitudinal carina. Epipod present on maxilliped 3 to pereopod 4��������������������������������������������������������������������������������������������������������������������������������������������������������2 Two cardiac sulci present (Fig. 9.15a). Maxilliped 3 with rudimentary exopod (Fig. 9.15d)����������������������������Bathycalliax Cardiac sulci absent. Maxilliped 3 exopod absent (Fig. 9.15e)������������������������������������������������������������������������� Vulcanocalliax Bathycalliax Sakai and Türkay, 1999

Vulcanocalliax Dworschak & Cunha, 2007

Diagnosis. Carapace, longitudinal carina absent; cardiac sulci present. Rostrum obsolete, flat, not reaching cornea. Major cheliped merus of male lower margin straight, with proximal row of denticles. Maximum cl. 13 mm. Slope. Temperate Northern E Pacific. 1 species. The only species Bathycalliax geomar Sakai and Türkay, 1999 was collected from a chemoautotrophic community associated with a subduction zone at 627 m (Poore et al. 2019; Sakai and Türkay 1999).

Diagnosis. Carapace, longitudinal carina absent; cardiac sulci absent. Rostrum obsolete or obtusely triangular, flat, not reaching cornea. Major cheliped merus of male with 1–2 small proximal teeth on lower margin. Maximum cl. 14 mm. Slope (1339 m). Temperate Northern E Atlantic (mud volcano).1 species (Dworschak and Cunha 2007; Poore et  al. 2019). The genus is also known from New Zealand.

Callianopsis de Saint Laurent, 1973 Diagnosis. Carapace, longitudinal carina strong on rostrum, weaker more posteriorly; cardiac sulci absent. Rostrum acute, produced, with dorsal ridge leading to gastric region. Major cheliped merus of male with weekly-toothed squarish blade with proximal erect spine on lower margin (also on minor cheliped). Maximum cl. 16 mm. Shelf–slope. Temperate Northern Atlantic, Temperate Northern Pacific, Western Indo-Pacific. 4 species (Dworschak and Poore 2018; Lin et al. 2007; Poore et al. 2019).

References Dworschak PC, Cunha MR (2007) A new subfamily, Vulcanocalliacinae n.subfam., for Vulcanocalliax arutyunovi n. gen., n. sp. from a mud volcano in the Gulf of Cádiz (Crustacea, Decapoda, Callianassidae). Zootaxa 1460, 35–46. doi:10.11646/zootaxa.1460.1.3 Dworschak PC, Poore GCB (2018) More cautionary tales: family, generic and species synonymies of recently published taxa of ghost and mud shrimps (Decapoda, Axiidea and Gebiidea). Zootaxa 4394, 61–76. doi:10.11646/ zootaxa.4394.1.3 Lin F-J, Komai T, Chan T-Y (2007) First record of the thalassinidean genus Callianopsis de Saint Laurent, 1973 (Decapoda, Ctenochelidae)

Fig. 9.15.  Callianopsidae. Carapace: a, Bathycalliax. Pleonite 6, telson, uropod: b Callianopsis. Uropod: c, Vulcanocalliax. Maxilliped 3: d, Bathycalliax; e, Vulcanocalliax.

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in the West Pacific, with the description of a new species from Taiwan. Crustaceana 80, 1193–1203. doi:10.1163/156854007782321191 Poore GCB, Dworschak PC, Robles R, Mantelatto FL, Felder DL (2019) A new classification of Callianassidae and related families (Crustacea: Decapoda: Axiidea) derived from a molecular phylogeny with morphological support. Memoirs of Museum Victoria 78, 73–146. doi:10.24199/​ j.mmv.2019.78.05 Sakai K, Türkay M (1999) A new subfamily, Bathycalliacinae n. subfam., for Bathycalliax geomar n. gen., n. sp. from deep water cold seeps off Oregon, USA. Senckenbergiana Biologica 79, 203–209.

Callichiridae Manning & Felder, 1991 ghost shrimps Figures 9.16–9.18, Plate 16i–q Callichirid ghost shrimps inhabit estuarine and marine sediments from the intertidal to slope depths. The family is cosmopolitan in temperate and tropical regions (Dworschak 2000a, 2005). They build permanent burrows with inhalent and exhalent openings on the seafloor (Abed-Navandi et al. 2005; Dworschak 2000b; Dworschak et al. 2006) or inhabit burrows under rocks (Sepahvand et al. 2014). The shrimps leave their burrows to collect plant detritus and sediment that they sequester in chambers in the burrows. The bioengineering effects of this behaviour in the biology and chemistry of sediments was extensive reviewed by Pillay and Branch (2011) and Dworschak et al. (2012). Callichirids differ from the other large callianassoid family, Callianassidae, in the shape of the anterior

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branchiostegal margin of the carapace. In callichirids the anterior branchiostegal lobe is sclerotised and articulates with the anterodorsal margin of the branchiostegite whereas in callianassids this anterior margin is simple or with a slight notch. Callichirids usually have well developed male pleopods 1 and 2, and pleopod 2 of both sexes is broad, not styliform as in callianassids. The epipod of maxilliped 1 has an acute anterior lobe as in all axiideans except callianassids (Poore et al. 2019). Keys to species in Sakai (2011) are useful for some genera but his higher taxonomy differs markedly from that of the most recent revision (Poore et al. 2019). Diagnosis. Carapace anterior branchiostegal margin interrupted by an oblique sclerotised ridge; anterior branchiostegal lobe sclerotised, well produced anteriorly beyond junction with oblique branchiostegal ridge which it meets level with linea thalassinica or with which it articulates by means of a virtual condyle. Scaphocerite rudimentary. Maxilliped 3 propodus about as wide as long or longer than wide. Uropodal exopod with elevated dorsal plate. Implicit generic attributes. Carapace anterolateral spines absent. Eyestalk flattened, contiguous. Antennular peduncle shorter than antennal peduncle. Maxilliped 3 exopod absent. Pereopod 3 propodus with strong broadly-rounded proximal lobe on lower margin, lower margin straight-­concave. Male pleopod 1 of 2 articles. Male pleopod 2 appendix interna digitiform.

Key to genera of Callichiridae 1. – 2. – 3. – 4. – 5. –

Eyestalk cylindrical, separate, with terminal spherical cornea, without terminal lobe (Fig. 9.16b). Maxilliped 3 merus with 3 spines along distal margin (Fig. 9.16m)���������������������������������������������������������� Calliapagurops Eyestalk flattened, contiguous, about twice as long as wide, with cornea subterminal and more or less elevated, usually with terminal lobe (Fig. 9.16a, c–k). Maxilliped 3 merus with 1 or without spines along distal margin (Fig. 9.16n–q)������������������������������������������������������������������������������������������������������������������������������������������������������2 Rostrum spine-like, anteriorly or upwardly directed. Anterolateral margins of carapace with conical spine, sometimes weakly calcified at base (Fig. 9.16d–f)������������������������������������������������������������������������������������������������������������������3 Rostrum obsolete or triangular, flat even if sharp. Anterolateral margins of carapace not produced (Fig. 9.16a, g–k) or rarely sharp (in Neocallichirus vigilax, Lepidophthalmus tridentatus)���������������������������������������������4 Major chelipeds with 3 spines on upper inner margin of merus and propodus, and 4–6 on lower margin of carpus (Fig. 9.17c). Rostrum with midventral ridge (Fig. 9.16e, f). Male pleopod 2 with partially articulating appendix masculina (Fig. 9.18n)��������������������������������������������������������������������������������������������������������� Glypturus Major chelipeds without spines on upper margin of merus and propodus or lower margin of carpus (Fig. 9.17a). Rostrum without midventral ridge (Fig. 9.16d). Male pleopod 2 with appendix masculina fused to endopod (Fig. 9.18o)������������������������������������������������������������������������������������������������������������������������������Corallianassa Maxilliped 3 exopod present, minute (Fig. 9.16n, o)�������������������������������������������������������������������������������������������������������������������5 Maxilliped 3 exopod absent������������������������������������������������������������������������������������������������������������������������������������������������������������8 Antennular peduncle much longer than antennal peduncle (Fig. 9.16h). Telson posterior margin with medial lobe (Fig. 9.18h)��������������������������������������������������������������������������������������������������������������������������������������������������������������6 Antennular peduncle shorter than antennal peduncle (Fig. 9.16e). Telson posterior margin convex or concave�������������� 7

220

Marine Decapod Crustacea

Male major cheliped merus with sinuous lower margin, widest proximally and excavate beyond, or with proximal tubercle bearing spine(s). Pereopod 3 propodus lower margin with obtuse notch between distal lobe and heel (Fig. 9.17m)������������������������������������������������������������������������������������������������������������������� Lepidophthalmus – Male major cheliped merus with denticulate blade, more prominent over proximal half of lower margin. Pereopod 3 propodus lower margin simply concave����������������������������������������������������������������������������������� Kraussillichirus 7. Male major cheliped fingers longer than square palm, with quadrant-shaped tooth in gape (Fig. 9.17d). Telson widest proximally, tapering to convex posterior margin (Fig. 9.18g)����������������������������������������Karumballichirus – Male major cheliped fingers shorter than rectangular palm (Fig. 9.17b). Telson widest at midpoint, posterior margin concave (Fig. 9.18f)����������������������������������������������������������������������������������������������������������������� Glypturoides 8. Antennular peduncle shorter or as long as antennal peduncle (Fig. 9.16a, j, k)���������������������������������������������������������������������9 – Antennular peduncle significantly longer than antennal peduncle���������������������������������������������������������������������������������������13 9. Major cheliped merus lower margin with simple proximal hook, dactylus massive (Fig. 9.17g). Maxilliped 3 dactylus with expanded setose apex (Fig. 9.16q)����������������������������������������������������������� Thailandcallichirus – Major cheliped merus lower margin denticulate or with toothed blade, dactylus not significantly broadened (Fig. 9.17f). Maxilliped 3 dactylus tapering�������������������������������������������������������������������������������������������������������10 10. Rostrum spine-like, thickened. Cornea subdistal, hemispherical, elevated, with minute mesiodistal plate (Fig. 9.16k). Female pleopod 2 endopod 3 times as long as wide��������������������������������������������������������Mucrollichirus – Rostrum flat, triangular. Cornea mid-distal, scarcely elevated, with obvious mesiodistal plate (Fig. 9.16j). Female pleopod 2 endopod 4 times as long as wide����������������������������������������������������������������������������������������11 11. Telson with convex parallel lateral margins, with obtuse angled apex, domed dorsally (Fig. 9.18i). Maxilliped 3 propodus longer than wide, free distal margin oblique. Male pleopod 1 of 1 slender article (Fig. 9.18l)��������Mocallichirus – Telson widest proximally or at midpoint (Fig. 9.18d, j). Maxilliped 3 propodus as long as wide, free distal margin nearly transverse. Male pleopod 1 of 2 articles (Fig. 9.18m)��������������������������������������������������������������������� 12 12. Telson converging over most of length from subproximal width to about half maximum width, with rounded posterolateral corners, posterior margin narrow (Fig. 9.18j). Pereopod 3 propodus with proximal lobe on lower margin slightly overlapping carpus (Fig. 9.17l)��������������������������������������������Neocallichirus 6.

Fig. 9.16.  Callichiridae. Rostrum, eyestalks, antennules, antennae: a, Audacallichirus; b, Calliapagurops; c, Callichirus; d, Corallianassa; e, f, Glypturus; g, Karumballichirus; h, i, Lepidophthalmus; j, Mocallichirus; k, Mucrollichirus. Maxilliped 1: l, Callichirus. Maxilliped 3: m, Calliapagurops; n, Glypturoides; n, Karumballichirus; p, Mocallichirus; q, Thailandcallichirus.

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

221

Fig. 9.17.  Callichiridae. Male major pereopod 1: a, Corallianassa; b, Glypturoides; c, Glypturus; d, Karumballichirus; e, Laticallichirus; f, Mucrollichirus; g, Thailandcallichirus. Minor pereopod 1: h, Balsscallichirus; i, Laticallichirus. Pereopod 3: j, Audacallichirus; k, Karumballichirus; m, Lepidophthalmus; n, Mucrollichirus; l, Neocallichirus.



Telson converging from subproximal width to about three-quarters maximum width, with strongly convex lateral margins and rounded distolateral corners, posterior margin wide, sinuous (Fig. 9.18d). Pereopod 3 propodus with prominent proximal lobe on lower margin appearing to overlap carpus (Fig. 9.17j)�����������������������������������������������������������������������������������������������������������������������������������������������������������Audacallichirus 13. Uropodal endopod strap-like, curved, longer than exopod. Pleonites 3–5 with dorsal pattern of symmetrical grooves. telson almost semicircular, thickened, with constriction on proximal region, with posterolateral swellings (Fig. 9.18a)�������������������������������������������������������������������������������������������������������������������������������������������������� Callichirus – Uropodal endopod ovate. Pleonites 3–5 without pattern of symmetrical grooves. telson convex-sided, widest proximally or near midpoint��������������������������������������������������������������������������������������������������������������������������������������14 14. Uropodal exopod about as wide as length of anterior margin. distal margin of telson convex, upper surface with long setae only (Fig. 9.18b)�������������������������������������������������������������������������������������������������������������Grynaminna – Uropodal exopod much wider than length of anterior margins. Distal margin of telson concave, upper surface usually with transverse row of short spiniform setae (Fig. 9.18e, c, q)����������������������������������������������������15 15. Minor cheliped palm about third as wide as major cheliped palm, carpus longer than wide, fingers straight, shorter than palm (Fig. 9.17h)����������������������������������������������������������������������������������������������������������Balsscallichirus – Minor cheliped palm at least half as wide as major cheliped palm, carpus as long as wide, fingers curved, pincer-like, much longer than palm (Fig. 9.17i)�������������������������������������������������������������������������������������������������������������������16 16. Pleonites 3–5 with anteriorly converging longitudinal grooves and oblique–transverse rows of long setae (Fig. 9.18c). Male major cheliped merus with lower margin almost straight. Female pleopod 2 uniramous (Fig. 9.18p)����������������������������������������������������������������������������������������������������������������������������� Michaelcallianassa – Pleonites 3–5 with lateral tufts of setae. Male major cheliped merus with convex denticulate blade (Fig. 9.17e). Female pleopod 2 biramous (Fig. 9.18q)������������������������������������������������������������������������������������� Laticallichirus

222

Marine Decapod Crustacea

Fig. 9.18.  Callichiridae. Pleon, telson, uropods: a, Callichirus; b, Grynaminna; c, Michaelcallianassa. Telson, uropod: d, Audacallichirus; e, Balsscallichirus; f, Glypturoides; g, Karumballichirus; h, Lepidophthalmus; i, Mocallichirus; j, Neocallichirus. Male pleopod 1: k, Glypturus; l, Mocallichirus; m, Neocallichirus. Male pleopod 2: n, Glypturus; o, Corallianassa. Female pleopod 2: p, Balsscallichirus; q, Laticallichirus. Pleopod 3: r, Callichirus.

Audacallichirus Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Maxilliped 3 ischium and merus 2–3 times as long as wide. Major cheliped merus of male lower margin convex or blade like, especially proximally, with small denticles. Male pleopod 1 article 2 distally expanded, with shallow apical notch between 2 rounded setose distal lobes. Male pleopod 2 appendix interna absent or obsolete distomesial lobe. Telson convex-sided, widest near midpoint; posterior margin trilobed, excavate each side of medial lobe. Maximum cl. 17 mm. Subtidal. Tropical W Atlantic, Central Indo-Pacific. 4 species (Ngoc-Ho 2014: description as Neocallichirus audax (De Man, 1911); Poore et al. 2019).

Balsscallichirus Sakai, 2011 Diagnosis. Antennular peduncle longer than antennal peduncle. Maxilliped 3 ischium and merus linear, 3 or more times as long as wide. Male major cheliped merus with lower margin almost straight or with prominent proximal denticulate blade-like tooth on lower margin. Male pleopod 2 appendix interna absent. Telson with transverse row of robust setae. Maximum cl. 10 mm. Intertidal–shelf. Tropical E Atlantic, Western Indo-Pacific, Temperate Southern Africa. 6 species. The genus has many synonyms (Poore et al. 2019). Its species live on the African coast, as far

north as Mauritania on the west, the entire east coast, and north to Pakistan.

Calliapagurops de Saint Laurent, 1973 Diagnosis. Rostrum spine-like. Carapace anterolateral spines prominent. Eyestalk cylindrical, with round terminal cornea. Maxilliped 3 ischium and merus length broad, less than twice as long as wide at their articulation. Male major cheliped merus with row of sharp oblique spines, usually 2 proximally and 1 or more along length of lower margin. Telson with transverse row of robust setae. Maximum cl. 18 mm. Subtidal, shelf. Tropical E Atlantic, Central Indo-Pacific. 1 species (Poore et al. 2019). Calliapagurops charcoti de Saint Laurent, 1973 is, unusually, a probable suspension feeder (Dworschak and Wirtz 2010).

Callichirus Stimpson, 1866 Diagnosis. Pleonites 3–5 with symmetrical pattern of deep curved transverse and longitudinal grooves plus paired tufts of dense setae, pleonite 6 with pair of deep longitudinal grooves on wide swollen anterior section, defined by lateral concavity. Uropodal endopod strap-like, curved, posterior margin concave. Telson thickened, with constriction on proximal region, with

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

posterolateral swellings; posterior margin with medial notch at end of longitudinal groove. Maximum cl. 30 mm (Pl. 16j). Estuarine, marine, intertidal, subtidal. Tropical Atlantic, Temperate South America, Temperate Southern Africa. 4 species (Poore et al. 2019). Species of Callichirus are among the best studied of all ghost shrimps, particularly in the Americas. Ecological studies have covered inter alia reproduction (Peiró et  al. 2014), population dynamics (Bilodeau et al. 2005; Botter-Carvalho et al. 2007) and the effects of salinity (Strasser and Felder 2001).

Corallianassa Manning, 1987 Diagnosis. Rostrum spine-like. Carapace anterolateral spines prominent. Major cheliped merus of male with row of sharp oblique spines, usually 2 proximally and 1 or more along length of lower margin. Male pleopod 1 article 2 ovoid, with shallow apical notch between rounded subdistal lobe and curved acute apex. Male pleopod 2 appendix interna free, articulating, distal on mesial endopod margin. Telson converging to about half basal width over most of length to rounded posterolateral corners; posterior margin straight or with slight medial lobe. Maximum cl. 15 mm (Pl. 16k). Subtidal. Tropical Atlantic, Indo-West Pacific, Temperate ­Australasia. 13 species (Komai et al. 2015: rediagnosis; Poore et al. 2019: rediagnosis). The permanent U-shaped burrows of C. longiventris (A. Milne-Edwards, 1870) can be up to 1.5 m deep (Dworschak et al. 2006).

223

Grynaminna Poore, 2000 Diagnosis. Antennular peduncle longer than antennal peduncle. Maxilliped 3 ischium and merus moderately long. Male major cheliped merus with convex tuberculate blade on most of lower margin. Male pleopod 1article 2 ovoid, with shallow apical notch between rounded subdistal lobe and curved acute apex. Male pleopod 2 appendix interna free, articulating, distal on mesial endopod margin. Telson converging to half basal over most of length to rounded posterolateral corners. Maximum cl. 17 mm. Subtidal. Central Indo-Pacific (Ryuku Is, Japan). 1 species (Poore 2000; Poore et al. 2019).

Karumballichirus Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Rostrum spine-like. Maxilliped 3 exopod present. Male major cheliped merus lower margin straight, with proximal row of denticles; gape in chela of both sexes with quadrant-shaped tooth. Male pleopod 1 article 2 ovoid, with shallow apical notch between rounded subdistal lobe and curved acute apex. Male pleopod 2 appendix interna free, articulating, distal on mesial endopod margin. Telson with convex lateral margins and rounded posterolateral corners. Maximum cl. 25 mm. Estuarine, marine, intertidal. Western and Central IndoPacific. 1 species (Dworschak 2008: illustrations; Poore et  al. 2019).

Glypturoides Sakai, 2011

Kraussillichirus Poore, Dworschak, Robles, Mantelatto & Felder, 2019

Diagnosis. Maxilliped 3 ischium and merus length broad, less than twice as long as wide at their articulation; exopod present. Male major cheliped merus lower margin blade like, especially proximally, with small denticles. Male pleopod 1 article 2 ovoid, with shallow apical notch between rounded subdistal lobe and curved acute apex. Telson posterior margin concave between rounded posterolateral corners. Maximum cl. 20 mm (Pl. 16l). Subtidal. Tropical W Atlantic. 1 species (Corsetti and Strasser 2003: population biology; Dobbs and Guckert 1988: ecology; Poore et al. 2019: rediagnosis).

Diagnosis. Antennular peduncle longer than antennal peduncle. Maxilliped 3 ischium and merus length broad, less than twice as long as wide at their articulation; exopod present. Male major cheliped merus lower margin with tuberculate blade, prominent proximally. Male pleopod 2 appendix interna absent. Telson with convex lateral margins and rounded posterolateral corners; posterior margin convex, with medial lobe. Maximum cl. 15 mm. Estuarine, marine, intertidal. Temperate Southern Africa. 1 species (Poore et al. 2019; Stebbing 1900: description of type species, Callianassa kraussi Stebbing, 1900).

Glypturus Stimpson, 1866

Laticallichirus Komai, Yokooka, Henmi & Itani, 2019

Diagnosis. Rostrum spine-like, with midventral ridge. Carapace anterolateral spines prominent. Cornea subdistal, hemispherical, elevated, with minute mesiodistal plate. Male major cheliped merus with row of sharp oblique spines, usually 2 proximally and 1 or more along length of lower margin; with 3 spines on upper inner margin of merus and propodus, and 4–6 on lower margin of carpus. Telson strongly diverging to beyond midpoint and semicircular distally; posterior margin broadly semicircular. Maximum cl. 34 mm (Pl. 16m, n). Intertidal, subtidal, shelf. Tropical Atlantic, Indo-West Pacific. 3 species (Komai et al. 2015; Poore et al. 2019). Species are common deep burrows in sand flats of coral reef lagoons where the shrimps are active herbivores (Poore and Suchanek 1988).

Diagnosis. Rostrum spine-like. Antennular peduncle longer than antennal peduncle. Maxilliped 3 ischium and merus less than twice as long as wide. Major cheliped merus of male lower margin with evenly convex tuberculate blade. Male pleopod 1 article 2 longer than wide, with slight apical notch. Male pleopod 2 appendix interna absent. Telson with convex lateral margins and rounded posterolateral corners; posterior margin concave between rounded posterolateral corners; transverse ridge with pairs of spiniform setae. Maximum cl. 18 mm. Estuarine. Temperate Northern W Pacific. 1 species. The type species was first described as a Pleistocene fossil but later discovered alive and redescribed in more detail (Komai et al. 2019; Poore et al. 2019).

224

Marine Decapod Crustacea

Lepidophthalmus Holmes, 1904 Diagnosis. Rostrum spine-like. Antennular peduncle longer than antennal peduncle. Maxilliped 3 dactylus curved, 3 times as long as wide, concave below; exopod present. Male major cheliped merus with sinuous lower margin, widest proximally, excavate beyond, or with proximal tubercle bearing spine(s) with or without more distal teeth along lower margin. Minor cheliped fingers usually with dense tufts of setae filling gape of fingers. Pereopod 3 propodus lower margin with obtuse notch between distal lobe and heel. Telson posterior margin trilobed, sometimes weakly. Maximum cl. 10 mm (Pl. 16o). Estuarine, marine, intertidal, subtidal. Temperate Northern Pacific, Tropical Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific. 17 species (Komai et al. 2018: redescription; Poore et al. 2019; Robles and Felder 2015: phylogeny).

Michaelcallianassa Sakai, 2002 Diagnosis. Pleonites 3–5 with anteriorly converging longitudinal grooves and oblique-transverse rows of long setae. Antennular peduncle longer than antennal peduncle. Male major cheliped merus with lower margin almost straight. Female pleopod 2 uniramous. Telson converging to about half basal width from near anterior width over most of length to rounded posterolateral corners; with transverse row of robust setae. Maximum cl. 10 mm. Shelf. Temperate Northern W Pacific, Central Indo-Pacific. 2 species (Liu and Liu 2009: second species; Poore et al. 2019).

Mocallichirus Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Maxilliped 3 ischium and merus length linear, 4 or more times as long as wide at their articulation. Male major cheliped merus lower margin with tuberculate blade, prominent proximally. Male pleopod 1 of 1 article. Telson with convex parallel lateral margins, with broadly rounded apex, domed; posterior margin broadly semicircular. Maximum cl. 10 mm. Subtidal. Western Indo-Pacific. 1 species (Poore et al. 2019).

Mucrollichirus Poore, Dworschak, Robles, Mantelatto & Felder, 2019 Diagnosis. Rostrum spine-like. Male major cheliped merus lower margin convex or blade like, especially proximally, with small denticles. Pereopod 3 propodus oval, lower margin convex, not proximally lobed (Fig. 9.18e). Male pleopod 1 article 2 ovoid, with shallow apical notch between rounded subdistal lobe and curved acute apex. Female pleopod 2, endopod 2–3 times as long as wide. Telson converging to about half basal width from near anterior width over most of length. Maximum cl. 10 mm (Pl. 16p). Subtidal. Central Indo-Pacific. 1 species (Poore et al. 2019).

Neocallichirus Sakai, 1988 Diagnosis. Male major cheliped merus lower margin convex or blade like, especially proximally, with small denticles. Uropodal

endopod with straight or slightly convex anterior margin ending in rounded angle, transverse or near-transverse distal margin continuous with curved posterior margin, as wide or wider than long. Telson converging to about half basal width over most of length. Maximum cl. 24 mm (Pl. 16q). Marine, intertidal, subtidal. Widespread in temperate and tropical seas. 33 species (Dworschak 2011a, 2011b; Hernáez et al. 2020; Poore et al. 2019).

Thailandcallichirus Sakai, 2011 Diagnosis. Rostrum spine-like. Maxilliped 3 exopod absent. Male major cheliped merus with proximal tubercle bearing spine with more distal teeth along lower margin, with massive propodus. Minor cheliped fingers with dense tufts of setae filling gape of fingers. Male pleopod 1 article 2 tapering, with small acute subdistal lobe, acute distal apex. Telson tapering from greatest width near base. Maximum cl. 9 mm. Estuarine, marine, intertidal. Central Indo-Pacific. 1 species (Poore et al. 2019; Sakai 1983: description of type species, Callianassa (Callichirus) ranongensis Sakai, 1983). References Abed-Navandi D, Koller H, Dworschak PC (2005) Nutritional ecology of thalassinidean shrimps constructing burrows with debris chambers: The distribution and use of macronutrients and micronutrients. Marine Biology Research 1, 202–215. doi:10.1080/​17451000510019123 Bilodeau AL, Felder DL, Neigel JE (2005) Population structure at two geographic scales in the burrowing crustacean Callichirus islagrande (Decapoda, Thalassinidea): historical and comtemporary barriers to planktonic dispersal. Evolution 59, 2125–2138. doi:10.1111/j.0014-3820.​ 2005.tb00923.x Botter-Carvalho ML, Santos PJP, Carvalho PVVC (2007) Population dynamics of Callichirus major (Say, 1818) (Crustacea, Thalassinidea) on a beach in northeastern Brazil. Estuarine, Coastal and Shelf Science 71, 508–516. doi:10.1016/j.ecss.2006.09.001 Corsetti JL, Strasser KM (2003) Population biology of the ghost shrimp Sergio trilobata (Biffar, 1970) (Crustacea: Decapoda: Thalassinidea). Gulf and Caribbean Research 15, 13–19. doi:10.18785/gcr.1501.03 Dobbs FC, Guckert JB (1988) Callianassa trilobata (Crustacea: Thalassinidea) influences abundance of meiofauna and biomass, composition, and physiologic state of microbial communities within its burrow. Marine Ecology Progress Series 45, 69–79. doi:10.3354/meps045069 Dworschak PC (2000a) Global diversity in the Thalassinidea (Decapoda). Journal of Crustacean Biology 20, 238–245. doi:10.1163/1937240X-90000025 Dworschak PC (2000b) On the burrows of Lepidophthalmus louisianensis (Schmitt, 1935) (Decapoda: Thalassinidea: Callianassidae). Senckenbergiana Maritima 30, 99–104. doi:10.1007/BF03042959 Dworschak PC (2005) Global diversity of Thalassinidea (Decapoda): an update (1998–2004). Nauplius 13, 57–63. Dworschak PC (2008) Neocallichirus kempi Sakai, 1999, a junior synonym of Callianassa karumba Poore & Griffin, 1979 (Decapoda: Callianassidae). Raffles Bulletin of Zoology 56, 75–84. Dworschak PC (2011a) Redescription of Callianassa jousseaumei Nobili, 1904, a junior subjective synonym of Callianassa indica de Man, 1905 with description of a new species of Neocallichirus (Decapoda: Axiidea: Callianassidae). Zootaxa 2746, 1–19. doi:10.11646/zootaxa.​2746.1.1 Dworschak PC (2011b) Redescription of Callianassa vigilax De Man, 1916, a subjective senior synonym of Neocallichirus denticulatus

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

Ngoc-Ho, 1994 (Crustacea: Decapoda: Callianassidae). Annalen des Naturhistorischen Museums in Wien, B 112, 137–151. Dworschak PC, Wirtz P (2010) Discovery of the rare burrowing shrimp Calliapagurops charcoti de Saint Laurent, 1973 (Decapoda: Axiidea: Callianassidae) in shallow water: first record of the infraorder for Madeira Island. Zootaxa 2691, 53–56. doi:10.11646/zootaxa.2691.1.3 Dworschak PC, Koller H, Abed-Navandi D (2006) Burrow structure, burrowing and feeding behaviour of Coralllianassa longiventris and Pestarella tyrrhena (Crustacea, Thalassinidea, Callianassidae). Marine Biology 148, 1369–1382. doi:10.1007/s00227-005-0161-8 Dworschak PC, Felder DF, Tudge CC (2012) Chapter 69. Infraorders Axiidea de Saint Laurent, 1979 and Gebiidea de Saint Laurent, 1979 (formerly known collectively as Thalassinidea). In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea. Complementary to the volumes translated from the French of the Traité de Zoologie [founded by P.-P. Grassé]. (Eds Schram FR, von Vaupel Klein JC) pp. 109–219. Brill, Leiden. Hernáez P, Windsor AM, Paula CA, Santana W (2020) A new species of the ghost shrimp genus Neocallichirus Sakai, 1988 (Decapoda: Axiidea: Callianassidae) from the southwestern Atlantic. Marine Biology Research 16, 50–60. doi:10.1080/17451000.2019.1703003 Komai T, Maenosono T, Osawa M (2015) Records of three species of callianassid ghost shrimp from the genera Glypturus Stimpson, 1866 and Corallianassa Manning, 1987 (Crustacea: Decapoda: Axiidea) from the Ryukyu Islands, Japan, with remarks on the taxonomic status of the two genera. Fauna Ryukyuana 27, 13–59. Komai T, Osawa M, Maenosono T, Fujita Y, Naruse T (2018) Records of the callianassid ghost shrimp Lepidophthalmus tridentatus (von Martens, 1868) (Crustacea: Decapoda: Axiidea: Callianassidae) from the Ryukyu Islands, Japan. Fauna Ryukyuana 42, 9–27. Komai T, Yokooka H, Henmi T, Itani G (2019) A new genus for “Neocallichirus” grandis Karasawa & Goda, 1996, a ghost shrimp species (Decapoda: Axiidea: Callianassidae) heretofore known only by fossil materials. Zootaxa 4604, 461–481. doi:10.11646/zootaxa.​ 4604.3.4 Liu W, Liu J-Y (2009) Michaelcallianassa sinica sp. nov. (Crusacea, Decapoda, Thalassinidea, Callianassidae) from the South China Sea. Zootaxa 2294, 39–46. doi:10.11646/zootaxa.2294.1.3 Ngoc-Ho N (2014) Six species of Axiidea and Gebiidea from the Indo-West Pacific (Crustacea, Decapoda). Zoosystema 36, 545–561. doi:10.5252/ z2014n3a1 Peiró DF, Wehrtmann IS, Mantelatto FL (2014) Reproductive strategy of the ghost shrimp Callichirus major (Crustacea: Axiidea: Callianassidae) from the southwestern Atlantic: sexual maturity of females, fecundity, egg features, and reproductive output. Invertebrate Reproduction & Development 58, 294–305. doi:10.1080/07924259.2014.944672 Pillay D, Branch GM (2011) Bioengineering effects of burrowing thalassinidean shrimps on marine soft-bottom ecosystems. In: Oceanography and Marine Biology: An Annual Review (Eds Gibson RN, Atkinson RJA, Gordon JDM) pp. 137–192. CRC Press, Boca Raton. Poore GCB (2000) A new genus and species of callianassid ghost shrimp from Kyushu, Japan (Decapoda: Thalassinidea). Journal of Crustacean Biology 20 (Special Issue 2), 150–156. Poore GCB, Suchanek TH (1988) Glypturus motupore, a new callianassid shrimp (Crustacea: Decapoda) from Papua New Guinea with notes on its ecology. Records of the Australian Museum 40, 197–204. doi:10.385 3/j.0067-1975.40.1988.154

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Poore GCB, Dworschak PC, Robles R, Mantelatto FL, Felder DL (2019) A new classification of Callianassidae and related families (Crustacea: Decapoda: Axiidea) derived from a molecular phylogeny with morphological support. Memoirs of Museum Victoria 78, 73–146. doi:10.24199/j.mmv.2019.78.05 Robles R, Felder DF (2015) Molecular phylogeny of the genus Lepidophthalmus (Decapoda, Callianassidae), with re-examination of its species composition. Zootaxa 4020, 453–472. doi:10.11646/zootaxa.4020.3.3 Sakai K (1983) On a new species of the genus Callianassa (Crustacea, Decapoda) from Thailand. Crustacean Research 12, 111–115. doi:10.18353/rcustacea.12.0_111 Sakai K (2011) Axioidea of the world and a reconsideration of the Callianassoidea (Decapoda, Thalassinidea, Callianassida). Crustaceana Monographs 13, 1–616. doi:10.1163/9789047424185 Sepahvand V, Sari A, Tudge CC, Bolouki M (2014) A study of burrow morphology in representative axiidean and gebiidean mud shrimps, from the Persian Gulf and Gulf of Oman, Iran. Nauplius 22, 137–144. doi:10.1590/S0104-64972014000200007 Stebbing TRR (1900) South African Crustacea. Marine Investigations in South Africa. Department of Agriculture, Cape Town 2, 14–64, pls 1–4. Strasser KM, Felder DF (2001) Effects of salinity on development in the ghost shrimp Callichirus islagrande and two populations of C. major (Crustacea: Decapoda: Thalassinidea). Gulf and Caribbean Research 13, 9–18. doi:10.18785/gcr.1301.01

Ctenochelidae Manning & Felder, 1991 ghost shrimps Figure 9.19, Plate 17a–c Ctenochelids are a diverse family of burrowing ghost shrimps united only by the unusual dactylus on pereopod 5. None is common. Apart from general observations about habitats nothing is known of their ecology. Some are from shallow sediments but others are known only from claws captured at depth. We treat genera formerly included in Ctenochelidae s.s. and Gourretiidae together as differences between them are difficult to isolate (Poore et al. 2019; Robles et al. 2020). Keys to species in Sakai (2011) are useful for some genera but his taxonomy differs markedly from that of the most recent comprehensive revision (Poore et al. 2019). Diagnosis. Scaphocerite simple, triangular (Fig. 9.19g, i). Maxilliped 3 dactylus slender, digitiform (Fig. 9.19u). Pereopod 5 subchelate, dactylus a twisted plate longer than finger (Fig. 9.19t). Uropodal exopod without elevated dorsal plate, or with margin divided by notch. (Fig. 9.19a–f). Implicit generic attributes. Maxilliped 3 merus with 1 mesiodistal spine; exopod absent. Major cheliped palm little longer than wide. Minor cheliped more slender, tapering; propodus and fingers together evenly tapering, fingers curved distally, of equal lengths.

Key to genera of Ctenochelidae 1. –

Major chelipeds with palm swollen, with narrow pectinate fingers longer than palm (Fig. 9.19k, l)���������������������������������2 Major cheliped with flattened propodi, upper and lower margins usually ridged, fingers rarely longer than palm (Fig. 9.19m, n)��������������������������������������������������������������������������������������������������������������������������������������������������������������������3

226

2. – 3. – 4.

Marine Decapod Crustacea

Rostrum spine-like, carinate (Fig. 9.19g, h). Cornea flat, with weak pigmentation. Pleonites 1 and 2 together half as long as pleonites 3–5 (Fig. 9.19a)����������������������������������������������������������������������������������������������� Ctenocheles Rostrum obsolete, flat (Fig. 9.19i). Cornea disc-like, strongly pigmented. Pleonites 1 and 2 together as long as pleonites 3–5 (Fig. 9.19b)������������������������������������������������������������������������������������������������������������������������������ Ctenocheloides Telson longer than wide, parallel-sided over most of length, posterior margin truncate (Fig. 9.19f). Carapace with anterolateral spines (Fig. 9.19j). Maxilliped 3 exopod absent�������������������������������������������������Laurentgourretia Telson shorter or longer than wide, tapering over most of length to rounded apex (Fig. 9.19c). Anterolateral angles of carapace obsolete (Fig. 9.19i). Maxilliped 3 palp exopod present (Fig. 9.19u) or absent���������������������������������4 Pleonite 6 with triangular anterolateral projections. Telson broadly rounded over posterior two-thirds (Fig. 9.19c)������������������������������������������������������������������������������������������������������������������������������������������������������������������� Dawsonius

Fig. 9.19.  Ctenochelidae. Pleon: a, Ctenocheles; b, Ctenocheloides. Pleonite 6, telson, uropod: c, Dawsonius; d, Gourretia; e, Paragourretia; f, Laurentgourretia. Rostrum, eyestalks, antennules, antennae: g, h, Ctenocheles; i, Ctenocheloides; j, Laurentgourretia. Major cheliped: k, Ctenocheles; l, Ctenocheloides; m, Kiictenocheloides; n, Gourretia. Minor cheliped: o, Ctenocheloides; p, Kiictenocheloides; q, r, Paragourretia; s, Gourretia. Pereopod 5 fingers: t, Gourretia. Maxilliped 3: u, Gourretia. Male pleopod 1: v, Gourretia; w, Laurentgourretia. Male pleopod 2: x, Gourretia; y, Laurentgourretia.

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

– 5. – 6. –

227

Pleonite 6 usually without triangular lateral projections. Telson tapering over posterior half to two-thirds (Fig. 9.19d, e)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������5 Major cheliped merus lower margin with tubercle at midpoint (Fig. 9.19m). Minor cheliped palm swollen fingers pectinate, spines erect (Fig. 9.19p). Maxilliped 3 exopod absent����������������������������������������������� Kiictenocheloides Major cheliped merus lower margin with straight or weakly convex blade on lower margin bearing a short proximal curved spine and 1 or more distal spines (Fig. 9.19n). Minor cheliped palm tapering, fingers with or without teeth (Fig. 9.19r, s). Maxilliped 3 exopod present (Fig. 9.19u)�����������������������������������������������������������������6 Minor cheliped with sharp proximally-directed teeth along cutting edges of fingers (Fig. 9.19s). Major cheliped carpus lower proximal margin swollen (Fig. 9.19n). Uropodal exopod without marginal notch (Fig. 9.19d)�������������������������������������������������������������������������������������������������������������������������������������������������������������������� Gourretia Minor cheliped without sharp teeth along cutting edges of fingers (Fig. 9.19r). Major cheliped carpus lower proximal margin tapering. Uropodal exopod usually with marginal notch, slight dorsal plate (Fig. 9.19e)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Paragourretia Ctenocheles Kishinouye, 1926

Diagnosis. Carapace longitudinal carina strong on rostrum, weaker more posteriorly. Major cheliped with barrel-shaped propodus, elongate pectinate fingers. Minor cheliped narrower, with elongate fingers. Pereopod 3 propodus elongate-oval, tapering, without proximal lobe on lower margin. Maximum cl. 17 mm. Subtidal, shelf. Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Australasia. 7 species (Matsuzawa and Hayashi 1997; Poore and Dworschak 2017; Poore et al. 2019).

Ctenocheloides Anker, 2010 Diagnosis. Maxilliped 3 merus with mesial spine at midpoint. Major cheliped with barrel-shaped propodus, elongate pectinate fingers. Minor cheliped narrower, with elongate fingers. Pereopod 3 propodus elongate-oval, tapering, without proximal lobe on lower margin. Maximum cl. 4 mm (Pl. 17a). Subtidal. Tropical W Atlantic, Western and Central IndoPacific. 3 species (Poore 2015; Poore et al. 2019).

Dawsonius Manning & Felder, 1991 Diagnosis. Pleonite 6 with lateral projections. Minor cheliped propodus and fingers barely tapering, fingers evenly curved, unarmed. Pereopod 3 propodus triangular, with straight upper margin, expanded proximally on lower margin. Maximum cl. 15 mm (Pl. 17b). Subtidal. Tropical W Atlantic. 1 species (Manning and Felder 1991; Poore et al. 2019).

Gourretia de Saint Laurent, 1973 Diagnosis. Maxilliped 3 exopod present. Minor cheliped propodus and fingers together evenly tapering, fingers straight, dactylus longer, with sharp spines along opposing edges. Pereopod 3 propodus oval, little longer than wide, with short rounded proximal lobe on straight lower margin, upper margin strongly convex. Maximum cl. 6 mm. Subtidal, shelf. Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Western Indo-Pacific. 8 species (Poore et  al. 2019; Sakai 2011: key to 6 species). Gourretia and

Paragourretia have in the past been synonymised but they can be distinguished by the dentition of the tapering minor cheliped, shape of the propodus of pereopod 3 and of the uropodal exopod (Poore et al. 2019).

Kiictenocheloides Sakai, 2013 Diagnosis. Major cheliped broad, with short fingers. Minor cheliped narrower, with long pectinate fingers. Pereopod 3 propodus elongate-oval, tapering, without proximal lobe on lower margin. Maximum cl. 3 mm. Subtidal. Temperate Northern W Pacific. 1 species (Komai 2013 as Ctenocheloides nomurai Komai, 2013; Poore et al. 2019).

Laurentgourretia Sakai, 2004 Diagnosis. Carapace anterolateral spines prominent. Maxilliped 3 merus with 2 or more mesiodistal spines; exopod absent. Minor cheliped propodus and fingers barely tapering, fingers evenly curved, unarmed. Pereopod 3 propodus oval, little longer than wide, with short rounded proximal lobe on lower margin, lower margin straight, upper margin strongly convex. Maximum cl. 4 mm. Subtidal. Central Indo-Pacific. 1 species (Poore et  al. 2019; Sakai 2004).

Paragourretia Sakai, 2004 Diagnosis. Maxilliped 3 exopod present. Pereopod 3 propodus linear, longer than wide, without proximal lobe on straight lower margin, upper margin moderately convex. Maximum cl. 8 mm (Pl. 17c). Subtidal. Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific. 9 species (Poore et al. 2019; Sakai 2011). References Komai T (2013) A new species of the ghost shrimp family Ctenochelidae (Crustacea: Decapoda: Axiidea) from Japan. Species Diversity 18, 45–55. doi:10.12782/sd.18.1.045 Manning RB, Felder DL (1991) Revision of the American Callianassidae (Crustacea: Decapoda: Thalassinidea). Proceedings of the Biological Society of Washington 104, 764–792.

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Matsuzawa K, Hayashi K-I (1997) Male of Ctenocheles balssi (Crustacea, Decapoda, Callianassidae) from off Muroto Peninsula, Shikoku, Japan. Suisan Daigakko Kenkyu Hokoku 46, 39–46. Poore GCB (2015) Ctenocheloides boucheti n. sp., a new ghost shrimp from Papua New Guinea (Decapoda, Axiidea, Ctenochelidae). Zootaxa 3955, 142–146. doi:10.11646/zootaxa.3955.1.10 Poore GCB, Dworschak PC (2017) Family, generic and species synonymies of recently published taxa of ghost shrimps (Decapoda, Axiidea, Eucalliacidae and Ctenochelidae): cautionary tales. Zootaxa 4294, 119–125. doi:10.11646/zootaxa.4294.1.6 Poore GCB, Dworschak PC, Robles R, Mantelatto FL, Felder DL (2019) A new classification of Callianassidae and related families (Crustacea: Decapoda: Axiidea) derived from a molecular phylogeny with morphological support. Memoirs of Museum Victoria 78, 73–146. doi:10.24199/j.mmv.2019.78.05 Robles R, Dworschak PC, Felder DL, Poore GCB, Mantelatto FL (2020) A molecular phylogeny of Callianassidae and related families (Crustacea: Decapoda: Axiidea) with morphological support. Invertebrate Systematics 34, 113–132. doi:10.1071/IS19021 Sakai K (2004) Dr. R. Plante’s collection of the families Callianassidae and Gourretiidae (Decapoda, Thalassinidea) from Madagascar, with the description of two new genera and one new species of the Gourretiidae Sakai, 1999 (new status) and two new species of the Callianassidae Dana, 1852. Crustaceana 77, 553–601. doi:10.1163/​1568540041718019 Sakai K (2011) Axioidea of the world and a reconsideration of the Callianassoidea (Decapoda, Thalassinidea, Callianassida). Crustaceana Monographs 13, 1–616. doi:10.1163/9789047424185

Eucalliacidae Manning & Felder, 1991 ghost shrimps Figures 9.20, 9.21, Plate 17d–f Eucalliacids are a family of ghost shrimps that shares with Callianopsidae a dilated, truncate dactylus on maxilliped 3, with a dense field of setae on its distal margin. They differ in possessing a dorsal plate on the uropodal exopod, and a smooth lower margin of the merus of the cheliped (Poore et  al. 2019). The shrimps inhabit permanent burrows usually on intertidal sandflats or shallow soft sediment but species of one genus inhabit shelf and slope depths. Diagnosis. Rostrum flat, triangular, shorter than eyestalks (Fig.  9.20a, b). Scaphocerite rudimentary. Maxilliped 3 propodus about as wide as long; dactylus broad, with dense field of setae on truncate distal margin (Fig. 9.21a–f). Uropodal exopod with elevated dorsal plate (Fig. 9.20d–g). Implicit generic attributes. Cardiac sulci absent. Major cheliped palm little longer than wide, minor cheliped more slender, tapering. Telson smooth dorsally.

Key to genera of Eucalliacidae 1.

– 2. –

3. – 4. – 5.

Major cheliped much larger than minor cheliped. Minor chela with fixed finger acute, shorter than dactylus (Fig. 9.20l, k). Pereopod 3 propodus oval, little longer than wide, with short rounded proximal lobe on lower margin, lower margin straight, upper margin strongly convex (Fig. 9.20n). Female pleopod 2 without appendix interna (Fig. 9.21u)�������������������������������������������������������������������������������������������������������������������2 Both chelipeds swollen and similar, minor slightly smaller if at all. Pereopod 3 propodus pentagonal, with strong broadly-rounded proximal lobe on lower margin, lower margin concave (Fig. 9.20m). Female pleopod 2 with or without appendix interna��������������������������������������������������������������������������������������������������������������������������3 Sternite 7 and pereopod 5 coxa-ischium glabrous. Maxilliped 3 ischium linear; exopod absent (Fig. 9.21b). Minor cheliped with tooth in gape between fingers (Fig. 9.20k). Male pleopod 1 article 2 sickle-shaped (Fig. 9.21g)�������������������������������������������������������������������������������������������������������������������������������������������������������������������������Calliax Sternite 7 and pereopod 5 coxa-ischium densely setose (Fig. 9.20c). Maxilliped 3 ischium with strong proximal lobe on lower margin, wider proximally than distally; exopod present (Fig. 9.21f). Minor cheliped with fixed finger shorter than dactylus (Fig. 9.20l). Male pleopod 1 stout, with apical notch (Fig. 9.21o)��������������������������������������������������������������������������������������������������������������������������������������������������������������Pseudocalliax Uropodal endopod triangular, anterior margin strongly convex, posterior margin straight (Fig. 9.20f). Cheliped carpus upper margin with 2 distal sharp spines (Fig. 9.20i). Male pleopod 2 appendix masculina 4 times as long as wide, attached near apex of endopod; appendix interna minute (Fig. 9.21q)������������������������Eucalliax Uropodal endopod more or less ovate (Fig. 9.20d, e). Cheliped carpus upper margin without 2 distal sharp spines. Male pleopod 2 appendix masculina longer than wide, attached mesially to endopod with appendix interna, shorter than or up to twice as long as endopod apex (Fig. 9.21p, r, s)������������������������������������������������4 Male pleopod 2 appendix masculina a lobe fused with appendix interna, subdistal on endopod margin; endopod broadly triangular (Fig. 9.21s)����������������������������������������������������������������������������������������������������������� Paraglypturus Male pleopod 2 appendix masculina a lobe fused with appendix interna, attached midway on endopod margin; endopod longer than wide (Fig. 9.21p, r)������������������������������������������������������������������������������������������������������������������5 Telson with transverse ridge; lateral margins convex (Fig. 9.20e)������������������������������������������������������������������� Eucalliaxiopsis

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

229

Fig. 9.20.  Eucalliacidae. Carapace: a, Calliaxina. Anterior carapace, eyestalk, antennule, antenna: b, Calliaxina. Posterior carapace, sternite 7, pleopod 1, coxa 4, basis of pereopod 5: c, Pseudocalliax. Telson, uropod: d, Calliaxina; e, Eucalliaxiopsis. Pleonite 6, telson, uropod: f, Eucalliax; g, Paraglypturus. Major cheliped: h, Paraglypturus; i, Eucalliax. Minor cheliped; j, Paraglypturus; k, Calliax; l, Pseudocalliax. Pereopod 3; m, Calliaxina; n, Calliax.

– 6. –

Telson without transverse ridge; lateral margins tapering (Fig. 9.20d)�����������������������������������������������������������������������������������6 Maxilliped 3 ischium and merus length less than twice as long as wide, without proximal lobe (Fig. 9.21c, d)����������������������������������������������������������������������������������������������������������������������������������������������������������������Calliaxina Maxilliped 3 ischium and merus length twice as long as wide, with strong proximal lobe on lower margin, wider proximally than distally (Fig. 9.21a)������������������������������������������������������������������������������������������������� Andamancalliax Andamancalliax Sakai, 2011

Diagnosis. Maxilliped 3 crista dentata a prominent toothed ridge extending beyond proximal margin of merus. Major cheliped palm little longer than wide, minor cheliped more slender, tapering. Male pleopod 1 article 1 with distal long setae, article 2 bladelike, with bifid apex. Telson widest proximally, curving to broad convex apex. Maximum cl. 3.5 mm. Shelf (31–158 m). Central Indo-Pacific. 2 species (Poore 2021; Sakai 2002).

Calliax de Saint Laurent, 1973 Diagnosis. Major chela of male with palm about twice as long as carpus, 1.5 times as long as wide, minor chela half as wide, with short fingers. Male pleopod 1 article 2 curved, simply curving to

sharp apex. Telson widest proximally, more or less semicircular or curving to broad convex apex. Maximum cl. 6 mm. Subtidal–slope (2–800 m). Temperate Northern Atlantic (Mediterranean), Western Indo-Pacific. 3 species (García Raso et al. 2019; Ngoc-Ho 2014; Poore et al. 2019).

Calliaxina Ngoc-Ho, 1973 Diagnosis. Carapace cardiac sulci present. Both chelipeds with palms as long as wide in both sexes, minor slightly smaller if at all. Male pleopod 1 article 1 with distal long setae, article 2 blade like, with bifid apex, sometimes obscure or article 1 with distal long setae, article 2 much shorter, digitiform (sometimes fused). Telson widest proximally, more or less semicircular or curving to broad convex apex. Maximum cl. 12 mm (Pl. 17d).

230

Marine Decapod Crustacea

Fig. 9.21.  Eucalliacidae. Maxilliped 3: a, Andamancalliax; b, Calliax; c, d, Calliaxina; e, Eucalliaxiopsis; f, Pseudocalliax. Male pleopod 1: g, Calliax; h–k, Calliaxina; l, m, Eucalliaxiopsis; n, Paraglypturus; o, Pseudocalliax. Male pleopod 2: p, Calliaxina; q, Eucalliax; r, Eucalliaxiopsis; s, Paraglypturus. Female pleopod 1: t Paraglypturus. Female pleopod 2: u, Calliax; v, Calliaxina; w, Paraglypturus. Female pleopod 3: x, Paraglypturus.

Intertidal, subtidal. Temperate Northern Atlantic, Temperate Northern Pacific, Indo-West Pacific. 5 species. The male pleopod 1 is highly variable (Fig. 9.21h–k) with forms overlapping those of Eucalliaxiopsis (Fig. 9.21l, m) (Poore 2021).

convex-sided, widest near midpoint; with transverse crest. Maximum cl. 20 mm. Subtidal. Tropical W Atlantic. 1 species (Manning and Felder 1991; Poore et al. 2019).

Eucalliax Manning & Felder, 1991

Eucalliaxiopsis Sakai, 2014

Diagnosis. Major chela of male with palm about twice as long as carpus, 1.5 times as long as wide; carpus upper margin with 2 distal sharp spines. Minor chela slightly smaller, with elongate fingers Male pleopod 1 article 1 with distal long setae, article 2 blade like, with bifid apex, sometimes obscure. Telson

Diagnosis. Carapace cardiac sulci present. Major chela of male with palm about twice as long as carpus, 1.5 times as long as wide, minor chela slightly smaller, with elongate fingers or both chelipeds with palms as long as wide in both sexes, minor slightly smaller if at all (rarely). Male pleopod 1 article 2 longer than

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

article 1, parallel-sided, setose or article 1 with distal long setae, article 2 blade like, with bifid apex, sometimes obscure or article 1 with distal long setae, article 2 much shorter, digitiform (sometimes fused). Telson convex-sided, widest near midpoint; with transverse crest. Maximum cl. 20 mm (Pl. 17e, f). Estuarine, marine, intertidal, subtidal. Tropical W Atlantic, Indo-West Pacific, Temperate Australia. 10 species (Poore 2021).

Paraglypturus Türkay & Sakai, 1995 Diagnosis. Major chela of male with palm about twice as long as carpus, 1.5 times as long as wide. Minor chela slightly smaller, with elongate fingers. Male pleopod 1 article 1 with distal long setae, article 2 blade like, with bifid apex, sometimes obscure. Telson widest proximally, more or less semicircular or curving to broad convex apex. Maximum cl. 10 mm. Shelf, slope (chemotrophic environments at 114, 544 m). Temperate Northern Pacific, Eastern Indo-Pacific. 2 species. Both were reported with a pigment patch on one of the two uropodal rami, but not the same one (Ahn et al. 2017; Poore et al. 2019; Türkay and Sakai 1995).

Pseudocalliax Sakai, 2014 Diagnosis. Thoracic sternite 7 and pereopod 5, coxa-ischium densely setose. Major chela of male with palm about twice as long as carpus, 1.5 times as long as wide, minor chela half as wide, with short fingers. Male pleopod 1 stout, article 1 setose, article 2 with shallow apical notch. Telson widest proximally, more or less semicircular or curving to broad convex apex. Maximum cl. 6 mm. Intertidal, subtidal. Temperate Australia. 1 species (Poore 2021). References Ahn D-H, Kim S-J, Ju S-J, Min G-S (2017) A new species of Paraglypturus (Crustacea: Decapoda: Axiidea: Callianassidae) from a vent field in the Tonga Arc of the south-western Pacific Ocean. Journal of the Marine Biological Association of the United Kingdom 97, 105–111. doi:10.1017/S0025315416000084 García Raso JE, García-Muñoz JE, Mateo-Ramírez A, López González N, Fernández-Salas LM, et al. (2019) Decapod crustaceans Eucalliacidae in chemoautotrophic bathyal bottoms of the Gulf of Cadiz (Atlantic Ocean), environmental characteristics and associated communities. Journal of the Marine Biological Association of the United Kingdom 99, 437–444. doi:10.1017/S0025315418000280 Manning RB, Felder DL (1991) Revision of the American Callianassidae (Crustacea: Decapoda: Thalassinidea). Proceedings of the Biological Society of Washington 104, 764–792.

231

Ngoc-Ho N (2014) Six species of Axiidea and Gebiidea from the IndoWest Pacific (Crustacea, Decapoda). Zoosystema 36, 545–561. doi:10.5252/z2014n3a1 Poore GCB (2021) Indo-West Pacific and Australian species of Eucalliacidae with descriptions of four new species (Crustacea, Axiidea). Memoirs of Museum Victoria 80, 1–41. Poore GCB, Dworschak PC, Robles R, Mantelatto FL, Felder DL (2019) A new classification of Callianassidae and related families (Crustacea: Decapoda: Axiidea) derived from a molecular phylogeny with morphological support. Memoirs of Museum Victoria 78, 73–146. doi:10.24199/j.mmv.2019.78.05 Sakai K (2002) Callianassidae (Decapoda, Thalassinidea) in the Andaman Sea, Thailand. Phuket Marine Biological Center Special Publication 23, 461–532. Türkay M, Sakai K (1995) Decapod crustaceans from a volcanic hot spring in the Marianas. Senckenbergiana Maritima 26, 25–35.

Micheleidae Sakai, 1992 ghost shrimps Figure 9.22 The Micheleidae are essentially a family of four genera (Poore 1997, 2020; Poore and Collins 2015). Many species are associated with coral environments where they are probably weak burrowers. The combination of ‘setal-rows’ and indistinct cervical groove are diagnostic. Setal rows are tight regular rows of short plumose setae, each seta sited in a pit-like structure, just behind the anterior margin of the carapace, on propodi of pereopods 2 and 3 and variously over the pleon (Kensley and Heard 1991). Setal rows are also seen in Strahlaxiidae, Callianideidae and some Ctenochelidae. Unlike these families, micheleids have equal chelipeds with cylindrical propodi and fine fingers (Fig. 9.22j, k). Diagnosis. Rostrum flat, triangular, shorter than eyestalks (Fig. 9.22a, e), or flat, thick, apically rounded, exceeding eyestalks (Fig.  9.22b–d, f, g). Carapace cervical groove indistinct; posterior margin of carapace with lateral lobes interacting with anterolateral lobes on pleonite 1 (Fig. 9.22a). Eyestalks cylindrical, with terminal subspherical cornea (Fig. 9.22a–g). Scaphocerite simple, triangular (Fig. 9.22b–f). Maxilla scaphognathite with long seta on posterior lobe extending into branchial chamber. Implicit generic characters. Rostrum prominent, flat, longer than eyestalks. Antennular article 1 approximately half length of entire peduncle. Pleopods 2–5 without marginal lamellae.

Key to genera of Micheleidae 1. –

Rostrum minute, triangular (Fig. 9.22e). Pleurobranchs usually absent above pereopod 2–4. Pleopodal lamellae usually present (Fig. 9.22m). Pereopods 3 and 4 with lateral spiniform setae (Fig. 9.22l)�����������������Michelea Rostrum prominent, flat (Fig. 9.22a–c, e, f). Pleurobranchs usually present above pereopod 2–4. Pleopodal lamellae absent. Pereopods 3 and 4 without lateral spiniform setae�����������������������������������������������������������������������������������2

232

Marine Decapod Crustacea

Fig. 9.22.  Micheleidae. Carapace: a, Michelea. Carapace, antennules, antennae, eyes: b, c, Marcusiaxius; d, Meticonaxius; e, Michelea; f, g, Tethisea. Pleonite 6, telson, uropod: h, Marcusiaxius; i, Tethisea. Cheliped: j, Marcusiaxius; k, Tethisea. Pereopod 3: l, Michelea. Pleopod 3: m, Michelea.

2. – 3. –

Uropodal endopod with anterior margin convex, ending by curving to rounded posterior margin. Pleonites 3–5 without setal-rows (Fig. 9.22i)������������������������������������������������������������������������������������������������������������������� Tethisea Uropodal endopod with straight anterior margin ending sharply. Pleonites 3–5 with setal-rows (Fig. 9.22h)����������������3 Eyes visible in dorsal view, rostrum narrowly rounded or acute (Fig. 9.22d). Cheliped fixed finger with major tooth two-thirds way along. Uropodal endopod at most 1.5 times as long as wide������������������������Meticonaxius Eyes not visible in dorsal view, rostrum broadly rounded, bullet-shaped (Fig. 9.22b, c). Cheliped fixed finger with major tooth one-third way along (Fig. 9.22j). Uropodal endopod about twice as long as wide (Fig. 9.22h)��������������������������������������������������������������������������������������������������������������������������������������������������������������Marcusiaxius Marcusiaxius Rodrigues & de Carvalho, 1972

Diagnosis. Eyestalks not visible in dorsal view, obscured by wide rostrum. Cheliped fixed finger with major tooth one-third way along. Uropodal exopod with straight anterior margin ending sharply, deeply curved posteriorly. Maximum cl. 21 mm. Shelf. Tropical Atlantic, Central Indo-Pacific. 6 species (Poore 1997; Poore and Collins 2015).

Meticonaxius De Man, 1905 Diagnosis. Eyestalks visible in dorsal view, rostrum narrow. Cheliped fixed finger with major tooth two-thirds way along. Uropodal exopod with straight anterior margin ending sharply, deeply curved posteriorly. Maximum cl. 17 mm.

Shelf–slope. Temperate Northern and Tropical Atlantic, Western and Central Indo-Pacific, Temperate Southern Africa. 12 species (Lin 2006; Poore 2020).

Michelea Kensley & Heard, 1991 Diagnosis. Rostrum triangular, minute. Antennular article 1 elongate, about three-quarters of entire peduncle. Cheliped fixed finger without or with insignificant teeth. Pleopods 2–5 with marginal flat lamellae (usually). Uropodal exopod oval. Maximum cl. 9 mm. Subtidal–shelf. Tropical Atlantic, Temperate Northern Pacific, Central Indo-Pacific, Temperate Australasia. 16 species (Liu and Liu 2012; Poore and Collins 2015).

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

Tethisea Poore, 1994 Diagnosis. Eyestalks not visible in dorsal view, obscured by wide rostrum. Cheliped fixed finger without or with insignificant teeth. Uropodal exopod with anterior margin concave, curving to rounded posterior margin. Maximum cl. 10 mm (Pl. 17g). Shelf, slope. Western and Central Indo-Pacific. 2 species (Poore and Collins 2015).

233

Diagnosis. Rostrum flat, triangular, shorter than eyestalks; median carina on rostrum only. Posterior margin of carapace with lateral lobes interacting with anterolateral lobes on pleonite 1 (Fig. 9.23a). Scaphocerite simple, triangular. Uropodal exopod without elevated dorsal plate (Fig. 9.23b). Paracalliax de Saint Laurent, 1979

References Kensley B, Heard RW (1991) An examination of the shrimp family Callianideidae (Crustacea: Decapoda: Thalassinidea). Proceedings of the Biological Society of Washington 104, 493–537. Lin F-J (2006) Two new axioids (Decapoda: Thalassinidea) from New Caledonia. Journal of Crustacean Biology 26, 234–241. doi:10.1651/C-2592.1 Liu W, Liu J-Y (2012) Michelea takeda sp. nov. (Crustacea, Decapoda, Axiidea, Micheleidae) from the South China Sea. Crustaceana Monographs 17, 175–181. Poore GCB (1997) A review of the thalassinidean families Callianideidae Kossmann, Micheleidae Sakai, and Thomassiniidae de Saint Laurent (Crustacea: Decapoda) with descriptions of fifteen new species. Zoosystema 19, 345–420. Poore GCB (2020) Axiid and micheleid lobsters from Indo-West Pacific deep-sea environments (Crustacea: Decapoda: Axiidea: Axiidae, Micheleidae). In: Corbari L, Chan T-Y, Ahyong ST (Eds) Deep-sea crustaceans from Papua New Guinea. Tropical Deep-Sea Benthos vol. 31. Mémoires du Muséum National d’Histoire Naturelle, Paris 213, 259–367. Poore GCB, Collins DJ (2015) Micheleidae (Crustacea: Decapoda: Axiidea): new family and generic synonymies, three new Australian species, and new records. Memoirs of Museum Victoria 73, 95–105. doi:10.24199/j.mmv.2015.73.08

Paracalliacidae Sakai, 2005 Figure 9.23 The only species differs from all other callianassoids in having pleonite 1 with dorsolateral lobes interacting and overlapping the posterolateral margin of the carapace and in pleopod 2 being similar to pleopods 3–5 (Fig. 9.23d, e) (Poore et al. 2019).

Maximum cl. 21 mm. Shelf (20–369 m). Temperate Northern E Atlantic. 1 species. The only species Paracalliax bollorei de Saint Laurent, 1979 is from the shelf off Mauritania (de Saint Laurent 1979) and has been collected once since (Dworschak and Poore 2018). References de Saint Laurent M (1979) Note 14, in de Saint Laurent, M. and Le Loeuff, P., Campagnes de la Calypso au large des côtes Atlantiques Africaines (1956 et 1959) (suite) 22. Crustacés Décapodes Thalassinidea. I. Upogebiidae et Callianassidae. In: Forest J (Ed.), Résultats Scientifiques des Campagnes de la Calypso. Fasc. 11 (22). Annales de l’Institut Océanographique, Monaco et Paris 55 Suppl., 95. Dworschak PC, Poore GCB (2018) More cautionary tales: family, generic and species synonymies of recently published taxa of ghost and mud shrimps (Decapoda, Axiidea and Gebiidea). Zootaxa 4394, 61–76. doi:10.11646/zootaxa.4394.1.3 Poore GCB, Dworschak PC, Robles R, Mantelatto FL, Felder DL (2019) A new classification of Callianassidae and related families (Crustacea: Decapoda: Axiidea) derived from a molecular phylogeny with morphological support. Memoirs of Museum Victoria 78, 73–146. doi:10.24199/j.mmv.2019.78.05

Strahlaxiidae Poore, 1994 mud shrimps Figure 9.24, Plate 17h–j Strahlaxiids are intertidal and shallow-water shrimps distinguished from axiids by the triangular rather than

Fig. 9.23.  Paracalliacidae, Paracalliax bollorei de Saint Laurent, 1979. a, posterior carapace, pleonites 1, 2; b, telson, uropod; c, maxilliped 3; d, e, female pleopods 1, 2.

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elongate exopods on pleopods 3–5 and a bifid rostrum. The family is cosmopolitan in temperate and tropical regions. Diagnosis. Rostrum triangular, apically bifid, with lateral teeth; median carina present as ridge or row of spines

extending over gastric region. Posterior margin of carapace with lateral lobes interacting with anterolateral lobes on pleonite 1. Eyestalks cylindrical, with terminal subspherical cornea. Scaphocerite elongate.

Key to genera of Strahlaxiidae 1. – 2. –

Scaphocerite with at least 1 spine on lower margin, sometimes laterally spinose (Fig. 9.24b). Telson with 2 or 3 transverse ridges (Fig. 9.20f). Pleurobranchs present above pereopod 2–4�������������������������������������������������� Neaxius Scaphocerite simple, straight (Fig. 9.20c). Telson smooth dorsally (Fig. 9.24g). Pleurobranchs absent above pereopod 2–4�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Rostrum rounded-triangular, with obscure marginal teeth (Fig. 9.24d), Male pleopod 1 simple��������������������Strahlaxius Rostrum almost semicircular, with narrow apical notch, with denticulate margin (Fig. 9.24a). Male pleopod 1 absent���������������������������������������������������������������������������������������������������������������������������������������������������������Neaxiopsis Neaxiopsis Sakai & de Saint Laurent, 1989

Diagnosis. Rostrum almost semicircular, with narrow apical notch, with denticulate margin. Scaphocerite simple, straight. Pleurobranchs absent from pereopods 2–4. Pleopod 1 of male absent. Telson smooth dorsally. Maximum cl. 28 mm. Subtidal–shelf. Tropical W Atlantic, Western and Central Indo-Pacific. 2 species (Poore and Dworschak 2018).

Neaxius Borradaile, 1903 Diagnosis. Rostrum narrow, apex with pair of spines separated by U-shaped notch. Scaphocerite with at least 1 spine on lower margin, sometimes more and sometimes laterally spinose. Pleurobranchs present above pereopod 2–4. Pleopod 1 of male

absent. Telson with 2 transverse ridges. Maximum cl. 40 mm (Pl. 17h). Intertidal, subtidal. Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific. 7 species (Ngoc-Ho 2006; Poore and Dworschak 2018). The best known species, N. acanthus (A. Milne-Edwards, 1878), inhabits coral reef areas (Kensley et al. 2000; Kneer et al. 2008).

Strahlaxius Sakai & de Saint Laurent, 1989 Diagnosis. Rostrum rounded-triangular, with obscure marginal teeth. Scaphocerite simple, straight. Pleurobranchs absent from pereopods 2–4. Pleopod 1 of male 1 simple article. Telson smooth dorsally. Maximum cl. 23 mm (Pl. 17i, j). Intertidal–subtidal. Central and Eastern Indo-Pacific, Temperate Australia. 2 species (Poore 2004).

Fig. 9.24.  Strahlaxiidae. Anterior carapace, antennules, antennae, eyes: a, Neaxiopsis; b, c, Neaxius; d, e, Strahlaxius. Pereopod 3: f, Neaxius. Pleopod 3: g, Neaxiopsis. Telson, uropod: h, Neaxiopsis; i, Neaxius; j, Strahlaxius.

9 – Axiidea – sponge shrimps, ghost shrimps and mud lobsters

References Kensley B, Lin F-J, Yu H-P (2000) Further records of thalassinidean shrimps from Taiwan (Decapoda: Axiidae and Calocarididae), with descriptions of three new species. Journal of Crustacean Biology 20, 207–217. doi:10.1163/1937240X-90000022 Kneer D, Asmus H, Vonk JA (2008) Seagrass as the main food source of Neaxius acanthus (Thalassinidea: Strahlaxiidae), its burrow associates, and of Corallianassa coutierei (Thalassinidea: Callianassidae). Estuarine, Coastal and Shelf Science 79, 620–630. doi:10.1016/j.ecss.2008.05.013

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Ngoc-Ho N (2006) A new species of Neaxius Borradaile, 1903 from the Gulf of Guinea (Crustacea, Decapoda, Thalassinidea, Strahlaxiidae). Zoosystema 28, 409–415. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Poore GCB, Dworschak PC (2018) The Indo-West Pacific species of Neaxiopsis and Neaxius (Crustacea: Axiidea: Strahlaxiidae). Memoirs the Museum Victoria 77, 15–28. doi:10.24199/j.mmv.2018.77.02

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10. Gebiidea – mud lobsters and sponge shrimps

Gebiideans are small lobsters with symmetrical chelate or subchelate first pereopods and subchelate or simple second pereopods. Gebiidea de Saint Laurent, 1979 is the second of two infraorders until recently considered part of Thalassinidea Latreille, 1831 (Poore et  al. 2014; Robles et  al. 2009). The other is Axiidea (Chapter 9). The two infraorders are probable sister taxa or not, according to the molecular data used (Timm and Bracken-Grissom 2015). Gebiideans are separated from axiideans in the possession of enlarged chelate or subchelate pereopods 1 and subchelate or simple pereopods 2 (Dworschak et al. 2012; de Saint Laurent 1979). Most species are cryptic burrowers in mud, sponges or corals. The infraorder comprises three diverse families with 20 genera, most in the family Upogebiidae. Axianassidae is treated as a synonym of Laomediidae.

Diagnosis. Sponge shrimps, mud lobsters. Carapace more or less cylindrical, longer than wide; epistome short, not fused to basal antennular and antennal articles, with narrow point of contact with lateral margin of carapace. Exoskeleton calcified, or chitinised (rarely). Thoracic sternite 8 free from thoracic sternites 1–7. Pleon elongate, muscular; pleonite 2 pleuron overlapping pleonites 1 and 3, or all pleura overlapping next most posterior pleuron. Antennular penduncle articles aligned linearly, main flagellum long, lash-like, aesthetascs more-or-less evenly distributed. Pereopod 1 large, chelate or subchelate; pereopod 2 subchelate or simple. Pereopod 5 shorter than others, subchelate. Uropods biramous.

Key to families of Gebiidea 1. – 2. –

Uropodal rami cylindrical. Midposterior carapace margin acute (Fig. 10.2)��������������������������������������������������Thalassinidae Uropodal rami flattened. Midposterior carapace margin concave������������������������������������������������������������������������������������������2 Rostrum thickened, bluntly rounded or sharply triangular, usually with erect marginal teeth. Maxilla 2 scaphognathite without posterior long setae. Uropodal exopod without transverse suture (Figs 10.3, 10.4)������������������������������������������������������������������������������������������������������������������������������������������������������� Upogebiidae Rostrum thin, rounded or triangular. Maxilla 2 scaphognathite ending posteriorly lobe with numerous long setae (Fig. 10.1i). Uropodal exopod with or without transverse suture (Fig. 10.1)������������������������������Laomediidae

References de Saint Laurent M (1979) Vers une nouvelle classification des Crustacés Décapodes Reptantia. Bulletin de l’Office Nationale de Pêche de Tunisie 3, 15–31. Dworschak PC, Felder DF, Tudge CC (2012) Chapter 69. Infraorders Axiidea de Saint Laurent, 1979 and Gebiidea de Saint Laurent, 1979 (formerly known collectively as Thalassinidea). In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea. Complementary to the volumes translated from the French of the Traité de Zoologie [founded by P.-P. Grassé]. (Eds Schram FR, von Vaupel Klein JC) pp. 109–219. Brill, Leiden. Poore GCB, Ahyong ST, Bracken-Grissom HD, Chan T-Y, Chu KH, et  al. (2014) On stabilising the names of the infraorders of thalassinidean shrimps, Axiidea de Saint Laurent, 1979 and Gebiidea de Saint Laurent, 1979 (Decapoda). Crustaceana 87, 1258–1272. doi:10.1163/​ 15685403-00003354 Robles R, Tudge CC, Dworschak PD, Poore GCB, Felder DL (2009) Molecular phylogeny of the Thalassinidea based on nuclear and mitochondrial genes. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 309–326. CRC Press, Boca Raton. Timm L, Bracken-Grissom HD (2015) The forest for the trees: evaluating molecular phylogenies with an emphasis on higher-level Decapoda. Journal of Crustacean Biology 35, 577–592. doi:10.1163/1937240X-00002371

Laomediidae Borradaile, 1903 mud shrimps Figure 10.1, Plate 18a–d Laomediid mud shrimps inhabit permanent burrows in intertidal and shallow-water estuarine and marine sediments. The family was diagnosed by Ngoc-Ho (1997) who recognised two subfamilies on the basis of a cladistic analysis of adult and larval morphological characters, Axianassinae from Laomediinae, which were subsequently treated as families. Komai (2014) reviewed molecular (Robles et al. 2009; Tsang et al. 2008; Tudge and Cunningham 2002) and morphological (Kensley and Heard 1990; Ngoc-Ho 1997) evidence for the separation of Axianassidae from Laomediidae, as did Anker and Pachelle (2016). Their genera cannot be conveniently divided into two families on the basis of obvious characters such as antennae and sutures on uropodal rami and are treated as one family here. Some species are known only from larvae (Goy and Provenzano 1979; Wear and Yaldwyn 1966).

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Diagnosis. Midposterior margin of carapace evenly concave. Rostrum bluntly rounded, margin rarely denticulate, or bluntly triangular, margin denticulate. Maxilla 2 scaphognathite with

several long setae on end of posterior lobe (Fig. 10.1i). Chelipeds fingers chelate, or simple, palm of propodus with prominent spines opposing dactylus. Pereopods 2 simple.

Key to genera of Laomediidae 1. – 2. – 3. – 4. – 5.

Antennular peduncle article 3 longer than first 2 articles combined. Antennal peduncle article 4 1.5 times or more as long as antennular peduncle (Fig. 10.1b–d, h)����������������������������������������������������������������������������������������������������2 Antennular peduncle article 3 short, as long as articles 1 plus 2 (Fig. 10.1e–g)����������������������������������������������������������������������5 Scaphocerite reduced (Fig. 10.1d). Uropodal exopod with suture, endopod with almost complete suture (Fig. 10.1p)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������Jaxea Scaphocerite triangular, acute, sometimes with median spine (Fig. 10.1b, c, h). Uropodal exopod with or without suture, endopod without or with partial transverse suture (Fig. 10.1n, o)����������������������������������������������������������3 Uropodal exopod with complete transverse suture (Fig. 10.1o)�������������������������������������������������������������������� Heteroaxianassa Uropodal exopod without transverse suture (Fig. 10.1n)�����������������������������������������������������������������������������������������������������������4 Rostrum with lateral and dorsal teeth (Fig. 10.1h). Maxilliped 3 with exopod (Fig. 10.1j)���������������������������������� Strianassa Rostrum without lateral and dorsal teeth (Fig. 10.1b). Maxilliped 3 without exopod������������������������������������������ Axianassa Chelipeds subchelate (Fig. 10.1m). Antennal peduncle almost as long as antennular peduncle, articles of similar lengths; scaphocerite broad, dentate (Fig. 10.1f)������������������������������������������������������������������������������������ Naushonia

Fig. 10.1.  Laomediidae. a, Laomedia healyi Yaldwyn & Wear, 1970. Front of carapace, antennules, antennae: b, Axianassa; c, Heteroaxianassa; d, Jaxea; e, Laomedia; f, Naushonia; g, Saintlaurentiella; h, Strianassa. Maxilla: i, Naushonia. Maxilliped 3: j, Strianassa. Pereopod 1: k, Jaxea; l, Laomedia; m, Naushonia. Telson, uropod: n, Axianassa; o, Heteroaxianassa; p, Jaxea; q, Laomedia.

10 – Gebiidea – mud lobsters and sponge shrimps

– 6. –

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Chelipeds chelate (Fig. 10.1l). Antennal peduncle longer than antennular peduncle, with elongate article 4; scaphocerite lanceolate or reduced (Fig. 10.1e, g)������������������������������������������������������������������������������������������������������������������5 Uropod exopod with suture. Scaphocerite lanceolate (Fig. 10.1g). Maxilliped 3 without exopod���������� Saintlaurentiella Uropodal rami both with suture (Fig. 10.1q). Scaphocerite reduced (Fig. 10.1e). Maxilliped 3 with exopod��������������������������������������������������������������������������������������������������������������������������������������������������������������������������� Laomedia Axianassa Schmitt, 1924

Diagnosis. Rostrum unarmed. Antennular peduncle article 3 longer than first two articles combined; antennal peduncle article 4 almost twice as long as antennule peduncle. Scaphocerite triangular, acute, sometimes with median spine. Maxilliped 3 without exopod. Chelipeds chelate. Uropodal rami both without transverse suture. Maximum cl. 12 mm (Pl. 18a). Intertidal–shelf (0–42 m), subtidal. Tropical W Atlantic, Temperate Northern Pacific, Indo-West Pacific, Tropical Eastern Pacific. 15 species (keys to species: Anker 2010; Anker and Pachelle 2016; Komai 2014; Komai et al. 2020).

Heteroaxianassa Sakai, 2016 Diagnosis. Rostrum with small marginal teeth. Antennular peduncle article 3 longer than first two articles combined; antennal peduncle article 4 almost twice as long as antennule peduncle. Scaphocerite triangular. Maxilliped 3 without exopod. Chelipeds chelate. Uropodal exopod with complete transverse suture, endopod without or with partial suture. Maximum cl. 4.7 mm. Subtidal (0.5–15 m). Temperate Northern W Pacific, Central and Eastern Indo-Pacific. 2 species (Anker 2020: discussion of differences from Axianassa; Komai et al. 2020: key to species).

Jaxea Nardo, 1847 Diagnosis. Rostrum unarmed. Antennular peduncle article 3 longer than first two articles combined; antennal peduncle article 4 almost twice as long as antennule peduncle. Scaphocerite reduced. Maxilliped 3 with exopod. Chelipeds chelate. Uropodal exopod with suture, endopod with almost complete suture. Maximum cl. 20 mm. Intertidal–slope (0–420 m). Temperate Northern Atlantic, Australasia (New Zealand). 2 species (Ngoc-Ho 2003).

Laomedia De Haan, 1841 Diagnosis. Rostrum unarmed. Antennular peduncle article 3 short; antennal peduncle article 4 short. Scaphocerite reduced. Maxilliped 3 with exopod. Chelipeds chelate. Uropodal rami both with transverse suture. Maximum cl. 30 mm (Pl. 18b). Estuarine–marine, intertidal–subtidal, associated with mangroves. Temperate Northern Atlantic, Central Indo-Pacific, Temperate E Australia. 4 species (Ngoc-Ho 1997; Sakai 2016: keys to species).

Naushonia Kingsley, 1897 Diagnosis. Rostrum with small marginal teeth. Antennular peduncle article 3 short; antennal peduncle almost as long as

antennular peduncle, articles of similar lengths. Scaphocerite broad, dentate. Maxilliped 3 with exopod. Chelipeds subchelate. Uropodal rami both with transverse suture. Maximum cl. 20 mm (Pl. 18c, d). Intertidal, subtidal, including marine anchialine caves (0–11 m). Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific, Temperate Australasia. 16 species (Komai and Anker 2010; 2015: rediagnosis, key to 14 species).

Saintlaurentiella Paiva, Tavares & Silva-Neto, 2010 Diagnosis. Rostrum unarmed. Antennular peduncle article 3 short; antennal peduncle article 4 long. Scaphocerite lanceolate. Maxilliped 3 without exopod. Chelipeds chelate. Uropodal exopod only with transverse suture. Maximum cl. 9 mm. Shelf (20–25 m). Tropical W Atlantic. 1 species (Le Loeuff and Intès 1974: as Laurentiella heterocheir Le Loeuff & Intès, 1974).

Strianassa Anker, 2020 Diagnosis. Rostrum with marginal and dorsal teeth. Antennular peduncle article 3 longer than first two articles combined; antennal peduncle article 4 almost twice as long as antennule peduncle. Scaphocerite triangular. Maxilliped 3 with exopod. Chelipeds dissimilar, chelate. Uropodal rami both without transverse suture. Maximum cl. 5.4 mm. Intertidal. Tropical Eastern Pacific (Panama). 1 species (Anker 2020). References Anker A (2010) The mud-shrimp genus Axianassa Schmitt, 1924 (Crustacea, Decapoda, Axianassidae) in the Indo-West Pacific, with description of a new species from French Polynesia. Zootaxa 2557, 49–59. doi:10.11646/zootaxa.2557.1.5 Anker A (2020) Strianassa lerayi gen. et sp. nov., a new laomediid mudshrimp from the eastern Pacific, with new records of Axianassa ngochoae Anker, 2010 and Heteroaxianassa heardi (Anker, 2011) in the western Pacific (Malacostraca: Decapoda: Gebiidea). Zootaxa 4820, 523–539. doi:10.11646/zootaxa.4820.3.6 Anker A, Pachelle PPG (2016) Mud-shrimps of the genus Axianassa Schmitt, 1924 from Panama, with description of two new species (Decapoda: Gebiidea: Laomediidae). Zootaxa 4111, 101–125. doi:10.11646/zootaxa.4111.2.1 Goy JW, Provenzano AJ (1979) Juvenile morphology of the rare burrowing mud shrimp Naushonia crangonoides Kingsley, with a review of the genus Naushonia (Decapoda: Thalassinidea: Laomediidae). Proceedings of the Biological Society of Washington 92, 339–359. Kensley B, Heard RW (1990) The genus Axianassa (Crustacea: Decapoda: Thalassinidea) in the Americas. Proceedings of the Biological Society of Washington 103, 558–572.

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Komai T (2014) A new species of the mud shrimp genus Axianassa (Crustacea: Decapoda: Gebiidea: Axianassidae) from Japan. Species Diversity 19, 141–149. doi:10.12782/sd.19.2.141 Komai T, Anker A (2010) Two new species of the laomediid genus Naushonia Kingsley, 1897 (Crustacea: Decapoda: Gebiidea) from the IndoWest Pacific. Zootaxa 2504, 31–46. doi:10.11646/zootaxa.2504.1.3 Komai T, Anker A (2015) Additional records of the laomediid mudshrimp genus Naushonia Kingsley, 1897 (Crustacea: Decapoda: Gebiidea), with a revised identification key. Zootaxa 3974, 341–360. doi:10.11646/zootaxa.3974.3.3 Komai T, Watanabe T, Matsui S, Tamego T (2020) A new species of the mud shrimp genus Axianassa Schmitt, 1924 (Decapoda: Gebiidea: Laomediidae) from Japan. Zootaxa 4852, 461–474. doi:10.11646/zootaxa.​4852.4.4 Le Loeuff P, Intès A (1974) Les Thalassinidea (Crustacea, Decapoda) du Golfe de Guinée systématique-écologie. Cahiers de l’Office de Recherches Scientifiques et Techniques Outre-Mer, série Océanographique 12, 17–69. Ngoc-Ho N (1997) The genus Laomedia De Haan, 1841 with description of a new species from Vietnam (Crustacea, Thalassinidae, Laomediidae). Zoosystema 19, 729–747. Ngoc-Ho N (2003) European and Mediterranean Thalassinidea (Crustacea, Decapoda). Zoosystema 25, 439–555. Robles R, Tudge CC, Dworschak PD, Poore GCB, Felder DL (2009) Molecular phylogeny of the Thalassinidea based on nuclear and mitochondrial genes. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 309–326. CRC Press, Boca Raton. Sakai K (2016) An aberrant specimen of the genus Laomedia De Haan, 1841, with an additional description of Laomedia astacina De Haan, 1841 and a new genus, Heteroaxianassa (Decapoda, Thalassinidea, Laomediidae). Crustaceana 89, 595–602. doi:10.1163/15685403-00003535 Tsang LM, Lin F-J, Chu KH, Chan T-Y (2008) Phylogeny of Thalassinidea (Crustacea, Decapoda) inferred from three rDNA sequences: implications for morphological evolution and superfamily classification. Journal of Zoological Systematics and Evolutionary Research 46, 216–223. doi:10.1111/j.1439-0469.2008.00459.x Tudge CC, Cunningham CW (2002) Molecular phylogeny of the mud lobsters and mud shrimps (Crustacea: Decapoda: Thalassinidea) using nuclear 18s rDNA and mitochondrial 16s rDNA. Invertebrate Systematics 16, 839–847. doi:10.1071/IS02012 Wear RG, Yaldwyn JC (1966) Studies on thalassinid Crustacea (Decapoda, Macrura Reptantia) with a description of a new Jaxea from New Zealand and an account of its larval development. Zoology Publications from Victoria University of Wellington 41, 1–27.

Thalassinidae Latreille, 1831 mud lobsters Figure 10.2 Mud lobsters of the only genus, Thalassina, are dominant and important members of tropical intertidal and shallow mangrove and seagrass communities. They are deep burrowers, usually estuarine but also found in fully marine environments (Dubey et al. 2012; Ng and Kang 1988; NgocHo and de Saint Laurent 2009). Their burrow openings are typically surrounded by a high volcano of mud. Diagnosis. Midposterior margin of carapace ending in sharp projection. Rostrum sharply triangular, with marginal ridge. Antennular peduncular article 3 about as long as proximal articles. Uropodal rami cylindrical. Maximum cl. to 80 mm.

Fig. 10.2.  Thalassinidae, Thalassina. a, telson, uropod; b, carapace, pleonite 1, dorsal.

Thalassina Latreille, 1806 Estuarine–marine, intertidal to few metres depth. Central IndoPacific. 10 species (Lin et al. 2016: key to species). Until recently only two species were recognised and the name T. anomala (Herbst, 1804) was widely used for the species throughout the Indo-West Pacific. Ngoc-Ho and de Saint Laurent (2009), Moh and Chong (2009) and Sakai and Türkay (2012) described additional species. References Dubey SK, Choudbury A, Chand BK, Trivedi RK (2012) Ecobiological study on burrowing mud lobster Thalassina anomala (Herbst, 1804) (Decapoda: Thalassinidea) in the intertidal mangrove mudflat of deltaic Sundarbans. Exploratory Animal and Medical Research 2, 70–75. Lin F-J, Komai T, Chan T-Y (2016) A new mud lobster of the genus Thalassina Latreille, 1806 (Crustacea: Decapoda: Gebiidea: Thalassinidae) from marine seagrass beds in Dongsha (Pratas) Island, South China Sea. Raffles Bulletin of Zoology 64, 98–105. Moh HH, Chong VC (2009) A new species of Thalassina (Crustacea: Decapoda: Thalassinidea) from Malaysia. Raffles Bulletin of Zoology 57, 465–473. Ng PKL, Kang N (1988) The mud lobster Thalassina. Nature Malaysiana 13, 28–31. Ngoc-Ho N, de Saint Laurent M (2009) The genus Thalassina Latreille, 1806 (Crustacea: Thalassinidea: Thalassinidae). Raffles Bulletin of Zoology Supplement 20, 121–158. Sakai K, Türkay M (2012) A review of the species of the genus Thalassina (Thalassinidea, Thalassinidae). Crustaceana 85, 1339–1376. doi:10.1163/​ 15685403-00003114

10 – Gebiidea – mud lobsters and sponge shrimps

Upogebiidae Borradaile, 1903 mud shrimps, sponge shrimps Figures 10.3, 10.4, Plate 18e–i Upogebiid shrimps are found under rocks, inside sponges or burrowing in sediments. Most are instantly recognised by their compact form and setose flat rostrum. The eastern Pacific species, Upogebia pugettensis (Dana, 1852), forms male–female pairs living in permanent U-shaped burrows up to 1 m deep in soft muds of tidal flats. There, they pump water through the burrow to feed on plankton and suspended detritus (Astall et  al. 1997; Atkinson and Taylor 1988; Nickell and Atkinson 1995; Williams 1986). Upogebia pusilla (Petagna, 1792) has been well studied in Europe (Conides et al. 2012; Dworschak 1983, 1987, 1988). The form of the rostrum, uropods and chelipeds, characters traditionally used to distinguish species and genera, displays considerable diversity. Notably, the one-time division of upogebiids into those with fully chelate chelipeds and those with subchelate is no longer useful for recognising genera. Most species (almost 200 world-wide) are members of the genus Upogebia and until recently, most species were

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referred to this genus. Poore (1994) provided a key to seven genera but more genera were added by Sakai (2006) in a world revision while synonymising others. Sakai’s genera were weakly defined and difficult to distinguish (Poore 2008). The allocation of species to genera was again revised in part by Sakai (2015) but inconsistences remain between type species and others included. The key and diagnoses here include only well diagnosed genera. Upogebia remains a polyphyletic catch-all genus for most species. Regional keys to species can be found for the Mediterranean (Ngoc-Ho 2003), Americas (Williams 1986, 1993), Atlantic and Indo-West Pacific (Sakai 2006), and southern Australia (Poore 2004). Diagnosis. Rostrum bluntly rounded or triangular, setose, usually with marginal teeth. Maxilla scaphognathite with rounded posterior lobe without long setae. Antennular peduncular article 3 about as long as proximal articles. Uropodal rami flat, both without transverse suture (Fig. 10.4) Implicit generic attributes. Rostrum with rounded apex, without inferior spines. Cheliped subchelate, fixed finger at most half as long as dactylus.

Key to genera of Upogebiidae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8.

Rostrum with 1 or more ventral spines (Fig. 10.3d, e)����������������������������������������������������������������������������������������������������������������2 Rostrum without (rarely with) ventral spines������������������������������������������������������������������������������������������������������������������������������3 Rostrum and projecting lateral crests with inferior spines (Fig. 10.3d). Uropodal endopod, outer margin with prominent defined squarish proximal lobe (Fig. 10.4a)������������������������������������������������������������Austinogebia Rostrum only with inferior spines (Fig. 10.3e). Uropodal endopod, outer margin simply curved���������������� Gebiacantha Uropodal rami slender, exopod ~3 times as long as wide (Fig. 10.4b, c)����������������������������������������������������������������������������������4 Uropodal rami broad, exopod at most twice as long as wide (Fig. 10.4e)��������������������������������������������������������������������������������5 Rostrum narrow, nearly parallel-sided, with pair of apical vertical spines. Anterior dorsal region of carapace and rostrum with pair of longitudinal rows of tubercles (Fig. 10.3h). Posterior margin of telson with V-shaped excavation (Fig. 10.4c). Chelipeds subchelate�������������������������������������������������������������������������� Mantisgebia Rostrum broad, with rounded apex. Anterior dorsal region of carapace and rostrum with broad spinose and/or setose area between lateral grooves. Posterior margin of telson convex. Chelipeds simple (Fig. 10.3p)����������� Neogebicula Anterior dorsal region of carapace and rostrum with low smooth median carina between pair of setose areas (Fig. 10.3m)�����������������������������������������������������������������������������������������������������������������������������������������������������Wolffogebia Anterior dorsal region of carapace and rostrum with broad spinose and/or setose area between lateral grooves, sometimes with narrow median groove������������������������������������������������������������������������������������������������������������������6 Anterior dorsal region of carapace and rostrum armed with stout, yellow-translucent tubercles with rounded tips; lateral crests not projecting (Fig. 10.3l)����������������������������������������������������������������������������������� Tuerkayogebia Anterior dorsal region of carapace and rostrum with broad spinose and/or setose area between lateral grooves, sometimes with narrow median groove; lateral crests projecting or not�����������������������������������������������������������7 Posterior margin of telson convex, broader than anterior width, pleonites 5 and 6 and tailfan operculiform (Fig. 10.4d)���������������������������������������������������������������������������������������������������������������������������������������������������������������Pomatogebia Posterior margin of telson more or less concave, straight or convex���������������������������������������������������������������������������������������8 Rostrum tapering, with single acute apical spine. Carapace lateral crests extending about half as long as rostrum, separate from rostrum by a broad U (Fig. 10.3f) (Gebicula uncertain)�������������������������������������������������������������9

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Marine Decapod Crustacea

Fig. 10.3.  Upogebiidae. a, Austinogebia spinifrons (Haswell, 1882); b, Gebicula exigua Alcock, 1901. Anterior carapace, rostrum: c, Aethogebia; d, Austinogebia; e, Gebiacantha. Carapace, dorsal: f, Acutigebia; g, Austinogebia; h, Mantisgebia; i–k, Upogebia spp.; l, Tuerkayogebia; m, Wolffogebia; Cheliped: n, Acutigebia; o, Austinogebia; p, Neogebicula; q, Upogebia.

– 9. – 10. –

Rostrum apex rounded, usually with pair of subapical spines. Carapace lateral crests extending less than half rostrum length (Fig. 10.3i–k)������������������������������������������������������������������������������������������������������������������������������������������10 Chelipeds subchelate, carpus and propodus without spines along upper and lower margins (Fig. 10.3n)�������Acutigebia Chelipeds simple, carpus and propodus with spines along upper and lower margins (Fig. 10.3b)����������������������� Gebicula Rostrum narrow, nearly parallel-sided, with pair of apical vertical spines. Carapace without projecting lateral crests (Fig. 10.3c)������������������������������������������������������������������������������������������������������������������������������������������� Aethogebia Rostrum broad, tapering, with rounded ape. Carapace lateral crests variously projecting (Fig. 10.3i–k)���������� Upogebia

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Fig. 10.4.  Upogebiidae. Telson, uropod: a, Austinogebia; b, Neogebicula; c, Mantisgebia; d, Pomatogebia; e, Upogebia.

Acutigebia Sakai, 1982

Gebicula Alcock, 1901

Diagnosis. Rostrum tapering, with single acute apical spine. Carapace lateral crests projecting about half as far as rostrum, separated from rostrum by a broad U. Maximum cl. 11 mm (Pl. 18e). Intertidal, subtidal. Temperate W Northern Pacific, Central Indo-Pacific, Temperate Australasia. 7 species. (Liu and Liu 2013b: key to species).

Diagnosis. Rostrum tapering, with single acute apical spine. Carapace lateral crests projecting about half as far as rostrum, separated from rostrum by narrow slot. Cheliped simple, with subdistal spine on lower margin of propodus. Uropodal rami slender, exopod ~3 times as long as wide. Maximum cl. 6 mm. Slope. Central Indo-Pacific. 1 species. Gebicula exigua Alcock, 1901 is the only species confidently included in this genus (Alcock 1901; Alcock and McArdle 1901).

Aethogebia Williams, 1993 Diagnosis. Rostrum narrow, nearly parallel-sided, with pair of apical vertical spines. Carapace lateral crests projecting not at all. Maximum cl. 4 mm. Subtidal. Tropical W Atlantic. 1 species (known from holotype only) (Williams 1993).

Austinogebia Ngoc-Ho, 2001 Diagnosis. Rostrum tapering, with single acute apical spine or ovoid, slightly wider at midpoint than at base, with pair of subapical spines, rostrum and projecting lateral crests with ventral spines; lateral crests projecting about half as far as rostrum, separated from rostrum by narrow slot. Uropodal endopod anterior margin with prominent defined squarish proximal lobe. Maximum cl. 15 mm. Subtidal–shelf. Temperate W Northern Pacific, Western IndoPacific, Temperate Australasia. 7 species (keys to species: Liu and Liu 2012; Ngoc-Ho 2001).

Gebiacantha Ngoc-Ho, 1989 Diagnosis. Rostrum ovoid, wider at midpoint than at base, with pair of subapical spines, with 1 or more ventral spines. Carapace lateral crests projecting as minute teeth, scarcely separate from rostrum. Posterior margin of telson concave to convex. Maximum cl. 12 mm (Pl. 18f). Intertidal–subtidal. Temperate W Northern Pacific, Tropical Atlantic, Western and Central Indo-Pacific. 20 species (Komai 2020; Ngoc-Ho 1989: key to 11 species).

Mantisgebia Sakai, 2006 Diagnosis. Rostrum narrow, nearly parallel-sided, with pair of apical vertical spines. Carapace lateral crests projecting much less than half as far as rostrum, widely separated; anterior dorsal region of carapace and rostrum with pair of longitudinal rows of tubercles. Uropodal rami slender, exopod ~3 times as long as wide. Posterior margin of telson with V-shaped excavation. Maximum cl. 7 mm. Shelf. Temperate Northern W Pacific, Central Indo-Pacific. 4 species (Liu and Liu 2013a).

Neogebicula Sakai, 1982 Diagnosis. Carapace lateral crests projecting much less than half as far as rostrum, widely separated. Cheliped simple, with small distal spine l on lower margin of propodus. Uropodal rami slender, exopod ~3 times as long as wide. Maximum cl. 3 mm. Subtidal. Western and Central Indo-Pacific. 19 species. Dworschak and Anker (2022) treated Paragebicula Sakai, 2006 as a synonym. Stated differences between Neogebicula and Paragebicula (Liu and Liu 2010; Sakai 2006) are not discernible over the range of morphologies of the species included.

Pomatogebia Williams & Ngoc-Ho, 1990 Diagnosis. Carapace lateral crests projecting much less than half as far as rostrum, widely separated. Posterior margin of telson

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convex, broader than anterior width, pleonites 5 and 6 plus tailfan operculiform. Maximum cl. 13 mm. Subtidal. Tropical W Atlantic. 3 species (Williams and NgocHo 1990). Williams (1986) noted that this rugose operculum seals the U-shaped tube in the coral reef where the shrimp is imprisoned.

Tuerkayogebia Sakai, 1982 Diagnosis. Carapace lateral crests projecting not at all; anterior dorsal region of carapace and rostrum armed with stout, yellowtranslucent tubercles with rounded tips. Cheliped chelate, fingers equal in length or almost so. Maximum cl. 10 mm. Subtidal. Temperate W Northern Pacific. 1 species (Sakai 1982).

Upogebia Leach, 1814 Diagnosis. Rostrum and projecting lateral crests without or with 1 or more ventral spines (rarely). Carapace lateral crests projecting about half as far as rostrum, separated from rostrum by a broad U, or about half as far as rostrum, separated from rostrum by narrow slot, or much less than half as far as rostrum, widely separated, or as minute teeth, scarcely separate from rostrum. Cheliped subchelate, fixed finger at most half as long as dactylus, or chelate, fingers equal in length or almost so. Maximum cl. 21 mm (Pl. 18g–i). Estuarine–marine, intertidal–slope. Cosmopolitan except Arctic and Southern Oceans. 125 species. Upogebia is by far the largest genus from which others have been hived off. Sakai’s (2006) keys are valuable but his taxonomy is not followed here. Arabigebicula Sakai, 2006, based on a single damaged specimen without chelipeds, is treated as a junior synonym.

Wolffogebia Sakai, 1982 Diagnosis. Carapace lateral crests projecting much less than half as far as rostrum, widely separated; anterior dorsal region of carapace and rostrum with low smooth median carina between pair of setose grooves. Maximum cl. 14 mm. Estuarine–marine, intertidal–subtidal; muddy mangrove environments. Temperate Northern W Pacific, Central IndoPacific. 6 species (Sakai 1982). The number of species attributed to this genus is disputed (Ngoc-Ho et al. 2001; Sakai and Lheknim 2014). References Alcock A (1901) A descriptive catalogue of the Indian deep-sea Crustacea Decapoda Macrura and Anomala, in the Indian Museum. Being a revised account of the deep-sea species collected by the Royal Indian Marine Survey Ship Investigator. Trustees of the Indian Museum, Calcutta. Alcock A, McArdle SB (1901) Illustrations of the zoology of the Royal Indian Marine Surveying Steamer Investigator, under the command of Commander T.H. Heming, R.N. Crustacea. – Part IX, Plates XLIX–LV Office of the Superintendent of Government Printing, India, Calcutta. Astall CM, Taylor AC, Atkinson RJA (1997) Behavioural and physiological implications of a burrow-dwelling lifestyle for two species of a

upogebiid mud-shrimp (Crustacea: Thalassinidea). Estuarine, Coastal and Shelf Science 44, 155–168. doi:10.1006/ecss.1996.0207 Atkinson RJA, Taylor AC (1988) Physiological ecology of burrowing decapods. Symposia of the Zoological Society of London 59, 201–226. Conides AJ, Nicolaidou A, Apostolopoulou M, Thessalou-Legaki M (2012) Growth, mortality and yield of the mudprawn Upogebia pusilla (Petagna, 1792) (Crustacea: Decapoda: Gebiidea) from western Greece. Acta Adriatica 53, 87–103. Dworschak PC (1983) The biology of Upogebia pusilla (Petagna) (Decapoda, Thalassinidea). I. The burrows. Pubblicazioni della Stazione Zoologica di Napoli I (Marina Ecologia) 4, 19–43. doi:10.1111/j.1439-0485.1983.​ tb00286.x Dworschak PC (1987) The biology of Upogebia pusilla (Petagna) (Decapoda, Thalassinidea) II. Environments and zonation. Pubblicazioni della Stazione Zoologica di Napoli I (Marina Ecologia) 8, 337–358. doi:10.1111/j.​1439-0485.1987.tb00193.x Dworschak PC (1988) The biology of Upogebia pusilla (Petagna) (Decapoda, Thalassinidea). III. Growth and production. Pubblicazioni della Stazione Zoologica di Napoli I (Marina Ecologia) 9, 51–77. doi:10.1111/j.​ 1439-0485.​1988.tb00198.x Dworschak PC, Anker A (2022) Axiidea (Crustacea: Callianassidae, Callichiridae and Ctenochelidae) and Gebiidea (Upogebiidae) collected during the Comprehensive Marine Biodiversity Survey of Singapore. Raffles Bulletin of Zoology 70, 108–133. doi:10.26107/RBZ-2022-0008 Komai T (2020) Redescription of a poorly known upogebiid mud shrimp Gebiacantha acanthochela (Sakai, 1967) and description of a new species of the genus from the Ryukyu Islands, Japan (Decapoda: Gebiidea). Zootaxa 4881, 307–322. Liu WL, Liu RY (2010) The subfamily Neogebiculinae (Decapoda, Thalassinidea, Upogebiidae) from South China Sea. In: Fransen CHJM, De Grave S, Ng PKL (Eds) Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume. Crustaceana Monographs 14, 389–401. Liu W, Liu J-Y (2012) A new species of the genus Austinogebia Ngoc-Ho, 2001 (Crustacea, Decapoda, Gebiidea, Upogebiidae) from northern China. Zootaxa 3243, 59–64. doi:10.11646/zootaxa.3243.1.4 Liu W, Liu J-Y (2013a) A new species of the genus Mantisgebia Sakai, 2006 (Crustacea, Decapoda, Gebiidea, Upogebiidae) from the South China Sea. Zootaxa 3637, 592–596. doi:10.11646/zootaxa.3637.5.6 Liu W, Liu J-Y (2013b) Two new species of Acutigebia (Crustacea: Decapoda: Gebiidea: Upogebiidae) from the South China Sea. Raffles Bulletin of Zoology 61, 571–577. Ngoc-Ho N (1989) Sur le genre Gebiacantha gen. nov., avec la description de cinq espèces nouvelles (Crustacea, Thalassinidea, Upogebiidae). Bulletin du Muséum National d’Histoire Naturelle 11, 117–145. Ngoc-Ho N (2001) Austinogebia, a new genus in the Upogebiidae and rediagnosis of its close relative, Gebiacantha Ngoc-Ho, 1989 (Crustacea: Decapoda: Thalassinidea). Hydrobiologia 449, 47–58. doi:10.1023/A:​ 1017516331683 Ngoc-Ho N (2003) European and Mediterranean Thalassinidea (Crustacea, Decapoda). Zoosystema 25, 439–555. Ngoc-Ho N, Ngoc-Dung D, Phi-Hung T (2001) The genus Wolffogebia Sakai, 1982 (Crustacea, Decapoda, Thalassinidea, Upogebiidae) with a new species from Vietnam. Zoosystema 23, 101–108. Nickell LA, Atkinson RJA (1995) Functional morphology of burrows and trophic modes of three thalassinidean shrimp species, and a new approach to the classification of thalassinidean burrow morphology. Marine Ecology Progress Series 128, 181–197. doi:10.3354/ meps128181 Poore GCB (1994) A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys to the families and genera. Memoirs of the Museum of Victoria 54, 79–120. doi:10.24199/j.mmv.1994.54.03

10 – Gebiidea – mud lobsters and sponge shrimps

Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Poore GCB (2008) Book review “Sakai, K. 2006. Upogebiidae of the world (Decapoda, Thalassinidea). Crustaceana Monographs 6, i–ix, 185 pp., 23 textfigs. Koninklijke Brill, NV, Leiden, The Netherlands, ISBN-13: 978 90 04 15150 8, hardcover, Euro98/U.S.$132. Journal of Crustacean Biology 28, 422–423. Sakai K (1982) Revision of Upogebiidae (Decapoda, Thalassinidea) in the Indo-West Pacific region. Researches on Crustacea Special Number 1, 1–106. Sakai K (2006) Upogebiidae of the world (Decapoda, Thalassinidea). Crustaceana Monographs 6, 1–185. Sakai K (2015) Upogebiidae Borradaile, 1903 from the Andaman Sea, collected by the Phuket Marine Biological Center, Thailand, with seven

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new species, presented in revised keys (Thalassinidea Latreille, 1831). Crustaceana 88, 577–609. doi:10.1163/15685403-00003425 Sakai K, Lheknim V (2014) Two new species of the genera Neocallichirus and Wolffogebia (Decapoda, Pleocyemata) from Thale Sap Songkhla, Songkhla Lagoon System, Songkhla Province, Thailand. Crustaceana 87, 91–100. doi:10.1163/15685403-00003269 Williams AB (1986) Mud shrimps, Upogebia, from the Eastern Pacific (Thalassinidea: Upogebiidae). San Diego Society of Natural History, Memoir 14, 1–60. Williams AB (1993) Mud shrimps, Upogebiidae, from the western Atlantic (Crustacea: Decapoda: Thalassinidea). Smithsonian Contributions to Zoology 544, 1–77. doi:10.5479/si.00810282.544 Williams AB, Ngoc-Ho N (1990) Pomatogebia, a new genus of thalassinidean shrimps for Western Hemisphere tropics (Crustacea: Upogebiidae). Proceedings of the Biological Society of Washington 103, 614–616.

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11. Achelata – spiny lobsters, slipper lobsters and bugs

Achelata are entirely marine and include a variety of lobsterlike crustaceans. Diagnostic features of the adults are the possession of a thickened, strong cylindrical or flattened carapace covering all of the head and thorax, absence of a prominent rostrum, and fusion of the basal antennular and antennal articles to the epistome. Other features uniting the Achelata are enlarged antennae, pereopods without chelae, and a unique knob attaching the carapace to the last thoracic somite. All the pereopods are simple or at most subchelate (which distinguishes them from ‘true’ lobsters, which have an enlarged cheliped). The monophyly of the Achelata is now well established (Ahyong and O’Meally 2004; Bracken-­Grissom et  al. 2014; Porter et al. 2005; Scholtz and Richter 1995). The Achelata Scholtz & Richter, 1995 are unique in passing their early larval stages as a pelagic phyllosoma larva (Fig. 11.1) (Baisre 1994). The phyllosoma is longlived, almost two years in some palinurids, and is known to travel widely in oceanic currents from the shore environments where the adults live (Chiswell and Booth 2017). The adult pleon (abdomen) is strong and ends as a broad tail fan formed from the telson and uropodal rami. This enables a quick reverse escape response by swimming. The meaty tail is also the reason that spiny lobsters in particular, slipper lobsters less so, are the basis of valuable fisheries world-wide. They are usually trapped individually using baited pots. Baisre (1994) was the first to publish a phylogeny of genera of the Achelata (which he called Palinuroidea) based on larval characters. Later workers largely confirmed his findings based on adult features (e.g. Dixon et al. 2003), extant and extinct forms (Karasawa et  al. 2013) and molecules (Davis et al. 2015; Palero et al. 2009). Diagnosis. Carapace more or less cylindrical or prismatic, longer than wide, or flat, usually with lateral edge, usually wider or as long as wide; epistome short, fused to anterior and lateral margin of carapace, basal antennular and antennal articles; basal antennular articles fused medially;

Fig. 11.1.  Phyllosoma of Palinurus elephas (Fabricius, 1787).

exoskeleton calcified. Thoracic sternites 1–8 fused. Pleon elongate, muscular; pleura not overlapping or overlapping next most posterior pleuron. Antennular peduncle articles aligned linearly, main flagellum short, compact or long, lash-like (Panulirus), aesthetascs concentrated on one side forming dense brush. Pereopods 1–5 simple, or pereopods 1 and 5 subchelate. Pereopod 5 of similar length to more anterior pereopods. Uropods biramous.

Key to families of Achelata 1. –

Carapace subcylindrical or prismatic. Antennal flagellum long, filamentous, cylindrical (Fig. 11.2)������������Palinuridae Carapace flattened. Antennal peduncle and flagellum short, flattened (Figs 11.3, 11.4)������������������������������������ Scyllaridae

References Ahyong ST, O’Meally D (2004) Phylogeny of the Decapoda Reptantia: resolution using three molecular loci and morphology. Raffles Bulletin of Zoology 52, 673–693.

Baisre JA (1994) Phyllosoma larvae and the phylogeny of Palinuroidea (Crustacea: Decapoda): a review. Australian Journal of Marine and Freshwater Research 45, 925–944. doi:10.1071/MF9940925 Bracken-Grissom HD, Ahyong ST, Wilkinson RD, Feldmann RM, Schweitzer CE, et  al. (2014) The emergence of the lobsters: phylogenetic

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relationships, morphological evolution and divergence time comparisons of an ancient group (Decapoda: Achelata, Astacidea, Glypheidea, Polychelida). Systematic Biology 63, 457–479. doi:10.1093/sysbio/syu008 Chiswell SM, Booth JD (2017) Evolution of long larval life in the ­Australasian rock lobster Jasus edwardsii. Marine Ecology Progress Series 576, 69–87. doi:10.3354/meps12233 Davis KE, Hesketh TW, Delmer C, Wills MA (2015) Towards a supertree of Arthropoda: a species-level supertree of the spiny, slipper and coral lobsters (Decapoda: Achelata). PLoS One 10, e0140110. doi:10.1371/ journal.pone.0140110 Dixon CJ, Ahyong ST, Schram FR (2003) A new hypothesis of decapod phylogeny. Crustaceana 76, 935–975. doi:10.1163/​156854003771997846 Haeckel E (1904) Kunst Formen der Natur. Bibiographisches Instituts, Leipzig und Wien. Karasawa H, Schweitzer CE, Feldmann RM (2013) Phylogeny and systematics of extant and extinct lobsters. Journal of Crustacean Biology 33, 78–123. doi:10.1163/1937240X-00002111 Palero F, Crandall KA, Abelló P, Macpherson E, Pascual A (2009) Phylogenetic relationships between spiny, slipper and coral lobsters (Crustacea, Decapoda, Achelata). Molecular Phylogenetics and Evolution 50, 152–162. doi:10.1016/j.ympev.2008.10.003 Porter ML, Pérez-Losada M, Crandall KA (2005) Model-based multi-locus estimation of decapod phylogeny and divergence times. Molecular Phylogenetics and Evolution 37, 355–369. doi:10.1016/j.ympev.2005.06.021 Scholtz G, Richter S (1995) Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca). Zoological Journal of the Linnean Society 113, 289–328. doi:10.1006/zjls.1995.0011

Palinuridae Latreille, 1802 spiny lobsters, rock lobsters Figure 11.2, Plate 19 Palinuridae are generally called spiny lobsters or rock lobsters, or in Australia and New Zealand crayfish or ‘crays’. They should not be confused with the widespread

freshwater crayfish or clawed lobsters of North America and Europe (infraorder Astacidea), which have large claws that palinurids lack. The pleon of most palinurids is large, muscular and edible, and lucrative fisheries exist for the commonest and largest species. Fishing is usually by the use of baited pots set from small boats close to shore. Because they are so commercially important, much is known about the biology and fisheries potential of spiny lobsters. They inhabit shallow nearshore rocky or reef environments, rarely to 1000 m depth. They are cryptic, hiding in rock crevices during the day and coming out at night to feed. Analyses of stomach contents tell us that they usually feed on small invertebrates and occasionally on algae. During development, palinurids pass through a translucent flat phyllosoma larva stage that lives in the plankton for up to 22 months feeding on gelatinous zooplankton (Johnston and Ritar 2001). In the Western Australian species, Panulirus cygnus George, 1962, the phyllosoma passes through nine instars lasting 9–11 months and is transported by currents well into the Indian Ocean before moulting to the puerulus stage, resembling a translucent miniature adult (Phillips et al. 1978). The puerulus swims across the continental shelf to settle in shallow inshore reef areas. George and Main (1967) recognised two lines of evolution in the Palinuridae based on the presence (in the Stridentes) or absence (in the Silentes) of a sound-producing organ, a stridulating apparatus. The stridulating apparatus consists of two parts, a movable plectrum attached to the

Fig. 11.2.  Palinuridae. Anterior carapace margins: a, Jasus; b, Justitia; c, Linuparus; d, Nupalirus; e, Palibythus; f, Palinurellus; g, Palinurus; h, Palinustus; i, j, Panulirus; k, Projasus; l, Puerulus; m, Sagmariasus. Pleons: n, Palibythus; o, Palinurellus. Pereopod 1 of male: p, Justitia.

11 – Achelata – spiny lobsters, slipper lobsters and bugs

last segment of the antennal peduncle, and a rigid file, located between the eyes on the side of the antennular plate (Patek 2002: fig. 1). Sound is produced by moving these two parts against each other. George (1997) revisited the evolution of the group, especially of Jasus and Panulirus, in the light of a more modem understanding of plate tectonics. In a study of the radiation of the family, Stridentes have been found to be monophyletic but Silentes paraphyletic (Tsang et al. 2009). The three genera restricted to the southern high latitudes (Jasus, Projasus and Sagmariasus) constitute the basal lineages in the spiny lobsters, suggesting a Southern Hemisphere origin for the group.

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There are 12 genera and 62 species of Palinuridae worldwide (Chan 2010). Another family, Synaxidae, was synonymised with Palinuridae by Davie (1990), a decision confirmed by Palero et al. (2009a) and Tsang et al. (2009). Holthuis (1991) provided illustrated keys to many species of commercial interest. Diagnosis. Carapace subcylindrical or prismatic. Antennal peduncle cylindrical, with long, filamentous flagellum. Implicit generic attributes. Supraocular horns prominent, upright. Carapace without strong longitudinal ridges. Pleonites 3–5 pleura triangular. Antennal flagella flexible.

Key to genera of Palinuridae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. – 9. – 10. – 11. –

Carapace prismatic, with strong coarse longitudinal ridges (Pl. 19d, f). Female pleopods 2 with stylamblys (second endopodal article) as long as in remaining pleopods���������������������������������������������������������������������������������������������2 Carapace subcylindrical (Pl. 19a–c, e), without strong longitudinal ridges but often with rows of spines. Female pleopods 2 with stylamblys absent or minute�����������������������������������������������������������������������������������������������������������3 Rostrum a broad 2- or 4-spined median projection of carapace between eye (Fig. 11.2c). Antennal flagella straight, inflexible�������������������������������������������������������������������������������������������������������������������������������������������������������Linuparus Supraorbital horns widely separate, small, flattened (Fig. 11.2l), rostrum absent. Antennal flagella flexible����Puerulus Rostrum broad, flat, triangular. Carapace supraocular spines small, low (Fig. 11.2e, f)������������������������������������������������������4 Rostrum absent or narrowly triangular, spine-like, sometimes distally bifid. Carapace supraocular horns prominent, upright (Fig. 11.2a, d, g–k, m)�������������������������������������������������������������������������������������������������������������������������������5 Antenna with stridulating apparatus formed by antenna and protruding antennular plate. Pleonal 3–5 pleura with strong anterior tooth followed by denticulate lobe (Fig. 11.2n)����������������������������������������������������� Palibythus Antenna without stridulating apparatus. Pleonal 3–5 pleura triangular (Fig. 11.2o)����������������������������������������Palinurellus Antennule with 2 very long flagella, much longer than peduncle���������������������������������������������������������������������������� Panulirus Antennule with 2 short flagella, shorter than half length of peduncle������������������������������������������������������������������������������������6 Rostrum triangular, as long as or longer than prominent curved simple supraocular spines (Fig. 11.2a, m)�������������������7 Rostrum, if present, shorter than supraocular horns�����������������������������������������������������������������������������������������������������������������8 Pleonites with dorsal scalloped sculpture (Pl. 19a)��������������������������������������������������������������������������������������������������������������Jasus Pleonites dorsally smooth (Pl. 19b)��������������������������������������������������������������������������������������������������������������������������� Sagmariasus With stridulating apparatus formed by antenna and protruding antennular plate��������������������������������������������������������������9 Without stridulating apparatus����������������������������������������������������������������������������������������������������������������������������������������������������10 Supraorbital horns tapering to sharp point (Fig. 11.2g). Antennular peduncle article 1 shorter than antennal peduncle����������������������������������������������������������������������������������������������������������������������������������������������������������������� Palinurus Supraorbital horns truncate, with anterior margin crenulate (Fig. 11.2h). Antennular peduncle reaching beyond antennal peduncle���������������������������������������������������������������������������������������������������������������������������������������� Palinustus Rostrum absent. Carapace supraocular horns as anterolaterally directed, flat, smooth, triangular plates (Fig. 11.2k). Dorsum of carapace with 2 submedian longitudinal rows of spines�����������������������������������������������Projasus Rostrum triangular or spine-like. Supraorbital horns prominent, dorsally dentate, directed fowards (Fig. 11.2d). Dorsum of carapace with few spines, squamosa��������������������������������������������������������������������������������������������11 Pereopods 1 enlarged in male, subchelate (Fig. 11.2p, Pl. 19c), with wide red crossbands, simple in female. Supraorbital horns with 2 teeth on upper margin. Rostrum flanked by several small spines����������������������������� Justitia Pereopods 1 not sexually dimorphic, simple, without coloured crossbands. Supraorbital horns with 3 teeth on upper margin (Fig. 11.2d). Rostrum without spines on either side flanked������������������������������������������������� Nupalirus

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Marine Decapod Crustacea

Justitia Holthuis, 1946 Japanese furrow lobster Diagnosis. Carapace dorsum ornamented with a strong, scale-like sculpture; with triangular or spine-like rostrum, flanked by several small spines; prominent anteriorly directed supraorbital horns with 2 teeth on upper margin. Pleonites, upper surface evenly orange-red, without transverse bands; pleonites 3–5 pleura with 2 simple, strong, sharp teeth. With stridulating apparatus. Pereopods 1 enlarged in male, subchelate, with wide red crossbands, simple in female. Maximum cl. 70 mm (Pl. 19c). Subtidal–slope (1–300 m). Temperate Northern Pacific, Tropical Atlantic, Indo-West Pacific. 1 species (Poupin 1994). Justitia longimana (H. Milne Edwards, 1837) is particularly colourful, and although rare, is collected for the aquarium trade (Calado et al. 2003).

Jasus Parker, 1883 spiny lobsters, rock lobsters, crayfish Diagnosis. Carapace dorsum irregularly spinose; with triangular rostrum, prominent supraorbital horns. Pleonites, upper surface with scalloped sculpture; pleonites 3–5 pleura with strong anterior tooth followed by denticulate lobe. Without stridulating apparatus. Maximum cl. 235 mm (Pls 1a, 19a). Intertidal–shelf (0–200 m). Temperate South America, Temperate Southern Africa, Temperate Australasia, Eastern IndoPacific (Pitcairn Island). 5 species (Holthuis 1991: key to species). All species are the basis of commercial fisheries. Much is known of the biology and fisheries of the species in Australia, New Zealand and South Africa (e.g. Jeffs et al. 2013).

Linuparus White, 1847 spear lobsters, box lobsters, barking lobsters, champagne lobsters Diagnosis. Carapace prismatic, with strong longitudinal ridges; rostrum a broad 2- or 4-spined median projection of carapace between eyes. Antennal flagella straight, inflexible; with stridulating apparatus. Maximum cl. 18 mm (Pl. 19d). Shelf–slope (30–414 m). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Australasia. 4 species (Holthuis 1991; Tsoi et al. 2011: key to species, biogeography).

Nupalirus Kubo, 1955 Diagnosis. Carapace dorsum ornamented with a strong, scalelike sculpture; small triangular rostrum without flanking spines; prominent anteriorly directed supraorbital horns, with 3 teeth on upper margin. Pleonites, upper surface with 6 transverse red bands; pleonites 3–5 pleura with 2 simple, strong, sharp teeth. Without stridulating apparatus. Maximum cl. 83 mm. Shelf (40–200 m). Temperate Northern Pacific, Indo-West Pacific. 3 species (Poupin 1994: described species of Nupalirus as the ‘J. japonica group’ of Justitia, key to species).

Palibythus Davie, 1990 Diagnosis. Carapace dorsum covered with small blunt tubercles and felt of short setae, with 2 short, submedian, post-rostral ridges and deep cervical grooves; with broad, flat, triangular rostrum, supraocular spines obsolete, low. Pleonites, upper surface tuberculate, with median carinae; pleonites 3–5 pleura with strong anterior tooth followed by denticulate lobe. With stridulating apparatus. Maximum cl. 69 mm. Shelf, slope (70–300 m). Central and Eastern Indo-Pacific. 1 species (Davie 1990).

Palinurellus von Martens, 1878 furry lobster Diagnosis. Carapace dorsum setose; with broad, flat, triangular rostrum, supraocular spines obsolete, low. Pleonites, upper surface tuberculate, setose. Without stridulating apparatus. Maximum cl. 71 mm. Subtidal, shelf (1–35 m). Western and Central Indo-Pacific. 2 species (Holthuis 1991).

Palinurus Weber, 1795 spiny lobsters, rock lobsters Diagnosis. Carapace dorsum with rows of few submedian and sublateral spines, generally spinulose; front concave, denticulate between flat triangular supraocular horns. With stridulating apparatus. Maximum cl. 220 mm. Intertidal–slope (0–600 m). Tropical Atlantic, Temperate Northern Pacific, Western Indo-Pacific, Temperate Southern Africa. 6 species (Groeneveld et al. 2006: key to species; Holthuis 1991: 5 species diagnoses). Three species are present in the Mediterranean and three in the North-eastern Atlantic – their phylogeny is well documented (Palero et  al. 2009b) as are fisheries for the species (e.g. Palero et al. 2010), especially Palinurus elephas (Fabricius, 1787).

Palinustus A. Milne-Edwards, 1880 blunthorn lobsters Diagnosis. Carapace dorsum with rows of few submedian and sublateral spines, generally spinulose; front truncate between flat, truncate, supraocular horns. With stridulating apparatus. Maximum cl. 48 mm. Shelf–bathyal (59–4122 m). Tropical Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific. 5 species (Chan and Yu 1995: key to species).

Panulirus White, 1847 spiny lobsters, rock lobsters Diagnosis. Carapace dorsum irregularly spinose; with straight front, prominent curved supraorbital horns. Pleonites 3–5 pleura

11 – Achelata – spiny lobsters, slipper lobsters and bugs

with strong anterior tooth followed by denticulate lobe. Without stridulating apparatus. Maximum cl. 200 mm (Pl. 19e). Intertidal–shelf (0–120 m). Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Tropical Eastern Pacific. 22 species (Giraldes and Smyth 2016: biogeography of 6 species in the Atlantic, keys to species, based on morphology and on live coloration; Holthuis 1991: key to 18 species; Sekiguchi and George 2005). Many species are commercially fished (Briones‐Fourzán and Lozano‐Álvarez 2013). Ptacek et  al. (2001), on the basis of mitochondrial genes, found two major lineages of species within Panulirus. One of these lineages equated to previously morphologically defined groups I and II and the other to groups III and IV (George and Main 1967; McWilliam 1995) but further separation of the four groups was not supported. George (2005) confirmed that the two lineages have different male copulatory structures.

Projasus George & Grindley, 1964 jagged lobsters Diagnosis. Carapace dorsum with 2 submedian and 2 lateral longitudinal rows of spines; with widely spaced flat, smooth, triangular supraocular plates, directed anterolaterally. Pleonites 3–5 pleura with 2 simple, strong, sharp teeth. Without stridulating apparatus. Maximum cl. 90 mm. Shelf–slope (175–880 m). Temperate South America (Chile), Temperate Southern Africa, Temperate Australasia. 2 species (George and Grindley 1964; Holthuis 1991).

Puerulus Ortmann, 1897 whip lobsters Diagnosis. Carapace prismatic, with strong longitudinal ridges, dorsum with median ridge behind cervical groove, often with spines or tubercles, without submedian rows; with supraorbital spines widely separate, small, flattened. Pleonites 3–5 pleura with 2 simple, strong, sharp teeth. With stridulating apparatus. Maximum cl. 74 mm (Pl. 19f). Shelf–slope (73–1608 m). Western and Central Indo-Pacific. 9 species (Chan et al. 2013: key to species). Puerulus is both a genus name as here and the term given to the first-settled stage of palinurids in general.

Sagmariasus Holthuis, 1991 eastern rock lobster (Australia), packhorse lobster (New Zealand) Diagnosis. Carapace dorsum with 2 large median teeth before cervical groove, posterior half closely set with numerous sharp spinules; with triangular rostrum, prominent supraorbital horns. Pleonites, upper surface smooth; pleonites 3–5 pleura with strong anterior tooth followed by denticulate lobe. Without stridulating apparatus. Maximum cl. 250 mm (Pl. 19b).

251

Intertidal–shelf (0–200 m). Temperate Australasia (Australia, New Zealand). 1 species. Around Sydney, Australia, S. verreauxi (H. Milne Edwards, 1851) forms the basis of a fluctuating commercial fishery (Montgomery and Liggins 2013). What is regarded as the same species in northern New Zealand is significantly genetically different from the Australian population (Brasher et al. 1992). Natural history and biology are well documented (Booth 2011). References Booth J (2011) Spiny lobsters through the eyes of the giant packhorse. Victoria University Press, Wellington. Brasher DJ, Ovenden JR, Booth JD, White RWG (1992) Genetic subdivison of Australian and New Zealand populations of Jasus verreauxi (Decapoda: Palinuridae) – preliminary evidence from the mitochondrial genome. New Zealand Journal of Marine and Freshwater Research 26, 53–58. doi:10.1080/00288330.1992.9516499 Briones‐Fourzán P, Lozano‐Álvarez E (2013) Essential habitats for Panulirus spiny lobsters. In Lobsters: biology, management, aquaculture and fisheries. (Ed. Phillips BF) pp. 186–220. Wiley-Blackwell, Chichester. Calado R, Lin J, Rhyne AL, Araújo R, Narciso L (2003) Marine ornamental decapods — popular, pricey, and poorly studied. Journal of Crustacean Biology 23, 963–973. doi:10.1651/C-2409 Chan T-Y (2010) Annotated checklist of the world’s marine lobsters (Crustacea: Decapoda: Astacidea, Glypheidea, Achelata, Polychelida). Raffles Bulletin of Zoology Supplement 23, 153–181. Chan T-Y, Yu H-P (1995) The rare lobster genus Palinustus A. Milne Edwards, 1880 (Decapoda: Palinuridae), with description of a new species. Journal of Crustacean Biology 15, 376–394. doi:10.2307/​1548964 Chan T-Y, Ma KY, Chu KH (2013) The deep-sea spiny lobster genus Puerulus Ortmann, 1897 (Crustacea, Decapoda, Palinuridae), with descriptions of five new species. In: Ahyong ST, Chan T-Y, Corbari L, Ng PKL (Eds), Tropical Deep-Sea Benthos Vol. 27. Mémoires du Muséum National d’Histoire Naturelle, Paris 204, 191–230. Davie PJF (1990) A new genus and species of marine crayfish, Palibythus magnificus, and new records of Palinurellus (Decapoda: Palinuridae) from the Pacific Ocean. Invertebrate Taxonomy 4, 685–695. doi:10.1071/IT9900685 George RW (1997) Tectonic plate movements and the evolution of Jasus and Panulirus spiny lobsters (Palinuridae). Marine and Freshwater Research 48, 1121–1130. doi:10.1071/MF97202 George RW (2005) Comparative morphology and evolution of the reproductive structures in spiny lobsters, Panulirus. New Zealand Journal of Marine and Freshwater Research 39, 493–501. doi:10.1080/00288330.200 5.9517328 George RW, Grindley JR (1964) Projasus — a new generic name for Parker’s crayfish, Jasus parkeri Stebbing (Palinuiridae: “Silentes”). Journal of the Royal Society of Western Australia 47, 87–90. George RW, Main AR (1967) The evolution of the spiny lobsters (Palinuridae): a study of evolution in the marine environment. Evolution 21, 803–820. doi:10.1111/j.1558-5646.1967.tb03435.x Giraldes BW, Smyth DM (2016) Recognizing Panulirus meripurpuratus sp. nov. (Decapoda: Palinuridae) in Brazil — systematic and biogeographic overview of Panulirus species in the Atlantic Ocean. Zootaxa 4107, 353–366. doi:10.11646/zootaxa.4107.3.4 Groeneveld JC, Griffiths CL, Van Dalsen AP (2006) A new species of spiny lobster, Panulirus barbarae (Decapoda, Palinuridae) from Walters Shoals on the Madagascar Ridge. Crustaceana 79, 821–833. doi:10.1163/156854006778008177 Holthuis LB (1991) FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue of species of interest to fisheries known to date. FAO Fisheries Synopsis 125, 1–292.

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Jeffs AG, Gardner C, Cockcroft A (2013) Jasus and Sagmariasus species. In Lobsters: biology, management, aquaculture and fisheries. (Ed. Phillips BF) pp. 259–288. Wiley-Blackwell, Chichester. Johnston DJ, Ritar A (2001) Mouthpart and foregut ontogeny in phyllosoma larvae of the spiny lobster Jasus edwardsii (Decapoda: Palinuridae). Marine and Freshwater Research 52, 1375–1386. doi:10.1071/MF01105 McCoy F (1890) Prodromus of the Zoology of Victoria: or figures and descriptions of the living species of all classes of the Victorian indigenous animals. Vol. 2. Decade 18. Government Printer, Melbourne. McWilliam PS (1995) Evolution in the phyllosoma and puerulus phases of the spiny lobster genus Panulirus White. Journal of Crustacean Biology 15, 542–557. doi:10.2307/1548775 Montgomery SS, Liggins GW (2013) Recovery of the eastern rock lobster Sagmariasus verreauxi off New South Wales, Australia. Marine Biology Research 9, 104–115. doi:10.1080/17451000.2012.727436 Palero F, Crandall KA, Abelló P, Macpherson E, Pascual A (2009a) Phylogenetic relationships between spiny, slipper and coral lobsters (Crustacea, Decapoda, Achelata). Molecular Phylogenetics and Evolution 50, 152–162. doi:10.1016/j.ympev.2008.10.003 Palero F, Lopes J, Abelló P, Macpherson E, Pascual M, et al. (2009b) Rapid radiation in spiny lobsters (Palinurus spp) as revealed by classic and ABC methods using mtDNA and microsatellite data. BMC Evolutionary Biology 9, 263. doi:10.1186/1471-2148-9-263 Palero F, Abelló P, Macpherson E, Matthee CA, Pascual M (2010) Genetic diversity levels in fishery-exploited spiny lobsters of the genus Palinurus (Decapoda: Achelata). Journal of Crustacean Biology 30, 658–663. doi:10.1651/09-3192.1 Patek S (2002) Squeaking with a sliding joint: mechanics and motor control of sound production in palinurid lobsters. Journal of Experimental Biology 205, 2375–2385. doi:10.1242/jeb.205.16.2375 Phillips BF, Rimmer DW, Reid DD (1978) Ecological investigations of the late stage Phyllosoma and Puerulus larvae of the Western Rock Lobster Panulirus longipes cygnus. Marine Biology 45, 347–357. doi:10.1007/ BF00391821 Poupin J (1994) The genus Justitia Holthuis, 1946, with the description of J. chani and J. vericeli spp. nov. (Crustacea: Decapoda: Palinuridea). Journal of Taiwan Museum 47, 37–56. Ptacek MB, Sarver SK, Childress MJ, Herrnkind WF (2001) Molecular phylogeny of the spiny lobster genus Panulirus (Decapoda: Palinuridae). Marine and Freshwater Research 52, 1037–1047. doi:10.1071/ MF01070 Sekiguchi H, George RW (2005) Description of Panulirus brunneiflagellum new species with notes on its biology, evolution, and fisheries. New Zealand Journal of Marine and Freshwater Research 39, 563–570. doi:10.1080/00288330.2005.9517335 Tsang LM, Chan T-Y, Cheung MK, Chu KH (2009) Molecular evidence for the Southern Hemisphere origin and deep sea diversification of

spiny lobsters (Crustacea: Decapoda: Palinuridae). Molecular Phylogenetics and Evolution 51, 304–311. doi:10.1016/j.ympev.​2009.01.015 Tsoi KH, Chan T-Y, Chu KH (2011) Phylogenetic and biogeographic analysis of the spear lobsters Linuparus (Decapoda: Palinuridae), with the description of a new species. Zoologischer Anzeiger 250, 302–315. doi:10.1016/j.jcz.2011.04.007

Scyllaridae Latreille, 1825 slipper lobsters, shovel-nosed lobsters Figures 11.3, 11.4, Plates 20, 21a-c Scyllarids, both adult and juvenile (Fig. 11.3g), are easily recognised by their flattened antennae that give some their common name of shovel-nosed lobsters. Local names vary but include slipper lobsters. The definitive works on their systematics are by Holthuis (1985, 2002, 2006). Several of the larger species are the target of pot and trawl fisheries (Holthuis 1991; Lavalli and Spanier 2007). Scyllarids are more diverse morphologically and ecologically than palinurid lobsters, which are associated with reefs. Like palinurids, the robust bodied arctidines and scyllarine scyllarids inhabit rock and coral reefs. Ibacine and thenine scyllarids, with flat, wide bodies, inhabit soft substrates and burrow into the sand or mud. Scyllarids have phyllosoma larvae like palinurids that have often been recorded associated with medusae (jelly fishes) in the open ocean (e.g. Wakabayashi et al. 2012). Of the four subfamilies recognised by Holthuis (1985), three were found to be monophyletic but one, Ibacinae, may be paraphyletic (Yang et  al. 2012). This finding was corroborated by Bracken-Grissom et al. (2014) and Palero et al. (2014) – both teams cast doubt on the monophyly of some scyllarine genera. The diagnoses and key to subfamilies and genera are based on those of Holthuis (1985, 1991, 2002). Relationships and morphological characterisation of genera was shown schematically by Yang et  al. (2012). Chan (2010) listed all species with their type localities. Diagnosis. Carapace flat; without prominent horns over eyestalks (supraocular spines). Antennal peduncle wide, flat, flagellum an unsegmented, wide flat plate.

Key to subfamilies and genera of Scyllaridae 1. – 2. – 3. –

Exopods of all maxillipeds with multiarticulate flagellum (Fig. 11.4q)����������������������������������������������������������������������������������2 Maxillipeds 1 and 3 exopods without flagellum; maxilliped 2 exopod flagellum transformed to a single laminate segment; maxilliped 3 exopod rudimentary����������������������������������������������������������������������������������������������������������6 Carapace strongly depressed, with a deep lateral cervical incision (Fig. 11.3c–e)������������������������������������������� Ibacinae … 3 Carapace highly vaulted, with shallow or no lateral cervical incision (Fig. 11.3a)�������������������������������������� Arctidinae … 5 Orbits entirely closed, slightly behind anterior margin of carapace. Cervical incision closed. Carapace with posteromedian tooth (Fig. 11.3c). Pleonite 5 without posteromedian spine (Fig. 11.3c)����������������������������������� Evibacus Orbits anteriorly open, on anterior margin of carapace. Cervical incision usually open, at least at the base. Carapace without posteromedian tooth (Fig. 11.3d, e)���������������������������������������������������������������������������������������������������������4

11 – Achelata – spiny lobsters, slipper lobsters and bugs

253

Fig. 11.3.  Scyllaridae. Dorsal views of carapace (and pleon). Arctidinae: a, Arctides; b, Scyllarides. Ibacinae: c, Evibacus; d, Ibacus; e, Parribacus. Scyllarinae: f, Antarctus; g, Crenarctus, puerulus stage; h, Eduarctus; i, Gibbularctus; j, Scammarctus. Theninae: k, Thenus.

4. – 5. – 6. – 7. –

Carapace dorsal surface smooth, punctate, not tuberculate, sometimes pubescent; with postrostral and branchial carinae. Orbits separated by distance shorter than or equal to distance between orbit and anterolateral angle. Pleonite 5 with posteromedian spine (Fig. 11.3d). Mandibular palp of 1 article����������������� Ibacus Carapace dorsal surface coarsely squamose-tuberculate, without postrostral or branchial ridges. Orbits separated by distance more than twice distance between orbit and anterolateral angle. Pleonite 5 without posteromedian spine (Fig. 11.3e). Mandibular palp 2-articled����������������������������������������������������������� Parribacus Pleonite 1 with dorsal transverse groove between anterior and posterior sections. Pleonites sculptured laterally. Carapace with postorbital spine (Fig. 11.3a)���������������������������������������������������������������������������������������������� Arctides Pleonite 1 without dorsal transverse groove. Pleonites with median ridge. Carapace without postorbital spine (Fig. 11.3b)���������������������������������������������������������������������������������������������������������������������������������������������������������Scyllarides Orbits on anterolateral angles of carapace; carapace strongly depressed (Fig. 11.3k)������������������������ Theninae … Thenus Orbits on anterior margin of carapace; carapace highly vaulted (Fig. 11.3f–j)���������������������������������������������Scyllarinae … 7 Pleonites without arborescent pattern of narrow grooves, with distinct, elevated median carina (Figs 11.3f, 11.4r)�����������������������8 Pleonites with arborescent pattern of narrow grooves, without distinct median carina (Pl. 20c, d, Fig. 11.4s, t)�������������� 11

254

Marine Decapod Crustacea

Fig. 11.4.  Scyllaridae. Thoracic sternum: a, Acantharctus; b, Gibbularctus. Anterior thoracic sternites: c, Bathyarctus; d, Biarctus; e, Crenarctus; f, Eduarctus; g, Galearctus; h, Petrarctus; i, Remiarctus; j, Scammarctus. Pereopod 2: k, Galearctus; l, Remiarctus. Pereopod 3: m, Chelarctus; n, Galearctus; o, Remiarctus. Pereopod 4: p, Scammarctus. Maxilliped 3, with exopod: q, Ibacus. Pleonites 1–3: r, Petrarctus; s, Biarctus; t, Galearctus. Carapace profile: u, Biarctus; v, Crenarctus; w, Scyllarus.

8.

Thoracic sternum anterior margin gutter-like, sunken, tapering, ending in depressed point, prolonged between bases of maxillipeds 3 (Fig. 11.4j). Pereopods 4, 5 unusually long, slender, pereopod 5 reaching base of antenna (Fig. 11.4p). Antennal article 4 with 1 oblique carina���������������������������������������������������������Scammarctus – Thoracic sternum anterior margin incised as V- or U-shape, truncate or convex. Pereopods 4, 5 not long, slender. Antennal article 4 with 1 or 2 oblique carinae (or row of tubercles)��������������������������������������������������������������������9 9. Thoracic sternum anterior margin truncate or convex, sometimes with median tubercle, without median incision (Fig. 11.4c)��������������������������������������������������������������������������������������������������������������������������������������������������Bathyarctus – Thoracic sternum anterior margin U- or V-shaped, with median incision (Fig. 11.4h)�����������������������������������������������������10 10. Antenna article 4 with 2 carinae, major one ending at distolateral angle. Pleonites 2–4 carinae of similar height (Fig. 11.3f)�������������������������������������������������������������������������������������������������������������������������������������������������������� Antarctus – Antenna article 4 with 1 carina, sometimes with row of tubercles on outer half. Pleonite 3 carina higher than that of pleonite 2 (Fig. 11.4r)�����������������������������������������������������������������������������������������������������������������������������Petrarctus 11. Antenna article 4 with major carina plus more lateral oblique row of tubercles (Fig. 11.3h, i)���������������������������������������� 12 – Antenna article 4 with 1 carina (Fig. 11.3 g)������������������������������������������������������������������������������������������������������������������������������13 12. Thoracic sternum anterior margin broadly U-shaped, with submedian tubercle each side of median incision (Fig. 11.4f). Pleonites with median carinae. Rostrum short, broad, truncate, constricted at base (Fig. 11.3h)��������������������������������������������������������������������������������������������������������������������������������������������������������������������Eduarctus – Thoracic sternum anterior margin V-shaped, without submedian tubercles (Fig. 11.4b). Pleonites without median carinae. Rostrum twice as wide as long, distally bilobed (Fig. 11.3i)����������������������������������������������Gibbularctus 13. Thoracic sternum with sharp median thorn on last segment (Fig. 11.4a)��������������������������������������������������������� Acantharctus – Thoracic sternum with at most central tubercle on last segment�������������������������������������������������������������������������������������������14

11 – Achelata – spiny lobsters, slipper lobsters and bugs

14. – 15. – 16. – 17. – 18. –

255

Pereopods 3 and 4 propodus with distinct distal tooth forming subchela with dactylus (Fig. 11.4m)��������������Chelarctus Pereopods 3 and 4 propodus simple (Fig. 11.4n, o)�������������������������������������������������������������������������������������������������������������������15 Pleonal pleura 2–4 ending in sharp posteriorly-directed point (Fig. 11.4t)��������������������������������������������������������������������������16 Pleonal pleura 2–4 apex blunt or rectangular-rounded (Fig. 11.4s)���������������������������������������������������������������������������������������17 Thoracic sternum anterior margin truncate, thickened (Fig. 11.4i). Pereopod 2 propodus as flat and wide as pereopod 3 propodus (Fig. 11.4l, o)������������������������������������������������������������������������������������������������������������������������Remiarctus Thoracic sternum anterior margin tongue-like, produced (Fig. 11.4g). Pereopod 2 propodus narrower than pereopod 3 propodus (Fig. 11.4k, n)������������������������������������������������������������������������������������������������������������������������Galearctus Thoracic sternum anterior margin with deep V-shaped incision formed of 2 anterolateral teeth, without additional tubercles (Fig. 11.4d). Pregastric tooth absent (Fig. 11.4u)������������������������������������������������������������������� Biarctus Thoracic sternum anterior margin U-shaped, incised, with 2 submedian tubercles (Fig. 11.4e). Pregastric tooth present (Fig. 11.4v, w)�����������������������������������������������������������������������������������������������������������������������������������������������������18 Pregastric and rostral teeth large, much larger than gastric and cardiac teeth (Fig. 11.4v)������������������������������� Crenarctus Gastric, pregastric and rostral teeth all well developed and pointed (Fig. 11.4w)����������������������������������������������������Scyllarus

Subfamily Arctidinae Holthuis, 1985 Diagnosis. Carapace highly vaulted, with or without small, shallow cervical incision. Maxillipeds all with multi-­ articulate flagellum.

surface smooth, tuberculate. Orbits separated by distance shorter than or equal to distance between orbit and anterolateral angle. Pleonite 5 without posteromedian spine. Maximum cl. 145 mm. Subtidal, shelf (2–90 m). Tropical Eastern Pacific. 1 species (Holthuis 1985).

Arctides Holthuis, 1960

Ibacus Leach, 1815

Spanish lobsters

Balmain bugs (Australian)

Diagnosis. Postorbial spine present. Pleonite 1 with dorsal transverse groove between anterior and posterior sections; pleonites 2–5 distinctly sculptured laterally. Maximum cl. 100 mm. Subtidal, shelf (1–146 m). Tropical Atlantic, Indo-West Pacific, Temperate Australasia. 3 species (Holthuis 2006: revision, key to species)

Diagnosis. Cervical incision usually open, at least at base; postrostral and branchial carinae distinct; carapace midline without posteromedian tooth; dorsal surface smooth, punctate, not tuberculate, sometimes pubescent. Orbits separated by distance shorter than or equal to distance between orbit and anterolateral angle. Pleonite 5 with posteromedian spine. Maximum cl. 80 mm (Pl. 21a). Subtidal–slope (4–750 m). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Australasia. 8 species (Brown and Holthuis 1998: key to species; Holthuis 1991). In southern Australia species are fished (Haddy et al. 2007).

Scyllarides Gill, 1898 Diagnosis. Postorbital spine absent. Pleonite 1 without dorsal transverse groove; pleonites 2–5 with median carina. Maximum cl. 170 mm (Pl. 20f). Intertidal–slope (0–380 m), Cosmopolitan except Arctic and Antarctic. 14 species (Holthuis 1991: diagnoses and key to 13 species; Holthuis 1993).

Subfamily Ibacinae Holthuis, 1985 Diagnosis. Carapace strongly depressed, with deep cervical incision on lateral margin. Maxillipeds all with multiarticulate flagellum. Evibacus Smith, 1869 shield fan lobster Diagnosis. Cervical incision closed; postrostral and branchial carinae distinct; carapace midline with posteromedian tooth; dorsal

Parribacus Dana, 1852 mitten lobsters Diagnosis. Cervical incision usually open, at least at base; postrostral and branchial carinae absent; carapace midline without posteromedian tooth; dorsal surface coarsely squamose-tuberculate, without postrostral or branchial ridges. Orbits separated by distance more than twice distance between orbit and anterolateral angle. Pleonite 5 without posteromedian spine. Maximum cl. 92 mm (Pl. 21b). Intertidal–shelf (0–23 m). Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific. 6 species (Holthuis 1991: diagnoses and key to species). The phyllosoma of Parribacus antarcticus (Lund, 1793) is 80 mm long, one of the largest marine larvae (Palero et al. 2014).

256

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Subfamily Scyllarinae Latreille, 1825

Biarctus Holthuis, 2002

The genus Scyllarus was subdivided by Holthuis (2002) who confined this genus to the Atlantic and introduced 12 new genera mostly confined to the Indo-West Pacific, Japan, temperate Australasia and Pacific South America. One genus has since been synonymised (Chan et  al. 2013). All the Indo-West Pacific genera and species were described in detail by Holthuis (2002). Genis-Armero et al. (2020) reallocated three species to other genera on the basis of molecular analysis (see below). Scyllarine slipper lobsters are too small to be fished. Diagnosis. Carapace highly vaulted, with weak cervical incision on lateral margin. Orbits on anterior margins of carapace. Maxillipeds 1 and 3 without flagellum, maxilliped 2 a single laminate segment.

Diagnosis. Carapace midline without pregastric tooth, with gastric and cardiac teeth. Thoracic sternum anterior margin with V-shaped incision formed of 2 anterolateral teeth. Pleonites 2–5 without median carinae; 2–4 with arborescent pattern of narrow grooves; pleonal pleura 2–4 apex blunt or rectangular-rounded. Antenna peduncle article 4 with 1 oblique dorsal carina. Maximum cl. 20 mm. Intertidal–shelf (0–73 m). Western and Central Indo-Pacific, Temperate Australasia. 4 species (Holthuis 2002).

Acantharctus Holthuis, 2002 Diagnosis. Carapace midline with pregastric, gastric and cardiac teeth. Rostrum without tooth, or with minute tooth. Thoracic sternum anterior margin U-shaped, incised, with 2 submedian tubercles; with strong median horn on last segment. Pleonites 2–4 with arborescent pattern of narrow grooves. Antenna peduncle article 4 with 1 oblique dorsal carina. Maximum cl. 30 mm. Shelf (20–57 m). Western and Central Indo-Pacific. 1 species. Holthuis (2002) included three species, the type species from the Indo-West Pacific and two from the Atlantic. The molecular analysis by Genis-Armero et al. (2020) placed the Atlantic species in Scyllarus and Crenarctus.

Antarctus Holthuis, 2002 Diagnosis. Carapace midline with pregastric, gastric and cardiac teeth. Rostrum with tooth. Thoracic sternum anterior margin with V-shaped incision formed of 2 anterolateral teeth. Pleonites 2–5 with prominent median carinae; 2–4 without clear sharp arborescent arrangement of narrow grooves. Antenna peduncle article 4 with 2 oblique dorsal carinae (major one ending at distolateral angle). Maximum cl. 48 mm (Pl. 20a). Shelf, slope (80–540 m). Temperate Australasia. 1 species (Holthuis 2002).

Chelarctus Holthuis, 2002 Diagnosis. Carapace midline with pregastric tooth, cardiac tooth represented by 2 low tubercles. Rostrum with tooth. Thoracic sternum anterior margin produced beyond anterolateral angles, tip with small median triangular incision. Pleonites 2–5 without median carinae; 2–4 with arborescent pattern of narrow grooves. Antenna peduncle article 4 with 1 oblique dorsal carina. Pereopods 3 and 4 propodi lower margin produced as tooth or spine, subchelate. Maximum cl. 72 mm (Pl. 20b). Subtidal–shelf (18–300 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 4 species (Yang and Chan 2012).

Crenarctus Holthuis, 2002 Diagnosis. Carapace midline with pregastric, gastric and cardiac teeth (pregastric, rostral teeth large, much larger than gastric and cardiac teeth). Rostrum with tooth. Thoracic sternum anterior margin U-shaped, incised, with 2 submedian tubercles; with pattern of short carinae and tubercles along sides. Pleonites 2–5 without median carinae; 2–4 with arborescent pattern of narrow grooves; pleonal pleura 2–4 apex blunt or rectangular-rounded. Antenna peduncle article 4 with 1 oblique dorsal carina, or with 2 oblique carinae, median one strong, ending in the top of the article, outer one shorter, ending in the lower lateral tooth. Maximum cl. 22 mm. Intertidal–slope (0–400 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate South America (Chile), Temperate Southern Africa, Temperate Australasia (Australia, New Zealand). 3 species (Holthuis 2002). Genis-Armero et al. (2020) added one species to Holthuis’s included species.

Bathyarctus Holthuis, 2002

Eduarctus Holthuis, 2002

Diagnosis. Carapace midline with pregastric, gastric and cardiac teeth. Rostrum without tooth. Thoracic sternum anterior margin straight, transverse, or arched, not incised in the middle. Pleonites 2–5 with prominent median carinae; 2–4 without clear sharp arborescent arrangement of narrow grooves; pleonite 4 carina slightly to distinctly higher than that of pleonite 3. Antenna peduncle article 4 with 2 oblique dorsal carinae. Maximum cl. 41 mm. Shelf, slope (26–732 m). Western and Central Indo-Pacific. 6 species (Holthuis 2002).

Diagnosis. Carapace midline with pregastric, gastric and cardiac teeth. Rostrum short, broad, truncate, constricted at base. Thoracic sternum anterior margin truncate, thickened, with minute median incision. Pleonites 2–5 with prominent median carinae, or with slightly elevated median carinae; 2–4 with arborescent pattern of narrow grooves. Antenna peduncle article 4 with major carina plus more lateral oblique row of tubercles. Maximum cl. 23 mm (Pl. 20c). Subtidal–slope (4–513 m). Temperate Northern W Pacific, IndoWest Pacific, Temperate Australasia. 8 species (Holthuis 2002).

11 – Achelata – spiny lobsters, slipper lobsters and bugs

Galearctus Holthuis, 2002 Diagnosis. Carapace midline without pregastric tooth, with gastric and cardiac teeth. Thoracic sternum anterior margin produced beyond anterolateral angles, tip with small median triangular incision. Pleonites 2–5 without median carinae; 2–4 without clear sharp arborescent arrangement of narrow grooves; pleonal pleura 2–4 ending in sharp posteriorly-directed point. Antenna peduncle article 4 with 1 oblique dorsal carina. Maximum cl. 39 mm. Subtidal–slope (5–680 m). Temperate Northern W Pacific, Central and Eastern Indo-Pacific, Temperate Australasia. 8 species (Yang and Chan 2010; Yang et al. 2011).

Gibbularctus Holthuis, 2002 Diagnosis. Carapace midline with pregastric, gastric and cardiac teeth. Rostrum twice as wide as long, distally bilobed. Thoracic sternum anterior margin with V-shaped incision formed of 2 anterolateral teeth. Pleonites 2–5 without median carinae; 2–4 with arborescent pattern of narrow grooves. Antenna peduncle article 4 with major carina plus more lateral oblique row of tubercles. Maximum cl. 15 mm. Subtidal, shelf (12–57 m). Western and Central Indo-Pacific. 1 species (Holthuis 2002).

Pleonites 2–5 with slightly elevated median carinae; 2–4 without clear sharp arborescent arrangement of narrow grooves. Antenna peduncle article 4 with 1 oblique dorsal carina. Pereopods 4, 5 especially long, slender, pereopod 5 reaching to base of antenna. Maximum cl. 33 mm (Pl. 20e). Shelf, slope (152–800 m). Western and Central Indo-Pacific. 1 species (Holthuis 2002).

Scyllarus Fabricius, 1775 Diagnosis. Carapace midline with pregastric, gastric and cardiac teeth. Rostrum with tooth. Thoracic sternum anterior margin U-shaped, incised, with 2 submedian tubercles. Pleonites 2–5 with prominent median carinae; 2–4 with arborescent pattern of narrow grooves; pleonal pleura 2–4 apex blunt or rectangularrounded. Antenna peduncle article 4 with 1 oblique dorsal carina. Maximum cl. 35 mm (Pl. 20d). Subtidal–slope (1–329 m). Temperate Northern and Tropical Atlantic, Temperate South America (SW Atlantic). 10 species. Genis-Armero et  al. (2017), using molecular evidence and morphology or late phyllosomas, suggested that two of the accepted species may be identical and have an amphi-Atlantic distribution. Genis-Armero et  al. (2020) provided a key to seven species of phyllosomas from the Atlantic.

Theninae Holthuis, 1985

Petrarctus Holthuis, 2002 Diagnosis. Carapace midline without pregastric tooth (or weak transverse carina or row of tubercles instead), with gastric and cardiac teeth. Rostrum without tooth. Thoracic sternum anterior margin U-shaped, with narrow median incision; without median tubercle and posterior tooth. Pleonites 2–5 with wide transverse median groove plus tubercles or ridges; 2–4 without clear sharp arborescent arrangement of narrow grooves; pleonite 3 carina distinctly higher than that of pleonite 2. Antenna peduncle article 4 with 1 oblique dorsal carina. Maximum cl. 27 mm. Subtidal–slope (5–415 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate ­Australasia. 6 species (Yang et al. 2008, 2017).

Remiarctus Holthuis, 2002 Diagnosis. Carapace midline without pregastric tooth, with gastric and cardiac teeth. Thoracic sternum anterior margin truncate, thickened, with minute median incision. Pleonites 2–4 with arborescent pattern of narrow grooves; pleonal pleura 2–4 ending in sharp posteriorly-directed point. Antenna peduncle article 4 with 1 oblique dorsal carina. Maximum cl. 21 mm. Subtidal, shelf (18–279 m). Temperate Northern W Pacific, Central Indo-Pacific. 1 species (Holthuis 2002).

Scammarctus Holthuis, 2002 Diagnosis. Carapace midline with pregastric, gastric and cardiac teeth. Rostrum without tooth. Thoracic sternum anterior part gutter-like, sunken, prolonged between bases of maxillipeds 3.

257

shovel-nosed lobsters, Moreton Bay bugs (Australia) Diagnosis. Carapace strongly depressed, with weak cervical incision on lateral margin. Orbits on anterolateral angles of carapace. Maxillipeds 1 and 3 without flagellum, maxilliped 2 a single laminate segment. Thenus Leach, 1816 Maximum cl. 94 mm (Pl. 21c). Subtidal, shelf (4–100 m). Western and Central Indo-Pacific. 5 species (Burton and Davie 2007). References Bracken-Grissom HD, Ahyong ST, Wilkinson RD, Feldmann RM, Schweitzer CE, et  al. (2014) The emergence of the lobsters: phylogenetic relationships, morphological evolution and divergence time comparisons of an ancient group (Decapoda: Achelata, Astacidea, Glypheidea, Polychelida). Systematic Biology 63, 457–479. doi:10.1093/sysbio/syu008 Brown DE, Holthuis LB (1998) The Australian species of the genus Ibacus (Crustacea: Decapoda: Scyllaridae), with description of a new species and addition of new records. Zoölogische Mededeelingen 72, 113–141. Burton TE, Davie PJF (2007) A revision of the shovel-nosed lobsters of the genus Thenus (Crustacea: Decapoda: Scyllaridae), with descriptions of three new species. Zootaxa 1429, 1–38. doi:10.11646/zootaxa.1429.1.1 Chan T-Y (2010) Annotated checklist of the world’s marine lobsters (Crustacea: Decapoda: Astacidea, Glypheidea, Achelata, Polychelida). Raffles Bulletin of Zoology Supplement 23, 153–181. Chan T-Y, Ahyong ST, Yang C-H (2013) Priority of the slipper lobster genus Crenarctus Holthuis, 2002, over Antipodarctus Holthuis, 2002 (Crustacea,

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Decapoda, Scyllaridae). Zootaxa 3701, 471–472. doi:10.11646/​zootaxa.​ 3701.4.7 De Man JG (1916) Eryonidae, Palinuridae, Scyllaridae and Nephropsidae. Siboga-Expéditie 23, 1–122, pls 1–4. Genis-Armero R, Guerao G, Abelló P, González-Gordillo JI, Cuesta JA, et  al. (2017) Possible amphi-Atlantic dispersal of Scyllarus lobsters (Crustacea: Scyllaridae): molecular and larval evidence. Zootaxa 4306, 325–328. doi:10.11646/zootaxa.4306.3.2 Genis-Armero R, González-Gordillo JI, Cuesta JA, Capaccioni-Azzati R, Palero F (2020) Revision of the West African species of Scyllarus Fabricius, 1775 (Decapoda: Achelata: Scyllaridae), with the description of three phyllosoma stages of S. caparti Holthuis, 1952 and an updated identification key. Journal of Crustacean Biology 40, 412–424. doi:10.1093/jcbiol/ruaa025 Haddy JA, Stewart J, Grahame KJ (2007) Fishery and biology of commercially exploited Australian fan lobsters (Ibacus spp.). In The biology and fisheries of the slipper lobster. (Eds Lavalli KL, Spanier E) pp. 359–376. CRC Press, Boca Raton. Holthuis LB (1985) A revision of the family Scyllaridae (Crustacea: Decapoda: Macrura). I. Subfamily Ibacinae. Zoölogische Verhandelingen 218, 1–130. Holthuis LB (1991) FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue of species of interest to fisheries known to date. FAO Fisheries Synopsis 125, 1–292. Holthuis LB (1993) Scyllarides obtusus spec. nov., the scyllarid lobster of Saint Helena, Central South Atlantic (Crustacea: Decapoda Reptantia: Scyllaridae). Zoölogische Mededeelingen 67, 505–515. Holthuis LB (2002) The Indo-Pacific scyllarine lobsters (Crustacea, Decapoda, Scyllaridae). Zoosystema 24, 499–683. Holthuis LB (2006) Revision of the genus Arctides Holthuis, 1960 (Crustacea, Decapoda, Scyllaridae). Zoosystema 28, 417–433. Lavalli KL, Spanier E (Eds) (2007) The Biology and Fisheries of the Slipper Lobster. CRC Press, Boca Raton.

McCoy F (1890) Prodromus of the Zoology of Victoria: or figures and descriptions of the living species of all classes of the Victorian indigenous animals. Vol. 2. Decade 18. Government Printer, Melbourne. Palero F, Guerao G, Hall M, Chan TY, Clark PF (2014) The ‘giant phyllosoma’ are larval stages of Parribacus antarcticus (Decapoda : Scyllaridae). Invertebrate Systematics 28, 258–276. doi:10.1071/IS13037 Wakabayashi K, Sato R, Hirai A, Ishii H, Akiba T, et al. (2012) Predation by the phyllosoma larva of Ibacus novemdentatus on various kinds of venomous jellyfish. Biological Bulletin 222, 1–5. doi:10.1086/ BBLv222n1p1 Yang C-H, Chan T-Y (2010) A new slipper lobster of the genus Galearctus Holthuis, 2002 (Decapoda, Scyllaridae) from Taiwan and Japan. In: Fransen CHJM, De Grave S, Ng PKL (Eds), Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume. Crustaceana Monographs 14, 735–745. Yang CH, Chan T-Y (2012) On the taxonomy of the slipper lobster Chelarctus cultrifer (Ortmann, 1897) (Crustacea: Decapoda: Scyllaridae), with description of a new species. Raffles Bulletin of Zoology 60, 449–460. Yang C, Chen I, Chan T-Y (2008) A new slipper lobster of the genus Petrarctus (Crustacea: Decapoda: Scyllaridae) from the West Pacific. Raffles Bulletin of Zoology Supplement 19, 71–81. Yang C-H, Chen I, Chan T-Y (2011) A new slipper lobster of the genus Galearctus Holthuis, 2002 (Crustacea, Decapoda, Scyllaridae) from New Caledonia. Zoosystema 33, 207–217. doi:10.5252/z2011n2a4 Yang C-H, Bracken-Grissom H, Kim D, Crandall KA, Chan T-Y (2012) Phylogenetic relationships, character evolution, and taxonomic implications within the slipper lobsters (Crustacea: Decapoda: Scyllaridae). Molecular Phylogenetics and Evolution 62, 237–250. doi:10.1016/j. ympev.2011.09.019 Yang CH, Kumar AB, Chan T-Y (2017) A new slipper lobster of the genus Petrarctus Holthuis, 2002 (Crustacea, Decapoda, Scyllaridae) from Southwest coast of India. Zootaxa 4329, 477–486. doi:10.11646/ zootaxa.4329.5.5

12. Polychelida – deep-sea lobsters

Lobsters of the infraorder Polychelida Scholtz & Richter, 1995 are unusual in having chelate pereopods 1–4, pereopod 5 being chelate in one or both sexes, and reduced eyes in extant species (Ahyong 2009; Scholtz and Richter 1995). The first pair of chelipeds are exceptionally long and thin. The group has a rich fossil history dating back to the Triassic but only one family is extant (Ahyong 2009; Audo et  al. 2021; Bracken-Grissom et  al. 2014; Galil 2000; Karasawa et  al. 2013). Polychelidan lobsters are essentially deep-water species but little is known of their benthic ecology (Abelló and Cartes 1992). Notably, although primarily benthic, adult polychelids can swim forwards in open water via beating of the pleopods, much like that of ‘natant’ shrimps (ArnésUrgellés et al. 2020). Polychelids have a midwater ‘eryoneicus’ larva (Fig. 12.1a, Pl. 21d), so named after the genus to which it was first assigned (Eller et al. 2016; Torres et al. 2014). Polychelidan taxonomy is well documented. Galil (2000) introduced two new genera and described all species then known. Her work was updated on the basis of a morphological phylogeny by Ahyong (2009) who recognised Stereomastis, previously synonymised with Polycheles, bringing the

number of genera to six. Molecular phylogenies suggest that Polycheles may be polyphyletic (Bracken-Grissom et  al. 2014; Rengaiyan et al. 2020). Diagnosis. Carapace flat, with lateral edge, longer than wide; epistome short, not fused to basal antennular and antennal articles, lateral margins with narrow point of contact with carapace margins. Exoskeleton chitinised. Thoracic sternites 1–8 fused. Pleon elongate, muscular; pleonite 2 pleuron overlapping pleonites 1 and 3. Antennular penduncle articles aligned linearly, main flagellum long, lashlike, aesthetascs more-or-less evenly distributed. Pereopods 1–4 or 1–5 chelate, first longest, thin. Pereopod 5 shorter than other pereopods. Uropods biramous.

Polychelidae Wood-Mason, 1875 Figure 12.1, Plate 21d–f Implicit generic attributes. Major chela fixed finger without perpendicular spine on inner margin. Pereopods 1–5 with epipod well developed, markedly longer than coxal width.

Key to genera of Polychelidae 1. – 2. – 3. – 4. – 5. –

Carapace ovate, slightly long than wide (Fig. 12.1b)��������������������������������������������������������������������������������������������������������Cardus Carapace distinctly longer than wide (Fig. 12.1c–e)�������������������������������������������������������������������������������������������������������������������2 Dorsal orbital sinus obsolete, shallowly concave (Fig. 12.1g). Major chela fixed finger with perpendicular spine on inner margin (Fig. 12.1m). Basal antennular article anterolateral margin unarmed�������������������Willemoesia Dorsal orbital sinus deep, slit-like, U- or V-shaped (Fig. 12.1b–f). Major chela fixed finger without perpendicular spine on inner margin. Basal antennular article anterolateral margin with 1 or more spines�������������������������3 Dorsal orbital sinus U-shaped (Fig. 12.1e). Pereopods 1–5 with epipod reduced, shorter than coxal width (Fig. 12.1q)����������������������������������������������������������������������������������������������������������������������������������������������������������������Stereomastis Dorsal orbital sinus V-shaped or slit-like (Fig. 12.1c, d, f). Pereopods 1–5 with epipod well developed, markedly longer than coxal width (Fig. 12.1p)�����������������������������������������������������������������������������������������������������������������������4 Basal antennular article anterolateral margin rounded, with 1–3 small spines (Fig. 12.1h). Maxilliped 3 epipod vestigial (Fig. 12.1l)���������������������������������������������������������������������������������������������������������������������������������������� Polycheles Basal antennular article anterolateral margin quadrate. Maxilliped 3 epipod well developed (Fig. 12.1j, k)�������������������5 Pereopods 2–4 fixed fingers and dactyli strongly curved (Fig. 12.1n). Basal antennular segment with 2 anterolateral spines. Maxilliped 3 epipod one-third to half length of ischium (Fig. 12.1j)���������������������������� Homeryon Pereopods 2–4 fixed fingers and dactyli straight or weakly curved (Fig. 12.1o). Basal antennular segment with 1 anterolateral spine. Maxilliped 3 epipod as long as or longer than ischium (Fig. 12.1k)����������������� Pentacheles Cardus Galil, 2000

Diagnosis. Carapace ovate, slightly longer than wide; dorsal orbital sinus deep, slit-like. Basal antennular article anterolateral margin

rounded, with 1 or 2 anterolateral spines. Maxilliped 3 epipod as long as or longer than ischium. Pereopods 2–4 fixed fingers and dactyli straight or weakly curved. Maximum cl. 30 mm.

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Fig. 12.1.  Polychelidae. a, eryoneicus larva; b, Cardus crucifer (Thomson, 1873); c, Homeryon ; d, Pentacheles laevis Bate, 1878; e, Stereomastis nana (Smith, 1884). Front: f, Polycheles; g, Willemoesia. Antennule basal articles: h, Polycheles; i, Stereomastis. Maxilliped 3: j, Homeryon; k, Pentacheles; l, Polycheles. m, pereopod 1 chela, Willemoesia. Pereopod 4 chela: n, Homeryon; o, Pentacheles. Pereopod 3: p, Polycheles; q, Stereomastis.

Slope–bathyal (549–2195 m). Temperate Northern and Tropical Atlantic. 1 species (Ahyong 2009: rediagnosis; Galil 2000: diagnosis, species redescription).

Homeryon Galil, 2000 Diagnosis. Carapace distinctly longer than wide; dorsal orbital sinus deep, V-shaped. Basal antennular article anterolateral margin quadrate, with 2 anterolateral spines. Maxilliped 3 epipod one-third to half length of ischium. Pereopods 2–4 fixed fingers and dactyli strongly curved. Maximum cl. 65 mm. Slope (520–1557 m). Temperate Northern W Pacific, Eastern Indo-Pacific. 2 species (Ahyong 2009: rediagnosis, key to species; Galil 2000: diagnosis, species redescriptions; Komai and Tsuchida 2014: figures of H. armarium Galil, 2000)

Pentacheles Bate, 1878 Diagnosis. Carapace distinctly longer than wide; dorsal orbital sinus deep, V-shaped. Basal antennular article anterolateral margin quadrate, with 1 anterolateral spine. Maxilliped 3 epipod as

long as or longer than ischium. Pereopods 2–4 fixed fingers and dactyli straight or weakly curved. Maximum cl. 77 mm. Slope–bathyal (347–3365 m). Cosmopolitan except polar seas. 5 species (Ahyong 2009: rediagnosis, key to species; Galil 2000: rediagnosis, species redescriptions).

Polycheles Heller, 1862 Diagnosis. Carapace distinctly longer than wide; dorsal orbital sinus deep, V-shaped. Basal antennular article anterolateral margin rounded, with 1–3 small spines. Maxilliped 3 epipod vestigial. Pereopods 2–4 fixed fingers and dactyli straight or weakly curved. Maximum cl. 55 mm (Pl. 12e). Shelf–bathyal (41–2195 m). Cosmopolitan except polar seas. 9 species (Ahyong 2009: rediagnosis, key to species; Galil 2000: rediagnosis, species redescriptions).

Stereomastis Bate, 1888 Diagnosis. Carapace distinctly longer than wide; dorsal orbital sinus U-shaped. Basal antennular article anterolateral margin

12 – Polychelida – deep-sea lobsters

rounded, with 1 or 2 small spines. Maxilliped 3 epipod vestigial. Pereopods 2–4 fixed fingers and dactyli straight or weakly curved; pereopods 1–5 with epipods reduced, shorter than coxal width. Maximum cl. 54 mm (Pl. 12f). Shelf–abyssal (144–4000 m). Cosmopolitan except polar seas. 19 species. Species of Stereomastis were treated as species of Polycheles by most authors including Galil (2000). The genus was revived and rediagnosed by Ahyong (2009) who provided a key to species.

Willemoesia Grote, 1873 Diagnosis. Carapace distinctly longer than wide; dorsal orbital sinus obsolete, shallowly concave. Basal antennular article anterolateral margin quadrate, without anterolateral spine. Maxilliped 3 epipod as long as or longer than ischium. Major chela fixed finger with perpendicular spine on inner margin. Pereopod 3 fixed finger and dactylus crossing; pereopod 4 fixed finger and dactylus weakly curved. Maximum cl. 81 mm. Slope–abyssal (1760–5124 m). Cosmopolitan except polar seas. 4 species (Ahyong 2009: rediagnosis, key to species; Galil 2000: diagnosis, species redescriptions). References Abelló P, Cartes JE (1992) Population characteristics of the deep-sea lobsters Polycheles typhlops and Stereomastis sculpta (Decapoda: Polychelidae) in a bathyal mud community of the Mediterranean Sea. Marine Biology 114, 109–117. Ahyong ST (2009) The polychelidan lobsters: phylogeny and systematics (Polychelida: Polychelidae). In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 309–326. CRC Press, Boca Raton. Arnés-Urgellés C, Buglass S, Ahyong ST, Salinas-de-León P, Wicksten MK, et  al. (2020) Arthropoda; Crustacea; Decapoda of deep-sea

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volcanic habitats of the Galapagos Marine Reserve, Tropical Eastern Pacific. Biodiversity Data Journal 8, e54482. doi:10.3897/BDJ.8.e54482 Audo D, Barriel V, Charbonnier S (2021) Phylogeny and evolutionary history of polychelidan lobsters. Journal of Systematic Palaeontology 19, 417–439. doi:10.1080/14772019.2021.1918773 Bracken-Grissom HD, Ahyong ST, Wilkinson RD, Feldmann RM, Schweitzer CE, et al. (2014) The emergence of the lobsters: phylogenetic relationships, morphological evolution and divergence time comparisons of an ancient group (Decapoda: Achelata, Astacidea, Glypheidea, Polychelida). Systematic Biology 63, 457–479. doi:10.1093/sysbio/ syu008 Eller S, Haug C, Haug JT (2016) Detailed description of a giant polychelidan eryoneicus-type larva with modern imaging techniques. Spixiana 39, 39–60. Galil BS (2000) Crustacea Decapoda: review of the genera and species of the family Polychelidae Wood-Mason, 1874. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 21. Mémoires du Muséum National d’Histoire Naturelle, Paris 184, 285–387. Karasawa H, Schweitzer CE, Feldmann RM (2013) Phylogeny and systematics of extant and extinct lobsters. Journal of Crustacean Biology 33, 78–123. doi:10.1163/1937240X-00002111 Komai T, Tsuchida S (2014) Deep-sea decapod crustaceans (Caridea, Polychelida, Anomura and Brachyura) collected from the Nikko seamounts, Mariana Arc, using a remotely operated vehicle “Hyper-Dolphin”. Zootaxa 3764, 279–316. doi:10.11646/zootaxa.3764.3.3 Rengaiyan P, Fernandes SO, Kurian PJ, Ingole B (2020) Molecular identification of deep-sea blind lobster Willemoesia forceps (Crustacea: Decapoda: Polychelidae) from the Central Indian Ridge. Mitochondrial DNA. Part B, Resources 5, 1013–1014. doi:10.1080/23802359.2020.1721345 Scholtz G, Richter S (1995) Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca). Zoological Journal of the Linnean Society 113, 289–328. doi:10.1006/zjls.1995.0011 Torres AP, Palero F, Dos Santos A, Abelló P, Blanco E, et al. (2014) Larval stages of the deep-sea lobster Polycheles typhlops (Decapoda, Polychelida) identified by DNA analysis: morphology, systematic, distribution and ecology. Helgoland Marine Research 68, 379–397. doi:10.1007/s10152-014-0397-0

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13. Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters The Anomura H. Milne Edwards, 1832 are a group of decapods, which, on first appearance, seem to have little in common; some look like crabs, others are hermit crabs, others resemble small lobsters, and others not easily placed in one or other of these well known crustacean types. The only easy character uniting them is the small folded fifth thoracic leg. To further complicate the issue, animals of any one of these forms cannot be easily be placed in one or other of the six superfamilies into which the infraorder is classified. Crab-like forms occur in the Paguroidea, Galatheoidea and Hippoidea. None of the lobster-like anomuran crustaceans have strong pleons like lobsters but some galatheoids are often called squat lobsters. And hermit crabs, easily recognisable as such, are distributed among six families. The anomurans form the sister group to the true crabs Brachyura, in a group Scholtz and Richter (1995) called Meiura, and which has since been corroborated by several phylogenetic analyses (Ahyong and O’Meally 2004; Dixon et al. 2003; Tsang et al. 2008; Wolfe et al. 2019). Molecular evidence goes some way to explaining the uncertainty surrounding the relationships of the established anomuran families and superfamilies. Pérez-Losada et  al. (2002) reviewed views based on morphology dating from the middle of the nineteenth century and provided a new interpretation based on 18S rDNA. They recovered the accepted composition of Galatheoidea (Porcellanidae, Chirostylidae, Galatheidae and Aeglidae) and differed only in the relative position of the families from earlier studies (Martin and Abele 1986; Tudge 1997a). Pérez-Losada et  al. (2002) and Ahyong and O’Meally (2004) found that Galatheoidea and Paguroidea are more closely related to each other than to Hippoidea, a view not shared by Morrison et al. (2002), who placed ­Hippoidea closest to Paguroidea. Extensive recent molecular evidence, sometimes combined with morphology, recovered polyphyly of Galatheoidea and Paguroidea and placed ­ Hippoidea as sister group to the the remaining anomurans (Ahyong et  al. 2009; Bracken-Grissom et  al. 2013; Schnabel et al. 2011; Tsang et al. 2011). Ahyong et al. (2011) reviewed the competing perspectives on the phylogeny and fossil record of the squat lobster groups. It has long been recognised that the larvae of the crab-like lithodids resemble those of the pagurid hermit crabs (Noever and Glenner 2018; Richter and Scholtz 1994). Cunningham et  al.’s (1992) cladogram based on DNA sequences from mitochondrial rRNA placed two genera of Lithodidae within the genus Pagurus, a finding that at least reinforces the

polyphyly of this large genus and of the Paguridae. It seems certain that carcinisation, evolution of a crab-like habitus, has evolved several times in the Anomura (Tsang et al. 2011). This phenomenon was discussed in detail by McLaughlin and Lemaitre (1997) but they disagreed with Cunningham et al.’s conclusions. More recently, detailed studies of internal structures have been undertaken to understand the evolutionary pathways taken by anomurans (e.g. Keiler et al. 2013; Keiler et al. 2015, 2016; Keiler et al. 2017). The name of this group has been debated, being sometimes termed Anomala, and has variously included several groups of anomalous decapods. McLaughlin and Holthuis (1985) agreed on the widely used name Anomura, which is now confined to six superfamilies: Aegloidea for South American freshwater squat lobsters (not included here), Galatheoidea and Chirostylidae for marine squat lobsters and porcelain crabs, Hippoidea for mole crabs, Paguroidea for hermit crabs and king crabs, and Lomisoidea for Lomis hirta (Lamarck, 1818) alone. The classification of Anomura has a convoluted history (Davie 2002; Martin and Abele 1986; Martin and Davis 2001; McLaughlin 1983a, 2003; McLaughlin and Lemaitre 2001); McLaughlin et al. 2007; De Grave et al. 2009). The enigmatic monotypic family Lomisidae has often been treated as a superfamily (Martin and Davis 2001; McLaughlin 1983b). Larval morphology suggests it is possibly related to Lithodidae, the structure of the spermatozoa suggests a basal position within Anomura (Tudge 1997b), but total evidence places the species close to Aegloidea and Chirostyloidea (Bracken-Grissom et  al. 2013; Tsang et  al. 2011). Its place in the key to superfamilies is quite artificial. Important questions have been raised over the monophyly of the hermit crabs and their classification in one or more superfamilies (Paguroidea, Coenobitoidea, Lithodoidea) (Bracken-Grissom et  al. 2013; McLaughlin et  al. 2010; McLaughlin et al. 2007; Tsang et al. 2011). Given the present-day uncertainty, the superfamilies Coenobitoidea and Lithodoidea are not recognised here; their members are placed with the Paguroidea. Diagnosis. Carapace more or less cylindrical, longer than wide, or flat, usually with lateral edge, usually wider or as long as wide, or flat, with lateral edge, longer than wide, or more or less globular; epistome short, not fused to basal antennular and antennal articles, lateral margins with narrow point of contact with carapace margins. Exoskeleton calcified, or chitinised. Thoracic sternite 8 free from thoracic

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sternites 1–7. Pleon short, flat, not or weakly folded under carapace, or coiled, unchitinised; pleonite 2 pleuron overlapping pleonites 1 and 3, or all pleonal pleura overlapping next most posterior pleuron, or not overlapping, or pleonites fused, without pleura. Maxilliped 3 with fewer than 7 free articles. Antennular peduncle articles folding in Z-shape or

linearly (Hippoidea), main flagellum usually short, thick, compact (except Coenobitidae, Hippoidea), aesthetascs concentrated on one side forming dense brush. Pereopod 1 chelate, claw-like; pereopods 2–4 usually simple, sometimes subchelate. Pereopod 5 shorter than others, subchelate or chelate. Uropods biramous or absent.

Key to marine superfamilies of Anomura 1. – 2. – 3. – 4. –

Pleon usually asymmetrical (Figs 13.18–20, 13.22, 13.27) (symmetrical in most Pylochelidae, Fig. 13.29). Pereopod 4 reduced, smaller than 2 and 3 (except in Lithodidae). Ocular scales present (Fig. 13.19, 13.23, 13.28) (reduced in Lithodidae)������������������������������������������������������������������������������������������������������������������������������� Paguroidea Pleon symmetrical. Pereopod 4 ambulatory, like 2 and 3 (Figs 13.13, 13.16). Ocular scales absent������������������������������������2 Uropods absent (or invisible in natural view and pleopod-like only in female) (Fig. 13.16)�����������������������������������������������������������������������������������������������������������������������������������������Lomisoidea (Lomis hirta) Uropods present and visible in natural position (Figs 13.3, 13.6)���������������������������������������������������������������������������������������������3 Pereopods 1 short, chelate or subchelate. Pereopods 2–4 with flattened dactyli. Telson longer than wide, not forming tail-fan with uropods (Figs 13.13)�����������������������������������������������������������������������������������������������������Hippoidea Pereopods 1 elongate chelipeds. Pereopods 2–4 with claw-like dactyli. Telson and uropods forming tail-fan����������������4 Sternal plate on thoracic somite 8, between last legs, well developed (Fig. 13.7l). Telson divided into median and paired plates, often in X-like pattern (Fig. 13.5r, 13.9f). Maxilliped 3 with or without epipod�������� Galatheoidea Sternal plate on thoracic somite 8, between last legs, absent, replaced entirely by arthrodial membrane (Fig. 13.1k, l). Telson transversely divided into 2 lobes (sometimes barely discernible), laterally incised (Fig. 13.1m, n, 13.2c). Maxilliped 3 without epipod�������������������������������������������������������������������������������������� Chirostyloidea

References Ahyong ST, O’Meally D (2004) Phylogeny of the Decapoda Reptantia: resolution using three molecular loci and morphology. Raffles Bulletin of Zoology 52, 673–693. Ahyong ST, Schnabel KE, Maas EW (2009) Anomuran phylogeny: new insights from molecular data. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 399–414. CRC Press, Boca Raton. Ahyong ST, Schnabel KE, Macpherson E (2011) Chapter 3. Phylogeny and fossil record of the marine squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 73–104. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Bracken-Grissom HD, Cannon ME, Cabezas P, Feldmann RM, Schweitzer CE, et  al. (2013) A comprehensive and integrative reconstruction of evolutionary history for Anomura (Crustacea: Decapoda). BMC Evolutionary Biology 13, 128 doi:10.1186/1471-2148-13-128. Cunningham CW, Blackstone NW, Buss LW (1992) Evolution of king crabs from hermit crab ancestors. Nature 355, 539–542. doi:10.1038/355539a0 Davie PJF (2002) Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells A, Houston WWK, Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing, Melbourne. Dixon CJ, Ahyong ST, Schram FR (2003) A new hypothesis of decapod phylogeny. Crustaceana 76, 935–975. doi:10.1163/156854003771997846 Keiler J, Richter S, Wirkner CS (2013) Evolutionary morphology of the hemolymph vascular system in hermit and king crabs (Crustacea: Decapoda: Anomala). Journal of Morphology 274, 759–778. doi:10.1002/jmor.20174 Keiler J, Richter S, Wirkner CS (2015) The anatomy of the king crab Hapalogaster mertensii Brandt, 1850 (Anomura: Paguroidea: Hapalogastridae) – new insights into the evolutionary transformation

of hermit crabs into king crabs. Contributions to Zoology 84, 149–165. doi:10.1163/18759866-08402004 Keiler J, Richter S, Wirkner CS (2016) Revealing their innermost secrets: an evolutionary perspective on the disparity of the organ systems in anomuran crabs (Crustacea: Decapoda: Anomura). Contributions to Zoology 85, 361–386. doi:10.1163/​18759866-08504001 Keiler J, Wirkner CS, Richter S (2017) One hundred years of carcinization – the evolution of the crab-like habitus in Anomura (Arthropoda: Crustacea). Biological Journal of the Linnean Society 121, 200–222. doi:10.1093/biolinnean/blw031 Martin JW, Abele LG (1986) Phylogenetic relationships of the genus Aegla (Decapoda: Anomura: Aeglidae), with comments on anomuran phylogeny. Journal of Crustacean Biology 6, 576–616. doi:10.2307/1548195 Martin JW, Davis GE (2001) An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County Science Series 39, 1–124. McLaughlin PA (1983a) Hermit crabs  – are they really polyphyletic? Journal of Crustacean Biology 3, 608–621. doi:10.2307/1547956 McLaughlin PA (1983b) A review of the phylogenetic position of the Lomidae (Crustacea: Decapoda: Anomala). Journal of Crustacean Biology 3, 431–447. doi:10.2307/1548143 McLaughlin PA (2003) Illustrated keys to families and genera of the superfamily Paguroidea (Crustacea: Decapoda: Anomura), with diagnoses of genera of Paguridae. Memoirs of Museum Victoria 60, 111–144. doi:10.24199/j.mmv.2003.60.16 McLaughlin PA, Holthuis LB (1985) Anomura versus Anomala. Crustaceana 49, 204–209. doi:10.1163/156854085X00468 McLaughlin PA, Lemaitre R (1997) Carcinization in the Anomura – fact or fiction? I. Evidence from adult morphology. Contributions to Zoology 67, 79–123. doi:10.1163/​18759866-06702001

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

McLaughlin PA, Lemaitre R (2001) A new family for a new genus and new species of hermit crab of the superfamily Paguroidea (Decapoda: Anomura) and its phylogenetic implications. Journal of Crustacean Biology 21, 1062–1076. doi:10.1163/20021975-99990198 McLaughlin PA, Lemaitre R, Sorhannus U (2007) Hermit crab phylogeny: a reappraisal and its “fall-out”. Journal of Crustacean Biology 27, 97–115. doi:10.1651/S-2675.1 McLaughlin PA, Komai T, Lemaitre R, Rahayu DL (2010) Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea) Part I – Lithodoidea, Lomisoidea and Paguroidea. Raffles Bulletin of Zoology Supplement 23, 5–107. Morrison CL, Harvey AW, Lavery S, Tieu K, Huang Y, et al. (2002) Mitochondrial gene rearrangements confirm the parallel evolution of the crab-like form. Proceedings. Biological Sciences 269, 345–350. doi:10.1098/​rspb.​2001.1886 Noever C, Glenner H (2018) The origin of king crabs: hermit crab ancestry under the magnifying glass. Zoological Journal of the Linnean Society 182, 300–318. doi:10.1093/zoolinnean/zlx033 Pérez-Losada M, Jara CG, Bond-Buckup G, Porter ML, Crandall KA (2002) Phylogenetic position of the freshwater anomuran family Aeglidae. Journal of Crustacean Biology 22, 670–676. doi:10.1163/ 20021975-99990279 Richter S, Scholtz G (1994) Morphological evidence for a hermit crab ancestry of lithodids (Crustacea, Decapoda, Anomala, Paguroidea). Zoologischer Anzeiger 233, 187–210. Schnabel KE, Ahyong ST, Maas EL (2011) Galatheoidea are not monophyletic  – molecular and morphological phylogeny of the squat lobsters (Decapoda: Anomura) with recognition of a new superfamily. Molecular Phylogenetics and Evolution 58, 157–168. doi:10.1016/j.ympev.​2010.11.011 Scholtz G, Richter S (1995) Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca). Zoological Journal of the Linnean Society 113, 289–328. doi:10.1006/zjls.1995.0011 Tsang LM, Ma KY, Ahyong ST, Chan T-Y, Chu KH (2008) Phylogeny of Decapoda using two nuclear protein-coding genes: Origin and evolution of the Repantia. Molecular Phylogenetics and Evolution 48, 359–368. doi:10.1016/j.ympev.2008.04.009 Tsang LM, Chan T-Y, Ahyong ST, Chu KH (2011) Hermit to king, or hermit to all: multiple transitions to crab-like forms from hermit crab ancestors. Systematic Biology 60, 616–629. doi:10.1093/sysbio/ syr063 Tudge CC (1997a) Phylogeny of the Anomura (Decapoda, Crustacea): Spermatozoa and spermatophore morphological evidence. Contributions to Zoology 67, 125–141. doi:10.1163/18759866-06702002 Tudge CC (1997b) Spermatological evidence supports the taxonomic placement of the Australian endemic hairy stone crab, Lomis hirta (Decapoda: Anomura: Lomidae). Memoirs of the Museum of Victoria 56, 235–244. doi:10.24199/j.mmv.1997.56.09 Wolfe JM, Breinholt JW, Crandall KA, Lemmon AR, Lemmon EM, et  al. (2019) A phylogenomic framework, evolutionary timeline and genomic resources for comparative studies of decapod crustaceans. Proceedings. Biological Sciences 286, 20190079. doi:10.1098/rspb.2019.0079

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Chirostyloidea Ortmann, 1892 Chirostyloid squat lobsters differ from galatheoid squat lobsters in lacking thoracic sternite 8, the telson transversely divided into two lobes by a suture, the antennal peduncle of five rather than four free articles, and the mandibular incisor margin strongly serrated or tridentate (it is entire in Galatheoidea). Chirostyloids tend to occur in deeper water than galatheoids. A detailed ecological account is reported for each family in turn. The biology of squat lobsters in general has been extensively reviewed (Poore et al. 2011). Baba et al. (2011a) dealt with morphology, Macpherson and Baba (2011) with taxonomy, Ahyong et al. (2011) with phylogeny and the fossil record, Baba et al. (2011b) with developmental biology, Schnabel et  al. (2011b) with distribution patterns, Lovrich and Thiel (2011) with ecology, physiology, feeding and trophic role, Thiel and Lovrich (2011) with agnostic behaviour and reproductive biology, Baeza (2011) with life in chemo-autotrophic environments, and Boyko and Williams (2011) with parasites and other symbionts. Until recently, the chirostyloids were grouped with the galatheid squat lobsters, all within Galatheoidea (e.g. Baba et  al. 2008) but phylogenetic studies show they belong in separate superfamilies, with chirostyloids more closely related to aegloids, lomisoids and hermit crabs (Ahyong et  al. 2009; Bracken-Grissom et  al. 2013; Schnabel et  al. 2011a; Tsang et al. 2011). The most important recent synopses of chirostyloids are Baba (2005, 2018). Since then, the superfamily has been reviewed again by Schnabel (2020) concentrating on the fauna of New Zealand. Her monograph covered ecology, taxonomy, identification and phylogeny. She and others before recognised four families. A molecular (COI) phylogeny supports the monophyly of all four families but not that of all chirostylid genera. Diagnosis. Sternal plastron consisting of thoracic sternites 3–7, thoracic sternite 8 absent (Fig. 13.1l). Pleon symmetrical; pleonites separate, calcified and distinct. Ocular scales absent. Antennal peduncle of 5 articulating articles. Mandibular incisor margin strongly serrated along its length, or tridentate. Pereopods 1 symmetrical, elongate, chelate. Pereopods 2–4 with claw-like dactyli. Pereopod 4 ambulatory, like 2 and 3. Uropods broad, part of tail-fan. Telson transversely folded, laterally incised (Figs 13.1m, n, 13.2c).

Key to families of Chirostyloidea 1. –

Insertion of pereopod 5 not visible ventrally, situated beneath sternal plastron. Sternite between left and right maxilliped 3 large, strongly produced anteriorly (Fig. 13.3d). Eyes strongly reduced (Fig. 13.3a). Antennal scale absent (Fig. 13.3a)����������������������������������������������������������������������������������������������������������������������������� Kiwaidae Insertion of pereopod 5 clearly visible ventrally. Sternite between left and right maxilliped 3 small, not produced anteriorly; eyes well developed. Antennal scale usually present������������������������������������������������������������������������2

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2. – 3.



Marine Decapod Crustacea

Supraocular spines present (Fig. 13.2)�������������������������������������������������������������������������������������������������������������������� Eumunididae Supraocular spines absent (Figs 13.1, 13.4)�����������������������������������������������������������������������������������������������������������������������������������7 Thoracic sternite 3 abruptly demarcated from preceding sternites 1 and 2 by distinct step forming well defined anterior margin at articulation with maxillipeds 3 (Fig. 13.1i–k). Maxilliped 3 pair widely separated, distal parts accommodated in excavated sternum between left and right maxillipeds 3 when folded. Pereopods 2–4 dactyli with terminal spine clearly demarcated by suture, often slightly movable������������������������������������Chirostylidae Thoracic sternite 3 confluent with sternites 1 and 2, without abrupt transverse step at maxilliped 3 articulations, produced and sloping anterodorsally, with pair of spines directly behind anterior margin. Maxilliped 3 pair close (Fig. 13.4b). Pereopods 2–4 ending in fixed, corneous unguis, demarcation between unguis and remaining dactylus diffuse������������������������������������������������������������������������������������������� Sternostylidae

References Ahyong ST, Schnabel KE, Maas EW (2009) Anomuran phylogeny: new insights from molecular data. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 399–414. CRC Press, Boca Raton. Ahyong ST, Schnabel KE, Macpherson E (2011) Chapter 3. Phylogeny and fossil record of the marine squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 73–104. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Baba K (2005) Deep-sea chirostylid and galatheid crustaceans (Decapoda: Anomura) from the Indo-West Pacific, with a list of species. Galathea Report 20, 1–317. Baba K (2018) Chirostylidae of the Western and Central Pacific: Uroptychus and a new genus (Crustacea: Decapoda: Anomura. In: Tropical Deep-Sea Benthos vol. 30. Mémoires du Muséum National d’Histoire Naturelle, Paris 212, 1–612. Baba K, Macpherson E, Poore GCB, Ahyong ST, Bermudez A, et al. (2008) Catalogue of squat lobsters of the world (Crustacea: Decapoda: Anomura—families Chirostylidae, Galatheidae and Kiwaidae). Zootaxa 1905, 1–220. doi:10.11646/zootaxa.1905.1.1 Baba K, Ahyong ST, Macpherson E (2011a) Chapter 1. Morphology of marine squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 1–37. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Baba K, Fujita Y, Wehrtmann IS, Scholtz G (2011b) Chapter 4. Developmental biology of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 105–148. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Baeza JA (2011) Chapter 8. Squat lobsters as symbionts and in chemo-autotrophic environments. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 249–270. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Boyko CB, Williams JD (2011) Chapter 9. Parasites and other symbionts of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 271–295. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Bracken-Grissom HD, Cannon ME, Cabezas P, Feldmann RM, Schweitzer CE, et  al. (2013) A comprehensive and integrative reconstruction of evolutionary history for Anomura (Crustacea: Decapoda). BMC Evolutionary Biology 13, 128 doi:10.1186/1471-2148-13-128. Lovrich GA, Thiel M (2011) Chapter 6. Ecology, physiology, feeding and trophic role of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 183–222. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Macpherson E, Baba K (2011) Chapter 2. Taxonomy of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J)

pp. 39–71. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Poore GCB, Ahyong ST, Taylor J (Eds) (2011) The biology of squat lobsters. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Schnabel K (2020) The marine fauna of New Zealand. Squat lobsters (Crustacea, Decapoda, Chirostyloidea). NIWA Biodiversity Memoir 132, 1–351. Schnabel KE, Ahyong ST, Maas EL (2011a) Galatheoidea are not monophyletic  – molecular and morphological phylogeny of the squat lobsters (Decapoda: Anomura) with recognition of a new superfamily. Molecular Phylogenetics and Evolution 58, 157–168. doi:10.1016/j.ympev.2010.11.011 Schnabel KE, Cabezas P, McCallum AW, Macpherson E, Ahyong ST, et al. (2011b) Chapter 5. World-wide distribution patterns of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 149–182. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Thiel M, Lovrich GA (2011) Chapter 7. Agonostic behaviour and reproductive biology of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 223–247. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Tsang LM, Chan T-Y, Ahyong ST, Chu KH (2011) Hermit to king, or hermit to all: multiple transitions to crab-like forms from hermit crab ancestors. Systematic Biology 60, 616–629. doi:10.1093/sysbio/syr063

Chirostylidae Ortmann, 1892 squat lobsters Figure 13.1, Plate 22a–d Chirostylids look superficially much like galatheids but differ in that the fifth legs attach freely and the sternal plate between them is absent. They resemble sternostylids even more (see below). Most species are common in deepwater habitats, usually associated with alcyonacean corals, rarely with antipatharians, pennatulaceans and crinoids, and a few are known from hydrothermal vent sites and cold seeps (Baba and de Saint Laurent 1992; Baba and W ­ illiams 1998; Baeza 2011; Dong and Li 2015; Martin and Haney 2005; Schnabel 2020). Shallow-water species are rare but underwater photographs sometimes show chirostylids associated with alcyonacean corals. They mimic their hosts’ colours (Kawamoto and Okuno 2003; Minemizu 2000). Carapace lengths range from as little as 2 mm to 20 mm.

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

The definitive early work on the systematics of Chirostylidae was by Baba (1988) who recognised five genera and more than 100 species world-wide. This number has now vastly increased to over 300 species, most in the Indo-West Pacific (e.g. Ahyong and Poore 2004; Baba 2018; Baba et al. 2008; McCallum and Poore 2013; Schnabel 2020; Schnabel 2009). The taxonomy of the family was reviewed by Baba (2005), Macpherson and Baba (2011) and by Schnabel (2020) who provided diagnoses and a key to genera. Most genera are currently well circumscribed, except for Hapaloptyx from South Africa, which is known only from the original description (Stebbing 1920).

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Diagnosis. Carapace without transverse setiferous striae; posterolateral margin not distinctly defined or greatly inflated. Supraocular spines absent. Thoracic sternite 3 abruptly demarcated from preceding sternites 1 and 2 by distinct step forming well defined anterior margin at articulation with maxillipeds 3, placing excavated sternum on more dorsal plane (Fig.  13.1i–k). Maxilliped 3 with crista dentata. Implicit generic attributes. Carapace posterolateral margin not distinctly defined or slightly excavated. Mandibular incisor margin serrate. Maxilliped 1 with flagellum.

Key to genera of Chirostylidae 1. – 2. – 3. –

Posterolateral margin of carapace strongly excavated (Fig. 13.1a, c). Base of eyestalks dorsally visible. Maxilliped 1 without flagellum������������������������������������������������������������������������������������������������������������������������������������������������2 Posterolateral margin of carapace barely or weakly excavated (Fig. 13.1b, d, e). Base of eyestalks barely visible in dorsal view. Maxilliped 1 with flagellum���������������������������������������������������������������������������������������������������������������3 Mandible with incisor margin not dentate ��������������������������������������������������������������������������������������������������������������� Hapaloptyx Mandible with incisor margin serrated����������������������������������������������������������������������������������������������������������������������Chirostylus Rostrum spiniform (Fig. 13.1b, d). Anterior margin of sternal plastron concave or sinuous behind excavated sternites 1–3�����������������������������������������������������������������������������������������������������������������������������������������������������������������������4 Rostrum flattish, narrowly or broadly triangular (Fig. 13.1e, f). Anterior margin of sternal plastron bearing distinct cliff behind excavated sternites 1–3���������������������������������������������������������������������������������������������������������������������������5

Fig. 13.1.  Chirostylidae. a, Chirostylus stellaris Osawa, 2007; b, Gastroptychus brevipropodus Baba, 1991; c, Hapaloptyx difficilis Stebbing, 1920; d, Heteroptychus epigaster Baba, 2004; e, Uroptychodes epigaster Baba, 2004; f, Uroptychus naso Van Dam, 1933. Pereopod 2: g, Uroptychus; h, Uroptychodes. Thoracic sternites 3, 4: i, Gastroptychus; j, Uroptychus. Thoracic sternum of female: k, Heteroptychus; l Uroptychus (sternites 3–7 indicated). Telson: m, Gastroptychus; n, Uroptychus.

268

4. – 5. –

Marine Decapod Crustacea

Rostrum flanked by sharp epigastric spines (Fig. 13.1b). Thoracic sternite 3 anterior margin with row of spines. Sternal plastron slightly excavate in both sexes (Fig. 13.1i)��������������������������������������������������������������Gastroptychus Rostrum without epigastric spines (Fig. 13.1d). Thoracic sternite 3 anterior margin without row of spines. Sternal plastron sexually dimorphic, in female strongly excavated, with median parts of sternites 5–7 absorbed into sternite 4 (Fig. 13.1k)������������������������������������������������������������������������������������������������������������������Heteroptychus Pereopod 2 more slender than pereopod 3, dactylus entire on flexor margin (Fig. 13.1h)�����������������������������Uroptychodes Pereopod 2 as broad as pereopod 3, dactylus with spines on flexor margin (Fig. 13.1g)������������������������������������Uroptychus Chirostylus Ortmann, 1892

Diagnosis. Rostrum, if present, small, spiniform, remote from epigastric spines. Carapace dorsal surface generally smooth; posterolateral margin strongly excavated. Thoracic sternite 3 anterior margin straight, transverse or concave (rarely), with row of spines. Eyestalk basal article visible in dorsal view. Maxilliped 1 without flagellum. Maximum cl. 7.3 mm. Subtidal–slope (10–270 m) (associated with soft and black corals). Temperate Northern W Pacific, Western and Central IndoPacific. 7 species (Baba 2009; Osawa 2007).

Gastroptychus Caullery, 1896 Diagnosis. Rostrum spiniform, remote from epigastric spines. Carapace dorsal surface spinose, without striae. Thoracic sternite 3 anterior margin sinuous, anteriorly produced or concave, with row or pair of spines. Maxilliped 1 with flagellum. Maximum cl. 21 mm (Pl. 22b). Shelf, slope (130–768 m). Tropical W Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate South America, Temperate Australasia. 9 species (Baba et al. 2018). Living specimens of Gastroptychus perarmatus (Haig, 1968) were observed living only on colonies of gorgonian corals and sponges in the northern Gulf of California by submarine cameras at 700 m depth (Hendrickx et al. 2013).

Hapaloptyx Stebbing, 1920 Diagnosis. Rostrum a small spine. Carapace dorsal surface smooth; posterolateral margin strongly excavated. Mandibular incisor margin smooth. Maxilliped 1 without flagellum. Maximum cl. 7 mm. Shelf (168 m). Temperate Southern Africa. 1 species (Stebbing 1920).

Heteroptychus Baba, 2018 Diagnosis. Rostrum triangular, flattish. Carapace dorsal surface smooth; pterygostomial flap, posterior half very low in the posterior half. Sternal plastron sexually dimorphic, in female strongly excavated, with median parts of sternites 5–7 absorbed into sternite 4 (left and right parts of sternites 5–7 discontinuous, interrupted by loss of median parts); thoracic sternite 3 anterior margin sinuous, anteriorly produced, without row of spines. Eyestalk basal article visible in dorsal view. Pereopod 2 as broad as pereopod 3. Maximum cl. 5 mm.

Slope–bathyal (331–2084 m). Temperate Northern W Pacific, Tropical W Atlantic, Central Indo-Pacific, Temperate Australasia (Norfolk Ridge). 11 species (Baba 2018: key to species).

Uroptychodes Baba, 2004 Diagnosis. Rostrum triangular, flattish. Carapace dorsal surface granulose or smooth. Thoracic sternite 3 anterior margin concave. Pereopod 2 more slender than pereopod 3; dactylus unarmed on flexor margin. Maximum cl. 12 mm (Pl. 22c). Shelf, slope (68–1110 m). Temperate Northern W Pacific, Central Indo-Pacific, Temperate Australasia (Norfolk Ridge, New Zealand, SE Australia). 14 species (Dong and Li 2010; Dong et al. 2021; Komai 2021: key to species; Schnabel 2020).

Uroptychus Henderson, 1888 Diagnosis. Rostrum triangular, flattish. Carapace dorsal surface granulose, spinose or smooth. Pterygostomial flap, posterior half at most third of anterior height. Sternal plastron slightly excavate in both sexes; thoracic sternite 3 anterior margin concave. Pereopod 2 as broad as pereopod 3; dactylus with spines on flexor margin. Maximum cl. 2–20 mm (Pl. 22a, d). Shelf–bathyal (20–2750 m). Temperate Northern W Pacific, Indo-West Pacific, Temperate Australasia. GOODS bathyal provinces: SE Pacific ridges, New Zealand/Kermadec, Indian. 285 species (Baba 2018: many new species, key to 219 species from the Indo-West and Central Pacific; Schnabel 2020: new species, key to 72 species from New Zealand). Many species are pale pink or orange but colour pattern can be taxonomically important (Poore and Andreakis 2011). References Ahyong ST, Poore GCB (2004) The Chirostylidae of southern Australia (Crustacea, Decapoda, Anomura). Zootaxa 436, 1–88. doi:10.11646/ zootaxa.436.1.1 Baba K (1988) Chirostylid and galatheid crustaceans (Decapoda: Anomura) of the “Albatross” Philippine Expedition, 1907–1910. Researches on Crustacea Special Number 2, 1–203. doi:10.18353/ rcrustaceasn.2.0_1 Baba K (2005) Deep-sea chirostylid and galatheid crustaceans (Decapoda: Anomura) from the Indo-West Pacific, with a list of species. Galathea Report 20, 1–317. Baba K (2009) A new species of squat lobster (Decapoda, Anomura, Chirostylidae) from the Philippines and Indonesia. Crustaceana 82, 795–802. doi:10.1163/156854009X427342

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

Baba K (2018) Chirostylidae of the Western and Central Pacific: Uroptychus and a new genus (Crustacea: Decapoda: Anomura. In: Tropical Deep-Sea Benthos vol. 30. Mémoires du Muséum National d’Histoire Naturelle, Paris 212, 1–612. Baba K, de Saint Laurent M (1992) Chirostylid and galatheid crustaceans (Decapoda: Anomura) from active thermal vent areas in the southwest Pacific. Scientia Marina 56, 321–332. Baba K, Williams AB (1998) New Galatheoidea (Crustacea, Decapoda, Anomura) from hydrothermal systems in the West Pacific Ocean: Bismarck Archipelago and Okinawa Trough. Zoosystema 20, 143–156. Baba K, Macpherson E, Poore GCB, Ahyong ST, Bermudez A, et al. (2008) Catalogue of squat lobsters of the world (Crustacea: Decapoda: Anomura—families Chirostylidae, Galatheidae and Kiwaidae). Zootaxa 1905, 1–220. doi:10.11646/zootaxa.1905.1.1 Baba K, Ahyong ST, Schnabel KE (2018) Rediagnosis of the squat lobster genus Gastroptychus Caullery, 1896, with a new genus Sternostylus and a new family Sternostylidae (Crustacea: Decapoda: Anomura: Chirostyloidea). Zootaxa 4524, 77–86. doi:10.11646/zootaxa.4524.1.5 Baeza JA (2011) Chapter 8. Squat lobsters as symbionts and in chemoautotrophic environments. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 249–270. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Dong C, Li X (2010) A new species and a new record of the genus Uroptychodes Baba, 2004 (Decapoda, Chirostylidae) from the East China Sea. Crustaceana 83(2), 207–213. doi:10.1163/001121609X12591347509167 Dong D, Li X (2015) Galatheid and chirostylid crustaceans (Decapoda: Anomura) from a cold seep environment in the northeastern South China Sea. Zootaxa 4057, 91–105. doi:10.11646/zootaxa.4057.1.5 Dong D, Gan Z, Li X (2021) Descriptions of eleven new species of squat lobsters (Crustacea: Anomura) from seamounts around the Yap and Mariana Trenches with notes on DNA barcodes and phylogeny. Zoological Journal of the Linnean Society 192, 306–355. doi:10.1093/​zoolinnean/ zlab003 Hendrickx ME, Hinojosa A, Ayón-Parente M (2013) In situ observations on the habitat and abundance of the squat lobster Gastroptychus perarmatus (Haig, 1968) (Crustacea: Decapoda: Chirostylidae) in the northern Gulf of California, Mexico. Journal of Threatened Taxa 5, 5228–5236. doi:10.11609/JoTT.o3133.5228-36 Kawamoto T, Okuno J (2003) Shrimps and crabs of Kume Island, Okinawa. 1st edn. Hankyu Communications, Tokyo. Komai T (2021) A new species of the squat lobster genus Uroptychodes Baba, 2004 (Decapoda: Anomura: Chirostylidae) from Japan. 2021 4966, 349–358. Macpherson E, Baba K (2011) Chapter 2. Taxonomy of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 39–71. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Martin JW, Haney TA (2005) Decapod crustaceans from hydrothermal vents and cold seeps: a review through 2005. Zoological Journal of the Linnean Society 145, 445–522. doi:10.1111/j.1096-3642.2005.00178.x McCallum AW, Poore GCB (2013) Chirostylidae of Australia’s western continental margin (Crustacea: Decapoda: Anomura), with the description of five new species. Zootaxa 3664, 149–175. doi:10.11646/zootaxa.3664.2.3 Minemizu R (2000) Marine decapod and stomatopod crustaceans mainly from Japan. Bun-ichi-sogo-shuppan, Tokyo. Osawa M (2007) A new species of Chirostylus Ortmann, 1892 (Crustacea: Decapoda: Anomura: Chirostylidae) from the Ryukyu Islands, southwestrn Japan, with a supplemental description of Chirostylus ortmanni Miyake & Baba, 1968. Zootaxa 1450, 31–43. doi:10.11646/zootaxa.​ 1450.1.3

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Poore GCB, Andreakis N (2011) Morphological, molecular and biogeographic evidence support two new species in the Uroptychus naso complex (Crustacea: Decapoda: Chirostylidae). Molecular Phylogenetics and Evolution 60, 152–169. doi:10.1016/j.ympev.2011.03.032 Schnabel KE (2009) A review of the New Zealand Chirostylidae (Anomura: Galatheoidea) with description of six new species from the Kermadec Islands. Zoological Journal of the Linnean Society 155, 542–582. doi:10.1111/j.1096-3642.2008.00449.x Schnabel K (2020) The marine fauna of New Zealand. Squat lobsters (Crustacea, Decapoda, Chirostyloidea). NIWA Biodiversity Memoir 132, 1–351. Stebbing TRR (1920) South African Crustacea (Part X of S.A. Crustacea, for the Marine Investigations in South Africa). Annals of the South African Museum 17, 231–272.

Eumunididae A. Milne Edwards & Bouvier, 1900 squat lobsters Figure 13.2, Plate 22e Eumunidid squat lobsters differ from chirostylids, sternostylids and kiwaids in having supraocular spines. All are deep-water species (shelf to slope depths) and primarily associated with lace corals (family Stylasteridae) (de Saint Laurent and Macpherson 1990; Schnabel 2020). Their slender rostral and supraocular spines, and transversely grooved carapace gives a superficial resemblance to galatheoid munidids, which are readily separated by chirostyloid superfamily characters, such as the absence of sternite 8 and the telson being weakly divided transversely only, rather than into multiple plates. Diagnosis. Carapace with transverse setiferous striae; posterolateral margin entire, not excavated. Supraocular spines present. Maxilliped 3 with crista dentata.

Fig. 13.2.  Eumunididae. Carapace: a, Eumunida spinosa Macpherson, 2006; b, Pseudomunida fragilis Haig, 1979. c, telson, Eumunida.

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Eumunida Smith, 1883 Diagnosis. Carapace lateral and mesial supraocular spines well developed; oblique row of hepatic spines present (Fig.  13.2a). Maximum cl. 62 mm (Pl. 22e). Shelf, slope (137–1320 m). Temperate Northern W Atlantic, Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific, Temperate Southern Africa, Temperate Australasia (137–1320 m). GOODS bathyal provinces: Northern N Pacific, North Atlantic. 34 species (de Saint Laurent and Poupin 1996; Schnabel 2020). Puillandre et al. (2011) discovered on the basis of molecular data that three species are possible synonyms and that another lineage is taxonomically unrecognised. In their major revision of the genus de Saint Laurent and Poupin (1996) recognised two subgenera, Eumunida s.s. and Eumunidopsis de Saint Laurent and Poupin (1996). While most species are associated with stylasterid lace corals, at least one species has been captured most commonly associated with the deep-sea reef-building scleractinian coral Lophelia pertusa (Caryophylliidae) but also on cold seeps (Kilgour and Shirley 2008).

In Résultats des Campagnes MUSORSTOM, volume 6. (Ed. A Crosnier). Mémoires du Museum National d’Histoire Naturelle, Paris (A) 145, 227–288. de Saint Laurent M, Poupin J (1996) Crustacea, Anomura: Les espèces indo-ouest pacifiques du genre Eumunida Smith, 1880 (Chirostylidae). Description de six espèces nouvelles. In Résultats des Campagnes MUSORSTOM, volume 15. (Ed. A Crosnier) Mémoires du Museum National d’Histoire Naturelle, Paris 168, 337–385. Haig J (1979) A new genus and species of Chirostylidae (Decapoda, Anomura, Galatheidea) from the Hawaiian Islands. Crustaceana 5(Supplement), 89–94. Kilgour MJ, Shirley TC (2008) Eumunida picta S.I. Smith, 1883, and Lophelia pertusa (Linnaeus, 1758): a relationship or just good friends? Crustaceana 81, 587–593. doi:10.1163/156854008784092166 Puillandre N, Macpherson E, Lambourdière J, Cruaud C, BoisselierDubayle M-C, et al. (2011) Barcoding type specimens helps to identify synonyms and an unnamed new species in Eumunida Smith, 1883 (Decapoda: Eumunididae). Invertebrate Systematics 25, 322–333. doi:10.1071/IS11022 Schnabel K (2020) The marine fauna of New Zealand. Squat lobsters (Crustacea, Decapoda, Chirostyloidea). NIWA Biodiversity Memoir 132, 1–351.

Pseudomunida Haig, 1979 Diagnosis. Carapace lateral supraocular spine very small or barely discernible; oblique row of hepatic spines absent (Fig.  13.2b). Maximum cl. 27 mm. Slope (969–1480 m). Central and Eastern Indo-Pacific. 1 species (Haig 1979; Schnabel 2020). References de Saint Laurent M, Macpherson E (1990) Crustacea Decapoda: Le genre Eumunida Smith, 1883 (Chirostylidae) dans les eaux néo-­caledoniennes.

Kiwaidae Macpherson, Jones & Segonzac, 2005 yeti crabs Figure 13.3, Plate 23a–c The discovery of a new family of hairy squat lobsters inhabiting apparently inhospitable hydrothermal vents caused considerable interest (Macpherson et  al. 2005). Not least because they were called ‘yeti crabs’. Since discovery of the

Fig. 13.3.  Kiwaidae. Kiwa hirsuta Macpherson, Jones & Segonzac, 2005: a, b, habitus, dorsal, lateral; c, right antenna; d, thoracic sternites 3, 4.

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

first species three more have been described in other similar environments and from cold seeps (Lee et al. 2016; Thatje et al. 2015; Thurber et al. 2011). Epibiotic bacteria living on the crabs’ thick setae are their main food source which they capture with modified setae on their third maxillipeds (Thurber et al. 2011). All known species of Kiwa live in chemosynthetic habitats, either cold methane seeps or hydrothermal vents, habitats that they probably colonised during the Paleogene (Ahyong and Roterman 2014). Roterman et  al. (2013) reviewed the biogeography of the only genus hypothesised that the three species from hydrothermal vents belong on a clade separate from the only species from a cold seep (Kiwa puravida Thurber, Jones & Schnabel, 2011). Phylogenetic analyses variously identify Eumunididae or Chirostylidae as sister to Kiwaidae, although the best supported analyses suggest eumunidids (Ahyong et al. 2011; Schnabel and Ahyong 2010). Diagnosis. Carapace dorsally smooth, without striae, with grooves clearly demarcating regions; posterolateral margin not greatly inflated. Supraocular spines small. Maxilliped 3 with crista dentata. Kiwa Macpherson, Jones & Segonzac, 2005 Maximum cl. 52 mm (Pl. 23a–c). Slope–bathyal (1007–2600 m). Tropical Eastern Pacific. GOODS bathyal provinces: SE Pacific ridges, Antarctic, Subantarctic. 4 species (Roterman et al. 2013). References Ahyong ST, Roterman CN (2014) Pristinaspinidae, a new family of Cretaceous kiwaiform stem-lineage squat lobster (Anomura, Chirostyloidea). In: Fraaije, R.H.B., Hyžný, M., Jagt, J.W.M., Krobicki, M. & Van Bakel, B.W.M. (eds.), Proceedings of the 5th Symposium on Mesozoic and Cenozoic Decapod Crustaceans, Krakow, Poland, 2013: A tribute to Pál Mihály Müller. Scripta Geologica 147, 125–133. Ahyong ST, Andreakis N, Taylor J (2011) Mitochondrial phylogeny of the deep-sea squat lobsters, Munidopsidae (Galatheoidea). Zoologischer Anzeiger 250, 367–377. doi:10.1016/j.jcz.2011.06.005 Lee SH, Won YJ, Lee WK (2016) A new species of yeti crab, genus Kiwa Macpherson, Jones and Segonzac, 2005 (Decapoda: Anomura: Kiwaidae), from a hydrothermal vent on the Australian-Antarctic Ridge. Journal of Crustacean Biology 36, 238–247. doi:10.1163/1937240X-00002418 Macpherson E, Jones W, Segonzac M (2005) A new squat lobster family of Galatheoidea (Crustacea, Decapoda, Anomura) from the hydrothermal vents of the Pacific-Antarctic Ridge. Zoosystema 27, 709–723. Roterman CN, Copley JT, Linse KT, Tyler PA, Rogers A (2013) The biogeography of the yeti crabs (Kiwaidae) with notes on the phylogeny of the Chirostyloidea (Decapoda: Anomura). Proceedings. Biological Sciences 280, 20130718. doi:10.1098/rspb.2013.0718 Schnabel KE, Ahyong ST (2010) A new classification of the Chirostyloidea (Crustacea: Decapoda: Anomura). Zootaxa 2687, 56–64. doi:10.11646/ zootaxa.2687.1.4 Thatje S, Marsh L, Roterman CN, Mavrogortdato MN, Linse K (2015) Adaptations to hydrothermal vent life in Kiwa tyleri, a new species of yeti crab from the East Scotia Ridge, Antarctica. PLoS One doi:10.1371/ journal.pone.0127621.

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Thurber AR, Jones WJ, Schnabel KE (2011) Dancing for food in the deep sea: bacterial farming by a new species of yeti crab. PLoS One 6, doi:10.1371/journal.pone.0026243.

Sternostylidae Baba, Ahyong & Schnabel, 2018 squat lobsters Figure 13.4, Plates 1b, 23d One genus of 13 species, Sternostylus, differs from Chirostylidae in having thoracic sternite 3 anteromedially produced and evenly sloping anterodorsally with a pair of spines directly behind the anterior margin, maxillipeds 3 adjacent and the dactyli of the pereopods 2–4 ending in an indistinctly demarcated corneous spine (Baba et  al. 2018; Schnabel 2020). Sternostylids were formerly placed in Gastroptychus but morphological and molecular analysis demonstrated they lie well outside of Gastroptychus sensu stricto and form the sister group to Chirostylidae (Baba et al. 2018; Bracken-Grissom et al. 2013). Diagnosis. Carapace without transverse setiferous striae; posterolateral margin not distinctly defined or greatly inflated. Supraocular spines absent (Fig.  13.4a). Thoracic sternite 3 confluent with sternites 1 and 2, without abrupt transverse step at maxilliped 3 articulations, produced and sloping anterodorsally, with pair of spines directly behind anterior margin (Fig. 13.4b). Sternostylus Baba, Ahyong & Schnabel, 2018 Maximum cl. 30 mm (Pls, 1b, 23d). Shelf, slope (229–1786 m). Temperate Northern and Tropical Atlantic, Western Indo-Pacific,

Fig. 13.4.  Sternostylidae. a, carapace: Sternostylus rogeri (Baba, 2000); b, anterior part of sternal plastron, maxillipeds 3, pereopod 1 coxae, Sternostylus.

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Tropical Eastern Pacific, Temperate South America (Pacific coast), Temperate Australasia. 14 species (Dong et  al. 2021; S­chnabel 2020). References Baba K, Ahyong ST, Schnabel KE (2018) Rediagnosis of the squat lobster genus Gastroptychus Caullery, 1896, with a new genus Sternostylus and a new family Sternostylidae (Crustacea: Decapoda: Anomura: Chirostyloidea). Zootaxa 4524, 77–86. doi:10.11646/zootaxa.4524.1.5 Bracken-Grissom HD, Cannon ME, Cabezas P, Feldmann RM, Schweitzer CE, et  al. (2013) A comprehensive and integrative reconstruction of evolutionary history for Anomura (Crustacea: Decapoda). BMC Evolutionary Biology 13, 128. doi:10.1186/1471-2148-13-128 Dong D, Gan Z, Li X (2021) Descriptions of eleven new species of squat lobsters (Crustacea: Anomura) from seamounts around the Yap and Mariana Trenches with notes on DNA barcodes and phylogeny. Zoological Journal of the Linnean Society 192, 306–355. doi:10.1093/​zoolinnean/ zlab003 Schnabel K (2020) The marine fauna of New Zealand. Squat lobsters (Crustacea, Decapoda, Chirostyloidea). NIWA Biodiversity Memoir 132, 1–351.

Galatheoidea Samouelle, 1819 The Galatheoidea comprise four families, some crab-like and others more lobster-like. They share with all anomurans only four pairs of well developed legs of which the first is a stronger cheliped. The fifth leg is weak and often hidden under the carapace. Galatheoids are symmetrical (unlike hermit crabs) and the uropods and telson form a broad tailfan at the end of the depressed pleon which folds up against itself or the thorax. Galatheoids differ from the other superfamily of marine squat lobsters, Chirostyloidea, in having a

well developed sternal plate between the last pereopods and a telson without a transverse suture. This separation is well supported by spermatazoal studies (Tudge 1997), larval data (Clark and Ng 2008) and genetics (Bracken-Grissom et  al. 2009). The relationships between galatheoid families and genera were reviewed by Ahyong et  al. (2010) who introduced a new family, diagnosed all families and listed the genera. Their work was incorporated into a reappraisal of galatheoid relationships by Schnabel et  al. (2011a) using both morphological and molecular data. Their rediagnosis of the superfamily it the basis of the diagnosis here. The biology of squat lobsters in general has been extensively reviewed (Poore et al. 2011). Baba et al. (2011a) dealt with morphology, Macpherson and Baba (2011) with taxonomy, Ahyong et al. (2011) with phylogeny and the fossil record, Baba et  al. (2011b) with developmental biology, ­Schnabel et  al. (2011b) with distribution patterns, Lovrich and Thiel (2011) with ecology, physiology, feeding and trophic role, Thiel and Lovrich (2011) with agnostic behaviour and reproductive biology, Baeza (2011) with life in chemo-autotrophic environments, Boyko and Williams (2011) with parasites and other symbionts, and Wehrtmann and Acuña (2011) with fisheries. Diagnosis. Sternal plastron consisting of thoracic sternites 3–7, thoracic sternite 8 independent. Pleon symmetrical; pleonites separate, calcified and distinct. Ocular scales absent. Antennal peduncle articles 1 and 2 fused. Mandibular incisor margin entire. Pereopods 1 symmetrical, elongate, chelate. Pereopods 2–4 with claw-like dactyli. Pereopod 4 ambulatory, like 2 and 3. Uropods broad, part of tail-fan. Telson divided into median and paired plates.

Key to families of Galatheoidea 1. – 2. – 3. –

Pleon folded up against thorax, body usually short, crab-like, sometimes elongated. Chelipeds moderately elongate, stout or flattened (Fig. 13.10). Maxilliped 3 operculiform, without crista dentata or epipod. Antennal peduncle directed laterally or inclined posteriorly����������������������������������������������������������������������� Porcellanidae Pleon folded up against itself (not against thorax), body lobster-like. Chelipeds greatly elongate, slender (Figs 13.5–13.9). Maxilliped 3 pediform, with crista dentata and epipod. Antennal peduncle directed anteriorly or anterolaterally�������������������������������������������������������������������������������������������������������������������������������������������������������3 Maxilliped 1 exopod without flagellum or flagellum strongly reduced���������������������������������������������������������� Munidopsidae Maxilliped 1 exopod with well developed flagellum (Fig. 13.8k)���������������������������������������������������������������������������������������������4 Front trispinous or trilobate, usually composed of slender rostrum flanked by supraorbital spines (Figs 13.6–13.8)�������������������������������������������������������������������������������������������������������������������������������������������������������������������� Munididae Rostrum broad, triangular (Fig. 13.5)����������������������������������������������������������������������������������������������������������������������� Galatheidae

References Ahyong ST, Baba K, Macpherson E, Poore GCB (2010) A new classification of the Galatheoidea (Crustacea: Decapoda: Anomura). Zootaxa 2676, 57–68. doi:10.11646/zootaxa.2676.1.4 Ahyong ST, Schnabel KE, Macpherson E (2011) Chapter 3. Phylogeny and fossil record of the marine squat lobsters. In The biology of squat

lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 73–104. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Baba K, Ahyong ST, Macpherson E (2011a) Chapter 1. Morphology of marine squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 1–37. CSIRO Publishing, Melbourne

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(also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Baba K, Fujita Y, Wehrtmann IS, Scholtz G (2011b) Chapter 4. Developmental biology of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 105–148. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Baeza JA (2011) Chapter 8. Squat lobsters as symbionts and in chemo-autotrophic environments. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 249–270. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Boyko CB, Williams JD (2011) Chapter 9. Parasites and other symbionts of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 271–295. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Bracken HD, Toon A, Felder DL, Martin JW, Finley M, et  al. (2009) The decapod tree of life: compiling the data and moving toward a consensus of decapod evolution. Arthropod Systematics & Phylogeny 67, 99–116. Clark PF, Ng PKL (2008) Lecithotrophic zoea of Chirostylus ortmanni Miyake and Baba, 1968 (Crustacea: Anomura: Galatheoidea: Chirostylidae) described from laboratory hatched material. Raffles Bulletin of Zoology 56, 85–94. Lovrich GA, Thiel M (2011) Chapter 6. Ecology, physiology, feeding and trophic role of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 183–222. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Macpherson E, Baba K (2011) Chapter 2. Taxonomy of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 39–71. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Poore GCB, Ahyong ST, Taylor J (Eds) (2011) The biology of squat lobsters. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Schnabel KE, Ahyong ST, Maas EL (2011a) Galatheoidea are not monophyletic  – molecular and morphological phylogeny of the squat lobsters (Decapoda: Anomura) with recognition of a new superfamily. Molecular Phylogenetics and Evolution 58, 157–168. doi:10.1016/j. ympev.2010.11.011 Schnabel KE, Cabezas P, McCallum AW, Macpherson E, Ahyong ST, et al. (2011b) Chapter 5. World-wide distribution patterns of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 149–182. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Thiel M, Lovrich GA (2011) Chapter 7. Agonostic behaviour and reproductive biology of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 223–247. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton).

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Tudge CC (1997) Phylogeny of the Anomura (Decapoda, Crustacea): spermatozoa and spermatophore morphological evidence. Contributions to Zoology 67, 125–141. doi:10.1163/​18759866-06702002 Wehrtmann IS, Acuña E (2011) Chapter 10. Squat lobster fisheries. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 297–322. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton).

Galatheidae Samouelle, 1819 squat lobsters Figure 13.5, Plate 24 Galatheids are commonly called squat lobsters although the names, plated lobsters, lobster krill and craylets, have been used. All the species are benthic on rough sedimentary bottoms from the intertidal to much greater depths. Galatheidae once included most squat lobsters but is now restricted to 12 genera, six other families being now recognised (Baba et  al. 2018; Macpherson and Baba 2011). Squat lobsters encountered by SCUBA divers are likely to be true galatheids. Many are associated with corals and are photogenic (Osawa 2015). Some North Atlantic species reach ~50 mm cl. but most species are much smaller, few reaching 5 mm cl. Most species occur in the Indo-West Pacific, and recent studies have revealed considerable diversity, particularly in the genera Galathea and Phylladiorhynchus (Macpherson and Robainas-Barcia 2015; Rodríguez-Flores et al. 2021; Schnabel and Ahyong 2019). Diagnosis. Galatheoidea. Body squat lobster-like. Rostrum triangular. Supraocular spines present, or absent. Carapace with transverse striae or tubercles. Tailfan well developed, not folded under preceding pleonites. Maxilliped 1 exopod flagellum well developed (Fig. 13.5m). Maxilliped 3 pediform; ischium and merus not expanded mesially. Maxilliped 3 with crista dentata; with epipod. Cheliped subcylindrical to ovate in cross-section. Implicit generic attributes. Supraocular spines absent. Antennular basal article with distomesial and distolateral spines. Male pleopod 1 present.

Key to genera of Galatheidae 1. – 2. – 3. – 4.

Lateral margin of rostrum nearly unarmed, at most finely serrate or with rudimentary teeth������������������������������������������2 Lateral margin of rostrum with distinct teeth�����������������������������������������������������������������������������������������������������������������������������6 Rostrum leaflet-like, margin usually smooth or finely serrate (Fig. 13.5j)�����������������������������������������������������������������������������3 Rostral margin with 2–5 very small or rudimentary teeth (Fig. 13.5e)�����������������������������������������������������������������������������������5 Carapace supraocular spines absent (Fig. 13.5j)��������������������������������������������������������������������������������������������������� Nanogalathea Carapace with distinct supraocular spine on each side of rostrum (Fig. 13.5a, k)����������������������������������������������������������������4 Carapace without outer orbital spine; epigastric spines absent (Fig. 13.5a). Male pleopods 1 and 2 present��������� Alainius

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– 5. – 6. – 7.

Marine Decapod Crustacea

Carapace with outer orbital spine; epigastric spines usually present (Fig. 13.5k). Male pleopods 1 only����������������������������������������������������������������������������������������������������������������������������������� Phylladiorhynchus Carapace lacking setiferous striae. Eyestalks narrow and elongate (Fig. 13.5e)������������������������������������������Fennerogalathea Carapace bearing setiferous striae. Eyestalks relatively broad and short (Fig. 13.5c)����������������������������������������Allomunida Uropodal endopod extremely broad���������������������������������������������������������������������������������������������������������������������������������������������7 Uropodal endopod about as broad as long�����������������������������������������������������������������������������������������������������������������������������������8 Maxilliped 3 carpus flexor margin with spine. Pereopods with plumose setae restricted to mesial and lateral margins of cheliped and extensor and flexor margins of pereopods 2–4. Pereopods 2–4 dactyli bearing 1 tooth on flexor margin (Fig. 13.5p)��������������������������������������������������������������������Lauriea

Fig. 13.5.  Galatheidae. a, Alainius crosnieri Baba, 1991; b, Allogalathea elegans (Adams & White, 1848); c, Allomunida magnicheles Baba, 1988; d, Coralliogalathea humilis Nobili, 1905; e, Fennerogalathea chacei Baba, 1988; f, Galathea consobrina De Man, 1902; g, Janetogalathea californiensis (Benedict, 1902); h, Lauriea siagiani Baba, 1994; i, Macrothea Macpherson & Cleva, 2010; j, Nanogalathea raymondi Tirmizi & Javed, 1980; k, Phylladiorhynchus pusillus (Henderson, 1885); l, Triodonthea setosa Macpherson & Robainas-Barcia (2013). Maxilliped 1: m, Galathea. Pereopod 2: n, Triodonthea; o, Galathea; p, Lauriea; q, Macrothea. Telson: r, Galathea.

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– 8. – 9. – 10. – 11. –

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Maxilliped 3 carpus flexor margin unarmed. Pereopods 1–4 covered with numerous long and plumose setae. Pereopods 2–4 dactyli bearing 2 teeth on flexor margin (Fig. 13.5n)���������������������������������� Triodonthea Carapace with orbit not excavated. Antennal peduncle article 1 fused with body (Fig. 13.5g)�������������������Janetogalathea Carapace with orbit excavated. Antennal peduncle article 1 free��������������������������������������������������������������������������������������������9 Rostrum extremely elongate, ventrally carinate, with 5–9 lateral spines (Fig. 13.5b)�������������������������������������� Allogalathea Rostrum moderate in length, usually flattish, with 2–5 lateral teeth������������������������������������������������������������������������������������10 Rostrum with 3 (rarely 4) distinct teeth on each side (Fig. 13.5d). Thoracic sternite 3 strongly produced, anteriorly pointed, as long as or longer than wide������������������������������������������������Coralliogalathea Rostrum with 2–5 (usually 4 or 5) distinct teeth on each side (Fig. 13.5f, i). Thoracic sternite 3 anteriorly blunt, variously shaped wider than long���������������������������������������������������������������������������11 Pereopods 2–4 dactyli sharply biunguiculate (Fig. 13.5q)����������������������������������������������������������������������������������������Macrothea Pereopods 2–4 dactyli flexor margin with row of seta-like movable spines (Fig. 13.5o)����������������������������������������Galathea Alainius Baba, 1991

Diagnosis. Rostrum without lateral teeth, with obvious supraocular spine on each side. Carapace epigastric spines absent; orbit without outer orbital spine, concave, not ventrally delimited by denticulate crest. Antennular basal article with distomesial and distolateral spines. Maxilliped 3 merus, submedian process on flexor margin strong spine-like. Male pleopods 1 and 2 present. Maximum cl. 4.3 mm. Shelf, slope (90–600 m). Eastern Indo-Pacific (New Caledonia). 1 species (Baba 1991).

Allogalathea Baba, 1969 Diagnosis. Rostrum with ventral carina, with 5–9 lateral teeth. Carapace orbital margin deeply concave. Antennular basal article with 3 terminal spines. Maxilliped 3 merus, submedian process on flexor margin strong spine-like or dentate. Maximum cl. 14 mm (Pl. 24a). Intertidal, shelf (0–94 m) (closely associated with crinoids). Temperate Northern W Pacific, Western and Central IndoPacific, Temperate Australia. 4 species. (Cabezas et  al. 2011: molecular, colour and morphological differences, key to species)

Allomunida Baba, 1988 Diagnosis. Rostrum unarmed or with 3 rudimentary lateral teeth in proximal half. Carapace orbital margin slightly concave, outer orbital spine present. Antennular basal article with distomesial and distolateral spines. Maxilliped 3 merus, submedian process on flexor margin obsolete or smooth. Maximum cl. 5.5 mm. Shelf–slope (22–472 m). Central Indo-Pacific. 1 species (Baba 1988).

Coralliogalathea Baba & Javed, 1974 Diagnosis. Rostrum with 3 (rarely 4) distinct lateral teeth. Carapace without supraocular spines; orbital margin deeply concave, with outer orbital spine. Thoracic sternite 3 strongly produced anteriorly pointed, triangular, chordate, as long as or longer than wide. Antennular basal article with 3 terminal spines. Maxilliped

3 merus, submedian process on flexor margin dentate. Male pleopod 1 absent. Maximum cl. 3 mm. Intertidal–shelf (0–100 m) (associated with corals). Indo-West Pacific. 6 species (Rodríguez-Flores et al. 2019).

Fennerogalathea Baba, 1988 Diagnosis. Rostrum with 2–5 distinct lateral teeth, with 2 pairs of epigastric spines. Carapace orbital margin oblique, with outer orbital spine; dorsal surface lacking setiferous striae. Antennular basal article with well developed distolateral and distodorsal spines, small distomesial spine. Maxilliped 3 merus, submedian process on flexor margin strong spine-like. Maximum cl. 8.4 mm. Shelf–bathyal (102–2417 m). Western and Central Indo-Pacific. GOODS bathyal province: Indian. 5 species (Rodríguez-Flores et al. 2017).

Galathea Fabricius, 1793 Diagnosis. Rostrum with 2–5 distinct lateral teeth. Carapace orbital margin deeply concave, outer orbital spine present, orbit ventrally delimited by denticulate crest. Thoracic sternite 3 anteriorly blunt, variously shaped wider than long. Antennular basal article with distomesial and distolateral spines. Maxilliped 3 merus, submedian process on flexor margin strong spine-like. Maximum cl. 5–15 mm (Pl. 24d–h). Intertidal–slope (0–1350 m). Cosmopolitan except polar seas. 177 species. Most species are in shallow waters (< 100 m) but a few reach 500 m or more, mostly in reef and coral environments. Macpherson and Robainas-Barcia (2015) revised Galathea and described 91 new species based on genetic and subtle morphological differences. They provided a key to 160 species from the Indian and Pacific Oceans but others occur elsewhere.

Janetogalathea Baba & Wicksten, 1997 Diagnosis. Rostrum with 3–4 distinct lateral teeth. Carapace frontal margin transverse or oblique; orbital margin weakly concave,

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Marine Decapod Crustacea

outer orbital spine absent. Antennular basal article with 3 well developed spines. Maxilliped 3 merus, submedian process on flexor margin dentate. Maximum cl. 36 mm. Shelf, slope (89–1015 m). Tropical Eastern Pacific. 1 species (Baba and Wicksten 1997).

submedian process on flexor margin dentate. Uropodal endopod wider than long. Maximum cl. 5 mm. Shelf (80–223 m). Central Indo-Pacific. 1 species (Macpherson and Robainas-Barcia 2013). References

Lauriea Baba, 1971 Diagnosis. Rostrum with 2–5 distinct lateral teeth. Carapace orbital margin slightly concave, without outer orbital spine. Antennular basal article with 3 terminal spines (middle on smallest). Maxilliped 3 merus, submedian process on flexor margin dentate. Uropodal endopod wider than long. Maximum cl. 4.5 mm (Pl. 24b). Intertidal–slope (0–355 m). Temperate Northern W Pacific, Indo-West Pacific, Temperate Australia. 7 species. (Macpherson and Robainas-Barcia 2013: key to species)

Macrothea Macpherson & Cleva, 2010 Diagnosis. Rostrum with 4 distinct lateral teeth. Carapace orbital margin oblique, weakly concave, with outer orbital angle. Thoracic sternite 3 subquadrate, blunt, wider than long Antennular basal article with 3 terminal spines. Maxilliped 3 merus, submedian process on flexor margin dentate. Uropodal endopod longer than broad. Maximum cl. 5 mm (Pl. 24c). Subtidal, shelf (15–77 m). Central Indo-Pacific. 1 species (Macpherson and Cleva 2010).

Nanogalathea Tirzimi & Javed, 1980 Diagnosis. Rostrum without lateral teeth. Carapace orbital margin deeply concave, outer orbital spine absent. Antennular basal article with 4 terminal spines. Maxilliped 3 merus, submedian process on flexor margin strong spine-like. Maximum cl. 2 mm. Shelf (77 m). Western Indo-Pacific. 1 species (Tirmizi and Javed 1980).

Phylladiorhynchus Baba, 1969 Diagnosis. Rostrum without lateral teeth, usually with tiny subterminal teeth, distinct supraocular tooth on each side. Carapace orbital margin deeply concave, outer orbital spinne present; epigastric spines usually present. Antennular basal article with 5 terminal spines. Maxilliped 3 merus, submedian process on flexor margin dentate. Male pleopod 1 absent. Maximum cl. 4 mm (Pl. 24i). Intertidal–slope (0–1246 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia. 55 species (Macpherson 2008; Schnabel and Ahyong 2019; Rodríguez-Flores et al. 2021: rediagnosis, key to species).

Triodonthea Macpherson & Robainas-Barcia, 2013 Diagnosis. Rostrum with 4 distinct lateral teeth. Carapace orbital margin slightly concave, with outer orbital spine. Antennular basal article with 3 terminal spines. Maxilliped 3 merus,

Baba K (1988) Chirostylid and galatheid crustaceans (Decapoda: Anomura) of the “Albatross” Philippine Expedition, 1907–1910. Researches on Crustacea Special Number 2, 1–203. doi:10.18353/rcrustaceasn.2.0_1 Baba K (1991) Crustacea Decapoda: Alainius gen. nov., Leiogalathea Baba, 1969, and Phylladiorhynchus Baba, 1969 (Galatheidae) from New Caledonia. In Résultats des Campagnes MUSORSTOM, volume 9. (Ed. A Crosnier). Mémoires du Muséum National d’Histoire Naturelle, Paris (A) 152, 479–491. Baba K, Wicksten MK (1997) Janetogalathea, a new genus of squat lobster, with redescription of its type species Galathea californiensis Benedict, 1902 (Anomura: Galatheidae). Crustacean Research 26, 38–46. doi:10.18353/crustacea.26.0_38 Baba K, Ahyong ST, Schnabel KE (2018) Rediagnosis of the squat lobster genus Gastroptychus Caullery, 1896, with a new genus Sternostylus and a new family Sternostylidae (Crustacea: Decapoda: Anomura: Chirostyloidea). Zootaxa 4524, 77–86. doi:10.11646/zootaxa.4524.1.5 Cabezas P, Macpherson E, Machordom A (2011) Allogalathea (Decapoda: Galatheidae): a monospecific genus of squat lobsters? Zoological Journal of the Linnean Society 162, 245–270. doi:10.1111/j.1096-3642.2010.00681.x Macpherson E (2008) Some new records of shallow-water galatheid crustaceans (Anomura: Galatheidae) from the Dampier Archipelago, Western Australia. Records of the Western Australian Museum 72(Supplement), 289–297. Macpherson E, Baba K (2011) Chapter 2. Taxonomy of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 39–71. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Macpherson E, Cleva R (2010) Shallow-water squat lobsters (Crustacea, Decapoda, Galatheidae) from Mayotte (Comoros Island), La Réunion and Madagascar, with the description of a new genus and two new species. Zootaxa 2612, 57–68. doi:10.11646/zootaxa.2612.1.4 Macpherson E, Robainas-Barcia A (2013) A new genus and some new species of the genus Lauriea Baba, 1971 (Crustacea, Decapoda, Galatheidae) from the Pacific and Indian Oceans, using molecular and morphological characters. Zootaxa 3599, 136–160. doi:10.11646/ zootaxa.3599.2.2 Macpherson E, Robainas-Barcia A (2015) Species of the genus Galathea Fabricius, 1793 (Crustacea, Decapoda, Galatheidae) from the Indian and Pacific Oceans, with descriptions of 92 new species. Zootaxa 3913, 1–335. doi:10.11646/zootaxa.3913.1.1 Osawa M (2015) A new strikingly-colored species of the genus Galathea Fabricius, 1793 (Crustacea: Decapoda: Anomura: Galatheidae) from the Ryukyu Islands, Japan. Zootaxa 4039, 591–598. doi:10.11646/ zootaxa.4039.4.9 Rodríguez-Flores PC, Machordom A, Macpherson E (2017) Three new species of squat lobsters of the genus Fennerogalathea Baba, 1988 (Decapoda: Galatheidae) from the Pacific Ocean. Zootaxa 4276, 46–60. doi:10.11646/zootaxa.4276.1.2 Rodríguez-Flores PC, Macpherson E, Buckley D, Machordom A (2019) High morphological similarity coupled with high genetic differentiation in new sympatric species of coral-reef squat lobsters (Crustacea: Decapoda: Galatheidae). Zoological Journal of the Linnean Society 185, 984–1017. doi:10.1093/zoolinnean/zly074 Rodríguez-Flores PC, Macpherson E, Machordom A (2021) Revision of the squat lobsters of the genus Phylladiorhynchus Baba, 1969 (Crustacea,

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Decapoda, Galatheidae) with the description of 41 new species. Zootaxa 5008, 1–159. doi:10.11646/zootaxa.5008.1.1 Schnabel KE, Ahyong ST (2019) The squat lobster genus Phylladiorhynchus Baba, 1969 in New Zealand and eastern Australia, with description of six new species. Zootaxa 4688, 301–347. doi:10.11646/zootaxa.4688.3.1 Tirmizi NM, Javed W (1980) Nanogalathea raymondi, a new genus and species of Galatheidae (Decapoda, Anomura) from the Bay of Bengal. Crustaceana 38, 127–130. doi:10.1163/156854080X00553

Munididae Ahyong, Baba, Macpherson & Poore, 2010 squat lobsters Figure 13.6–13.8, Plate 25 Munidid squat lobsters typically have a spiniform rostrum with one, sometimes two, supraocular spines on each side. In some genera these are reduced to small tubercles (Ahyong et al. 2010). Munididae are the largest family of squat lobsters, the genus Munida being especially diverse. The family ranges

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from intertidal to bathyal environments but most species concentrate on the continental shelf or slope. Phylogenetic relationships between the genera were investigated by Machordom and Macpherson (2004) and Macpherson and Machordom (2005), and summarised by Ahyong et al. (2011). The key to genera is expanded from that in Macpherson and Baba (2011). Keys to most genera from the Indo-West Pacific can be found in Baba (2005) but these have largely been superseded. Diagnosis. Galatheoidea. Body squat lobster-like. Rostrum spiniform (rarely reduced to tubercle). Carapace supraocular spines present; dorsal surface with transverse striae. Tailfan well developed, folded under preceding pleonites. Maxilliped 1 exopod flagellum well developed (Fig. 13.8k). Maxilliped 3 pediform; ischium and merus not expanded mesially. Maxilliped 3 with crista dentata; with epipod. Cheliped subcylindrical to ovate in cross-section. Implicit generic attributes. Supraocular spines absent. Carapace dorsal surface with long, continuous striae.

Key to genera of Munididae 1. –

Carapace mesogastric region without laterally compressed, anteriorly bluff process�����������������������������������������������������������2 Carapace mesogastric region with laterally compressed anteriorly bluff process or abnormally developed gastric spine (Fig. 13.7f–j)���������������������������������������������������������������������������������������������������������������������������������������������������������17 2. Carapace without any dorsal spine (Fig. 13.6b)��������������������������������������������������������������������������������������������������������Anoplonida – Carapace with dorsal spines�����������������������������������������������������������������������������������������������������������������������������������������������������������3 3. Rostrum with dorsal and ventral spines (Fig. 13.7k)�������������������������������������������������������������������������������������������� Cervimunida – Rostrum lacking dorsal and ventral spines����������������������������������������������������������������������������������������������������������������������������������4 4. Rostral spine flanked by 2 lateral spines on each side (Fig. 13.7b)�������������������������������������������������������������������������Sadayoshia – Rostral spine flanked by 1 spine (supraocular) on each side (Fig. 13.7c)���������������������������������������������������������������������������������5 5. Carapace pterygostomial flap largely visible in dorsal view (Fig. 13.6o)������������������������������������������������������������Pleuroncodes – Carapace pterygostomial flap not visible in dorsal view�����������������������������������������������������������������������������������������������������������6 6. Epipods present on pereopods 1–3. Maxilliped 3 carpus with spine at flexor distal margin (Fig. 13.8n)������� Raymunida – Epipods absent from pereopods 1–3. Maxilliped 3 carpus unarmed on flexor margin (Fig. 13.8l, m)������������������������������7 7. Pereopods 2–4 meri unarmed dorsally, dactyli smooth on flexor margin (Fig. 13.8o). Carapace with outer orbital spine (Fig. 13.8a)�������������������������������������������������������������������������������������������������������������������������������� Anomoeomunida – Pereopods 2–4 meri armed with dorsal spines, dactyli with spine-like setae on flexor margin (Fig. 13.8p). Carapace with outer orbital spine a low angle or absent (Fig. 13.8b)����������������������������������������������������������������������������������8 8. Antennal peduncle slender, articles 1 and 2 spineless (Fig. 13.8f)����������������������������������������������������������������������������� Enriquea – Antennal peduncle broad, articles 1 and 2 each with distomesial spine (Fig. 13.8i)�������������������������������������������������������������9 9. Orbit with mound mesial to its lateral limit (Fig. 13.7e)������������������������������������������������������������������������������������������� Torbenella – Orbit without mound mesial to its lateral limit�������������������������������������������������������������������������������������������������������������������������10 10. Carapace orbital margin deeply concave (Fig. 13.6f). Maxilliped 3 merus subrhomboidal (Fig. 13.8l)�����������Crosnierita – Carapace orbital margin weakly concave, transverse or oblique. Maxilliped 3 merus elongate (Fig. 13.8m)�����������������11 11. Carapace without distinct transverse ridges and striae (Fig. 13.6h, m, n)��������������������������������������������������������������������������� 12 – Carapace with distinct transverse ridges or striae (Figs 13.6a–g, j–l, o, 13.7c, d, f, h–j, l)�������������������������������������������������������� 14 12. Antennal article 1 distomesial spine never exceeding distal spines of antennular article 1 (Fig. 13.8i)���������������������������13

278

Marine Decapod Crustacea

Fig. 13.6.  Munididae. Dorsal view: a, Agononida aequabilis Macpherson, 2006; b, Anoplonida cracentis Baba and de Saint Laurent, 1996; c, Babamunida callista (Macpherson, 1994); d, Bathymunida longipes Van Dam, 1938; e, Cervimunida princeps Benedict, 1902; f, Crosnierita dicata Macpherson, 1998; g, Enriquea leviantennata (Baba, 1988); h, Hendersonida granulata (Henderson, 1885); i, Heteronida aspinirostris (Khodkina, 1981); j, Munida aequalis Ahyong and Poore, 2004; k, Neonida grandis Baba & de Saint Laurent, 1996; l, Onconida modica Baba and de Saint Laurent, 1996; m, Paramunida antipodes Ahyong and Poore, 2004; n, Plesionida psila Baba & de Saint Laurent, 1996; o, Pleuroncodes monodon (H. Milne Edwards, 1837).

– 13. – 14. – 15. – 16. –

Antennal article 1 distomesial spine long, clearly exceeding distal spines of antennular article 1 (Fig. 13.8g, h)������������� 15 Pleonites 2–3 each with strong median hump-like process (Figs 13.6i, 13.7g)�����������������������������������������������������Heteronida Pleonites 2–4 without hump-like process, with paired spines (Fig. 13.6n)������������������������������������������������������������ Plesionida Carapace dorsal surface covered with spinules (Fig. 13.6m). Antennal peduncle article 2 distomesial spine never reaching end of anterior prolongation of article 1 (Fig. 13.8h)��������������������������������������������������������������Paramunida Carapace dorsal surface granulose (Fig. 13.6h). Antennal peduncle article 2 distomesial spine long, almost reaching end of anterior prolongation of article 1 (Fig. 13.8g)���������������������������������������������������������������������Hendersonida Lateral parts of thoracic sternite 7 each with 1 or 2 seta-like spines (Fig. 13.8r)������������������������������������������������������ Setanida Lateral parts of thoracic sternite 7 without seta-like spines����������������������������������������������������������������������������������������������������16 Pleonite 4 usually with more than 2 spines on anterior ridge (Fig. 13.6a) (except A. tenuipes). Gonopod 1 absent�������������������������������������������������������������������������������������������������������������������������������������������������������������������������� Agononida Pleonite 4 usually unarmed (Fig. 13.6c, j) (except M. babai and M. gilii). Gonopods 1 and 2 present�����������������������������17

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

279

17.

Epistome with ridge arising from marginal ridge of mouth, laterally leading to ventral margin of orbit distinctly anterior to base of antennal peduncle (Fig. 13.8c)���������������������������������������������������������������������������Babamunida – Epistome with ridge arising from marginal ridge of mouth, laterally leading to level of antennal gland aperture (Fig. 13.8d)��������������������������������������������������������������������������������������������������������������������������������������������������������Munida 18. Antennal basal article 1 distomesial spine long overreaching remaining articles���������������������������������������������������������������19 – Antennal basal article 1 distomesial spine short, not overreaching article 2��������������������������������������������������������������������� 20 19. Pleonite 2 and usually pleonite 3 unarmed (Fig. 13.7i)����������������������������������������������������������������������������������������������� Onconida – Pleonites 2 and 3 spinose (Fig. 13.7d)��������������������������������������������������������������������������������������������������������������������������Tasmanida 20. Antennule basal article with 2 spines, one above the other, at distolateral angle (Fig. 13.8j)���������������������������������Neonida – Antennule basal article with single spine at distolateral angle���������������������������������������������������������������������������Bathymunida

Fig. 13.7.  Munididae. Dorsal view: a, Raymunida formosanus Lin, Chan & Chu, 2004; b, Sadayoshia edwardsii (Miers, 1884); c, Setanida cristata (Macpherson, 2006); d, Tasmanida norfolkae Ahyong, 2007; e, Torbenella orbis (Baba, 2005). Lateral view: f, Bathymunida dissimilis Baba & de Saint Laurent, 1996; g, Heteronida aspinirostris (Khodkina, 1981); h, Neonida grandis Baba & de Saint Laurent, 1996; i, Onconida alaini Baba & de Saint Laurent, 1996; j, Tasmanida norfolkae Ahyong, 2007; k, Cervimunida. l, thoracic sternum, Munida (sternites 3–8 numbered).

Fig. 13.8.  Munididae. Front of carapace: a, Anomoeomunida; b, Enriquea. Epistome, antennules, eyes: c, Babamunida; d, Munida. Antennal and antennular peduncles, ventral: e, Crosnierita; f, Enriquea; g, Hendersonida; h, Paramunida; i, Plesionida. Antennular peduncle article 1, lateral: j, Neonida. Maxilliped 1: k, Munida. Maxilliped 3, ischium–carpus (dactylus): l, Crosnierita; m, Plesionida; n, Raymunida. Pereopod 2: o, Anomoeomunida; p, Enriquea. Pereopod 5, propodus, dactylus: q, Onconida. Thoracic sternum: r, Setanida.

280

Marine Decapod Crustacea

Agononida Baba & de Saint Laurent, 1996

Bathymunida Balss, 1914

Diagnosis. Rostrum spiniform, remote from supraocular spines. Carapace with ventral orbital margin not visible in dorsal view; with pair of epigastric spines; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 each with 2 elevated transverse ridges, each anterior ridge often armed with spines. Male pleopod 1 absent. Maximum cl. 35 mm (Pl. 25d). Shelf, slope (40–1232 m). Tropical W Atlantic, Temperate Northern W Pacific, Indo-West Pacific, Temperate Australasia. 41 species (Baba and de Saint Laurent 1996; Macpherson et al. 2020: records of several species). Some species of Agononida of the IndoWest Pacific present a classic case of cryptic species revealed by molecular and thorough morphological examination. The apparently widespread species Agononida incerta (Henderson, 1888) was shown to comprise ten species with limited but overlapping ranges (Poore and Andreakis 2012, 2014).

Diagnosis. Rostrum base broad, rostral and supraocular spines widely separated; supraocular spines obvious or obsolete. Carapace with transverse striae usually interrupted, often reduced, with pair of small epigastric spines, keel-like process on gastric and cardiac regions; mesogastric region with prominent median ridge-like process or elevated region. Pleonites 2–4 each having anterior ridge with pair of submedian spines. Antennular basal article with distolateral spine (longer than distomesial). Male pleopod 1 absent. Maximum cl. 6 mm (Pl. 25f). Shelf, slope (85–610 m). Temperate Northern W Pacific, IndoWest Pacific. 15 species (Baba and de Saint Laurent 1996: key to 13 species).

Anomoeomunida Baba, 1993 Diagnosis. Rostrum spiniform, dorsally with low rounded ridge. Carapace with some small epigastric, hepatic and parahepatic spines; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 unarmed. Antennular basal article with 3 terminal spines (small). Pereopods 2–4 meri unarmed dorsally; dactyli flexor margin smooth, unarmed. Male pleopod 1 present. Maximum cl. 5.5 mm. Subtidal, shelf (11–38 m). Tropical W Atlantic. 1 species (Baba 1993).

Anoplonida Baba & de Saint Laurent, 1996 Diagnosis. Rostrum stout, laterally ridged, broad at base. Carapacewith 1 pair of obsolete supraocular spines; without gastric spines; mesogastric region with prominent median ridge-like process or elevated region. Pleonites 2 and 3 each with 2 pairs of spines. Antennular basal article with 2 distolateral spines. Male pleopod 1 absent. Maximum cl. 5 mm. Shelf, slope (135–424 m). Central Indo-Pacific. 3 species (Baba and de Saint Laurent 1996; Macpherson and Baba 2006).

Babamunida Cabezas, Macpherson & Machordom, 2008 Diagnosis. Rostrum spiniform, carinate dorsally, clearly overreaching supraocular spines; supraocular spines carinate dorsally, not exceeding end of corneas; deep longitudinal grooves between rostrum and supraocular spines. Carapace ventral orbital margin with well developed mesial spine; with pair of epigastric spines directly behind ocular peduncles usually flanked by spines; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 unarmed. Antennular basal article with 2 distolateral spines. Male pleopod 1 present. Maximum cl. 19 mm. Shelf, slope (37–590 m) (associated with corals and sponges). Indo-West Pacific. 8 species (Baba 2011: key to 7 species; Cabezas et  al. 2008: 5 species on genetic and morphological evidence; Macpherson et al. 2020).

Cervimunida Benedict, 1902 yellow squat lobster Diagnosis. Rostrum spiniform, flanked by supraocular spine, arched in lateral view, with dorsal and ventral teeth. Carapace with pair of epigastric spines directly behind ocular peduncles usually flanked by spines; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 each having anterior ridge with pair of submedian spines. Antennular basal article with 2 lateral spines. Male pleopod 1 present. Maximum cl. 39 mm (Pl. 25a). Intertidal–slope (0–500 m). Temperate Northern Pacific, Central Indo-Pacific, Temperate South America (Pacific coast). 2 species. Cervimunida johni is the target of a commercial bottom-trawl fishery in Chile and has been well studied for this reason (Thiel and Lovrich 2011; Wehrtmann and Acuña 2011).

Crosnierita Macpherson, 1998 Diagnosis. Rostrum spiniform, overreaching supraocular spines. Carapace ventral orbital margin not visible in dorsal view; with transverse striae granulated; with pair of epigastric and postcervical spines, sometimes with median gastric and cardiac spines; mesogastric region with prominent median ridge-like process or elevated region. Pleonites 2–4 each having 2 transverse ridges, each anterior ridge with 4–6 spines. Antennular basal article with 2 lateral spines. Male pleopod 1 absent. Maximum cl. 9 mm. Shelf, slope (80–610 m). Central Indo-Pacific. 6 species (Ahyong et al. 2013: key to 5 species; Macpherson et al. 2020).

Enriquea Baba, 2005 Diagnosis. Rostrum spiniform, overreaching supraocular spines. Carapace dorsal surface granulose and striated, with small dorsal spines; prominent anterolateral spines directed forward, reaching midlength of ocular peduncle; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 each having 2 transverse ridges, both with spines. Antennular basal article with 1 small lateral spine. Antennal peduncle article 2 distomesial spine absent. Male pleopod 1 present. Maximum cl. 20 mm. Slope (300–1250 m). Central Indo-Pacific. 1 species (Baba 2005).

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

Hendersonida Cabezas & Macpherson, 2014 Diagnosis. Rostrum short, basally subtriangular, distally ending in spine; supraocular spines short, stout. Carapace dorsal surface granulose, spinulose, without striations; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 each having 2 transverse ridges, both with spines. Antennular basal article with 2 obsolete lateral spines. Antennal peduncle article 2 distomesial spine long, almost reaching end of anterior prolongation of article 1. Male pleopod 1 absent. Maximum cl. 14 mm. Slope (395–650 m). Central Indo-Pacific. 1 species (Cabezas and Macpherson 2014).

Heteronida Baba & de Saint Laurent, 1996 Diagnosis. Rostrum base broad, rostral spine rudimentary, supraocular spines blunt. Carapace dorsal surface granulose, without transverse striations with laterally compressed, anteriorly bluff strong process, epigastric spines obsolescent; mesogastric region with prominent median ridge-like process or elevated region. Pleonites 2 and 3 each with strong median process flanked by low lateral process. Antennular basal article with 1 small lateral spine. Male pleopod 1 absent. Maximum cl. 4.2 mm. Slope (205–930 m). Central and Eastern Indo-Pacific. 3 species (Macpherson and Baba 2006: differentiation of species).

Munida Leach, 1820 Diagnosis. Rostrum spiniform; shallow longitudinal grooves between rostral and supraocular spines. Carapace with pair of epigastric spines directly behind ocular peduncles usually flanked by additional spines; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 with setiferous transverse striae. Antennular basal article with 2 lateral spines. Male pleopod 1 present. Maximum cl. 30 mm (Pl. 25c). Intertidal–bathyal (0–2165 m). Cosmopolitan except Arctic. 306 species. Munida is the largest genus of squat lobsters (Komai 2014). Baba (2005) provided keys to 160 species from the IndoWest Pacific and Southern Ocean and 16 species from the eastern Pacific but some of these are now in other genera. The most recent substantial works described 16 new species from Japan (Komai 2011, 2012) and eight new species each from the western Indian Ocean (Macpherson et al. 2017) and Papua New Guinea (Macpherson et  al. 2020). One species, Munida gregaria (Fabricius, 1793), sometimes called ‘lobster-krill’, occurs off southern South America and New Zealand where it swarms in such large numbers that it is responsible for ‘red tides’ (Zeldis 1985). Munida gregaria was said to be a pelagic form, while a separate but related morphological species, M. subrugosa Dana, 1852, was associated with the benthic phase. The two have now been shown to belong to one species (Pérez-Barros et al. 2008).

Neonida Baba & de Saint Laurent, 1996 Diagnosis. Rostrum stout, flanked by small supraocular spines. Carapace with prominent gastric and cardiac processes, without

281

epigastric spines; mesogastric region with prominent median ridge-like process or elevated region. Pleonites 2–4 strigose, with 2 transverse ridges, each anterior ridge elevated, with 2 strong submedian spines. Antennular basal article with 2 distolateral spines, one above other. Male pleopod 1 absent. Maximum cl. 8 mm. Slope (273–500 m). Central Indo-Pacific. 1 species (Baba and de Saint Laurent 1996).

Onconida Baba & de Saint Laurent, 1996 Diagnosis. Rostrum stout, flanked by small supraocular spines. Carapace with median gastric process, small epigastric spines, small postcervical spines; mesogastric region with prominent median ridge-like process or elevated region. Pleonites 2 and 3 unarmed, pleonite 3 sometimes with spine; pleonite 4 anterior margin with 2 spines. Cheliped with red spot on distal portion of merus, or juncture between merus and carpus. Male pleopod 1 absent. Maximum cl. 7 mm. Shelf–slope (125–800 m). Western and Central Indo-Pacific. 6 species (Ahyong et al. 2013: key to species)

Paramunida Baba, 1988 Diagnosis. Rostrum short, basally subtriangular, distally ending in spine; supraocular spines short, stout. Carapace dorsal surface covered with spinules, with obsolescent, short or scale-like striae, spinulose; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 each having 2 transverse ridges, both with spines. Antennular basal article with 2 obsolete lateral spines. Antennal peduncle article 2 distomesial spine never reaching end of anterior prolongation of article 1. Male pleopod 1 absent. Maximum cl. 14 mm. Shelf, slope (50–1630 m). Central and Eastern Indo-Pacific, Temperate Australasia. 45 species (Cabezas and Macpherson 2014: key to 40 species; Cabezas et al. 2010: rediagnosis; Machordom and Macpherson 2004).

Plesionida Baba & de Saint Laurent, 1996 Diagnosis. Rostrum spiniform, overreaching obsolete supraocular spines. Carapace dorsal surface granulose, without transverse striae; mesogastric region with prominent median ridge-like process or elevated region. Pleonites 2–4 each with 2 submedian spines. Male pleopod 1 absent. Maximum cl. 11.4 mm. Slope (393–600 m). Western and Central Indo-Pacific. 4 species (Ahyong et al. 2013: key to species).

Pleuroncodes Stimpson, 1860 Diagnosis. Rostrum spiniform, overreaching supraocular spines. Carapace without gastric spines; mesogastric region without prominent median process (region may be spinose); pterygostomial flap visible in dorsal view. Pleonites 2–4 unarmed. Antennular basal article with 2 lateral spines. Male pleopod 1 present. Maximum cl. 44 mm.

282

Marine Decapod Crustacea

Subtidal–slope (18–523 m). Tropical Eastern Pacific, Temperate South America. 2 species. Both species of Pleuroncodes, P. monodon (H. Milne Edwards, 1837) and P. planipes Stimpson, 1860, are of commercial interest in Latin America where they are known as ‘langostinos’. Both are fished using bottom trawls, the ‘tails’ for human consumption and the exoskeletons (meal) as food for shrimp cultivation and as a source of pigmentation during salmon and trout culture (Wehrtmann and Acuña 2011). Both species are known to migrate up and down the continental margin (Thiel and Lovrich 2011).

Raymunida Macpherson & Machordom, 2000 Diagnosis. Rostrum spiniform, overreaching supraocular spines. Carapace with epigastric spines and 4 spines on branchial lateral margin; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 unarmed. Antennular basal article with 2 lateral spines. Male pleopod 1 present. Uropodal endopod with several long marginal spines. Maximum cl. 13 mm (Pl. 25b). Subtidal–slope (5–549 m). Temperate Northern W Pacific, Indo-West Pacific. 13 species (Macpherson and Machordom 2000; 2001: key to 7 species).

Sadayoshia Baba, 1969 Diagnosis. Rostrum spiniform, base broad, rostral and supraocular spines widely separated; with 2 pairs of supraocular spines. Carapace with 6–10 epigastric spines, 1 parahepatic and 1 anterior branchial spine on each side, laterally with 2 spines in front of and 5 behind anterior cervical groove; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 unarmed. Antennular basal article with 4 terminal spines, last article distal margin fringed with long setae. Antennal peduncle article 2 distomesial spine absent. Male pleopod 1 present. Maximum cl. 11 mm (Pl. 25e). Subtidal, shelf (5–90 m). Temperate Northern W Pacific, IndoWest Pacific. 14 species Macpherson and Baba (2010, 2012).

Setanida Macpherson, 2006 Diagnosis. Rostrum spiniform, overreaching supraocular spines. Orbit ventral margin visible in dorsal view as an acute distal angle and lateral spine. Carapace with pair of epigastric spines; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 unarmed. Antennular basal article with 2 lateral spines. Male pleopod 1 absent. Uropodal endopod with marginal spinules and several long spines. Maximum cl. 3 mm. Shelf (120–203 m). Eastern Indo-Pacific. 1 species (Macpherson 2006).

Tasmanida Ahyong, 2007 Diagnosis. Rostrum stout, longer than supraocular spines. Carapace with long, continuous transverse striae, with pair of epigastric and postcervical spines, and prominent median gastric

process; mesogastric region with prominent median ridge-like process or elevated region. Pleonites 2–4 with 2 pairs of spines. Antennular basal article with 1 distomesial and 2 distolateral spines. Male pleopod 1 absent. Maximum cl. 8 mm. Slope (521–539 m). Central Indo-Pacific (Norfolk Ridge). 1 species (Ahyong 2007).

Torbenella Baba, 2008 Diagnosis. Rostrum spiniform, remote from supraocular spines. Orbit with mound mesial to lateral limit. Carapace with small epigastric and hepatic spines, median gastric and cardiac regions unarmed; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 each having anterior ridge with pair of submedian spines. Antennular basal article with short distomesial and distolateral spines. Male pleopod 1 absent. Maximum cl. 12 mm. Shelf, slope (120–950 m). Western and Central Indo-Pacific. 3 species (Ahyong et al. 2013: key to species). References Ahyong ST (2007) Decapod Crustacea collected by the NORFANZ Expedition: Galatheidae and Polychelidae. Zootaxa 1593, 1–54. Ahyong ST, Baba K, Macpherson E, Poore GCB (2010) A new classification of the Galatheoidea (Crustacea: Decapoda: Anomura). Zootaxa 2676, 57–68. doi:10.11646/zootaxa.2676.1.4 Ahyong ST, Schnabel KE, Macpherson E (2011) Chapter 3. Phylogeny and fossil record of the marine squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 73–104. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Ahyong ST, Taylor J, McCallum AW (2013) New species and new records of deepwater munidid squat lobsters from north-western Australia: Onconida, Bathymunida, Crosnierita, Plesionida and Torbenella. Zootaxa 3734, 23–37. doi:10.11646/zootaxa.3734.1.3 Baba K (1993) Anomoeomunida, a new genus proposed for Phylladiorhynchus caribensis Mayo, 1972 (Crustacea: Decapoda: Galatheidae). Proceedings of the Biological Society of Washington 106, 102–105. Baba K (2005) Deep-sea chirostylid and galatheid crustaceans (Decapoda: Anomura) from the Indo-West Pacific, with a list of species. Galathea Report 20, 1–317. Baba K (2011) Babamunida debrae, a new species of squat lobster (Crustacea: Anomura: Galatheoidea: Munididae) from the Hawaiian Islands. Zootaxa 2812, 21–27. doi:10.11646/zootaxa.2812.1.2 Baba K, de Saint Laurent M (1996) Crustacea Decapoda: Revision of the genus Bathymunida Balss, 1914, and description of six new related genera (Galatheidae). In Résultats des Campagnes MUSORSTOM, volume 15. (Ed. A Crosnier). Mémoires du Muséum National d’Histoire Naturelle, Paris 168, 433–502. Cabezas P, Macpherson E (2014) A new species of Paramunida Baba, 1988 from the Central Pacific Ocean and a new genus to accommodate P. granulata (Henderson, 1885). ZooKeys 425, 15–32. doi:10.3897/zookeys.425.7882 Cabezas P, Macpherson E, Machordom A (2008) A new genus of squat lobster (Decapoda: Anomura: Galatheidae) from the South West Pacific and Indian Ocean inferred from morphological and molecular evidence. Journal of Crustacean Biology 28, 68–75. doi:10.1651/07-2823R.1 Cabezas P, Macpherson E, Machordom A (2010) Taxonomic revision of the genus Paramunida Baba, 1988 (Crustacea: Decapoda: Galatheidae): a morphological and molecular approach. Zootaxa 2712, 1–60. doi:10.11646/​ zootaxa.​2712.1.1

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

Komai T (2011) Squat lobsters of the genus Munida (Crustacea: Decapoda: Anomura: Munididae) from the Ogasawara Islands, with descriptions of four new species. Memoirs of the National Science Museum, Tokyo 47, 339–365. Komai T (2012) Squat Lobsters of the genus Munida Leach, 1820 (Crustacea: Decapoda: Anomura: Munididae) from the Sagami Sea and Izu Islands, Central Japan. Bulletin of the National Science Museum, Ser. Zoology (Jena, Germany) 12, 1–69. Komai T (2014) A new bathyal species of the squat lobster genus Munida Leach, 1820 (Crustacea: Decapoda: Anomura: Munididae) from Japan. Natural History Research 13, 11–18. Machordom A, Macpherson E (2004) Rapid radiation and cryptic speciation in galatheid crabs of the genus Munida and related genera in the South West Pacific: molecular and morphological evidence. Molecular Phylogenetics and Evolution 33, 259–279. doi:10.1016/j.ympev.2004.​ 06.001 Macpherson E (2006) Galatheidae (Crustacea, Decapoda) from the Austral Islands, Central Pacific. In Tropical Deep-Sea Benthos, volume 24 (Eds B Richer de Forges and JL Justine) Mémoires du Muséum National d’Histoire Naturelle, Paris 193, 285–333. Macpherson E, Rodríguez-Flores PC, Machordom A (2020) Squat lobsters of the families Munididae and Munidopsidae from Papua New Guinea. In: Corbari, L., Chan, T-Y and Ahyong, ST (eds) Papua New Guinea. Tropical Deep-Sea Benthos vol. 31. Mémoires du Muséum National d’Histoire Naturelle, Paris 213, 11–120. Macpherson E, Baba K (2006) New species and records of small galatheids (Crustacea, Decapoda, Galatheidae) from the southwest and central Pacific Ocean. Zoosystema 28, 443–456. Macpherson E, Baba K (2010) Revision of the genus Sadayoshia (Anomura, Galatheidae), with descriptions of four new species. In Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume. (Eds C Fransen, S De Grave and PKL Ng) Crustaceana Monographs 14, 415–452. Macpherson E, Baba K (2011) Chapter 2. Taxonomy of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 39–71. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Macpherson E, Baba K (2012) The squat lobsters of the genus Sadayoshia Baba, 1969 (Crustacea: Decapoda: Anomura: Munididae): new records including six new species from the Pacific Ocean. Zootaxa 3589, 30–48. doi:10.11646/zootaxa.3589.1.2 Macpherson E, Machordom A (2000) Raymunida, new genus (Decapoda: Anomura: Galatheidae) from the Indian and Pacific Oceans. Journal of Crustacean Biology 20, 253–258. doi:10.1163/1937240X-90000027 Macpherson E, Machordom A (2001) Phylogenetic relationships of species of Raymunida (Decapoda: Galatheidae) based on morphology and mitochondrial cytochrome oxidase sequences, with the recognition of four new species. Journal of Crustacean Biology 21, 696–714. doi:10.1163/​20021975-99990168 Macpherson E, Machordom A (2005) Use of morphological and molecular data to identify three new sibling species of the genus Munida Leach, 1820 (Crustacea, Decapoda, Galatheidae) from New Caledonia. Journal of Natural History 39, 819–834. doi:10.1080/00222930400002473 Macpherson E, Rodríguez-Flores PC, Machordom A (2017) New sibling species and new occurrences of squat lobsters (Crustacea, Decapoda) from the western Indian Ocean. European Journal of Taxonomy 343, 1–61. doi:10.5852/​ejt.2017.343 Pérez-Barros P, D’Amato ME, Guzman NV, Lovrich GA (2008) Taxonomic status of two South American sympatric squat lobsters, Munida gregaria and Munida subrugosa (Crustacea: Decapoda: Galatheidae), challenged by DNA sequence information. Biological Journal of the Linnean Society. Linnean Society of London 94, 421–434. doi:10.1111/j.​ 1095-8312.2008.00987.x

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Poore GCB, Andreakis N (2012) The Agononida incerta species complex unravelled (Crustacea: Decapoda: Anomura: Munididae). Zootaxa 3429, 1–29. Poore GCB, Andreakis N (2014) More species of the Agononida incerta complex revealed by molecules and morphology (Crustacea: Decapoda: Anomura: Munididae). Zootaxa 3860, 201–225. doi:10.11646/​ zootaxa.3860.3.1 Thiel M, Lovrich GA (2011) Chapter 7. Agonostic behaviour and reproductive biology of squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 223–247. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Wehrtmann IS, Acuña E (2011) Chapter 10. Squat lobster fisheries. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 297–322. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Zeldis JR (1985) Ecology of Munida gregaria (Decapoda, Anomura): distribution and abundance, population dynamics and fisheries. Marine Ecology Progress Series 22, 77–99. doi:10.3354/meps022077

Munidopsidae Ortmann, 1898 squat lobsters Figure 13.9, Plate 26 Munidopsids are a family of largely deep-sea squat lobsters. Most occur at slope to bathyal depths, but the family also includes the deepest occurring squat lobster, M. taiwanica Osawa, Lin & Chan, 2008, recorded at 5491 m in the Mariana Trench (Dong and Li 2018). Munidopsidae also boasts the only anchialine squat lobster, M. polymorpha Koelbel, 1892, living in shallow water caves in the Canary Islands. The phylogenetic separation of munidopsids from other galatheoid families is well attested (Ahyong et  al. 2011b; Bracken-Grissom et al. 2013; Palero et al. 2010), but the single morphological feature always separating munidopsids from galatheids and munidids is the absence of a flagellum on the exopod of maxilliped 1. Nevertheless, munidopsids almost always have reduced eyes, whereas the eyes are usually well developed in galatheids and munidids. Leiogalathea was recognised to be a munidopsid on genetic evidence (Ahyong et al. 2009); its 18 species have a reduced flagellum. The largest genus, Munidopsis, includes more than 270 species arrayed in many seemingly discrete species groups (some with currently synonymised names), that may ultimately be recognised as separate genera (Ahyong et  al. 2011a). Diagnosis. Galatheoidea. Body squat lobster-like. Rostrum triangular, quadrate or spiniform. Supraocular spines absent. Carapace smooth, tuberculate, spinose or with transverse striae. Tailfan well developed, folded under preceding pleonites. Maxilliped 1 exopod flagellum absent or reduced. Maxilliped 3 pediform; ischium and merus not expanded mesially. Maxilliped 3 with crista dentata; with epipod. Cheliped subcylindrical to ovate in cross-section.

284

Marine Decapod Crustacea

Fig. 13.9.  Munidopsidae. a, Galacantha subrostrata Macpherson, 2007; b, Leiogalathea laevirostris (Balss, 1913); c, Munidopsis andamanica MacGilchrist, 1905; d, Munidopsis pericalla Macpherson, 2007; e, Munidopsis amapa Poore, 2014. f, telson, Munidopsis.

Key to genera of Munidopsidae 1. – 2. – 3. –

Eyes well developed. Carapace with obsolescent transverse striae bearing coarse setae (Fig. 13.9b). Maxilliped 1 exopod flagellum reduced������������������������������������������������������������������������������������������������������������� Leiogalathea Eyes reduced. Carapace without transverse striae and coarse setae. Maxilliped 1 exopod flagellum absent�������������������2 Ventral surface of cephalothorax with mat of long silky setae (Pl. 26f). Maxilliped 3 with rudimentary epipod������������������������������������������������������������������������������������������������������������������������������������������������������������������������������ Shinkaia Ventral surface of cephalothorax without mat of long silky setae. Maxilliped 3 with well developed epipod������������������3 Carapace (mesogastric and cardiac regions) and pleonites 2–4 with row of prominent spines in midline (Fig. 13.9a)����������������������������������������������������������������������������������������������������������������������������������������������������������������� Galacantha Carapace and pleon without complete row of spines in midline (Fig. 13.9c–e)�������������������������������������������������� Munidopsis Galacantha A. Milne-Edwards, 1880

Diagnosis. Rostrum triangular, flattish. Carapace dorsal surface without transverse setiferous striae; mesogastric region with extremely strong erect laterally-compressed spine; cardiac region with moderately strong spine. Pleonites 2–4 each with strong sharp median spine. Cornea not pigmented. Maxilliped 1 exopod flagellum absent. Maxilliped 3 with epipod. Maximum cl. 37 mm (Pl. 26a). Slope–bathyal (277–3800 m). Cosmopolitan. 11 species (Macpherson 2007: key to 9 species; Osawa et al. 2013).

Leiogalathea Baba, 1969 Diagnosis. Rostrum triangular, flattish. Carapace dorsal surface with obsolete transverse striae bearing coarse setae; mesogastric

region without prominent median process. Pleonites 2–4 unarmed. Cornea well pigmented. Maxilliped 1 exopod flagellum reduced. Maxilliped 3 with epipod. Maximum cl. 5.5 mm (Pl. 26b). Shelf, slope (160–1642 m). Tropical W Atlantic, Indo-West Pacific. 18 species (Rodríguez-Flores et al. 2019: key to species).

Munidopsis Whiteaves, 1874 Diagnosis. Rostrum spiniform to triangular, flattish. Carapace dorsal surface without transverse setiferous striae; mesogastric region without prominent median process (region may be spinose). Pleonites 2–4 variously armed but without row of single median spines. Cornea not pigmented or well pigmented. Maxilliped 1 exopod flagellum absent. Maxilliped 3 merus at most 1.5 times as long as broad; with epipod. Maximum cl. 70 mm (Pl. 26c–e).

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

Shelf–abyssal (75–5491 m), marine anchialine caves (2–8 m). Cosmopolitan except Arctic, Antarctic. 272 species. Numerous species have been described during the last decade or so from the Indo-West Pacific and temperate Australasia (e.g. Ahyong 2014; Macpherson 2007; Macpherson et  al. 2020; Marin 2020; Osawa et  al. 2008; Osawa et  al. 2013) and the Americas (e.g. Poore 2014; Rodríguez-Flores et al. 2018). Munidopsis polymorpha Keolbel, 1892 is unusual in that it is endemic to an anchialine cave system of the Corona lava tube in Lanzarote (Canary Islands) where it is considered ‘endangered’ (Cabezas et al. 2012; Wilkens et al. 1990).

Shinkaia Baba & Williams, 1998 Diagnosis. Rostrum prominent, flattened dorsally, straplike in outline. Carapace dorsal surface without transverse setiferous striae; mesogastric region without prominent median process (region may be spinose); cephalothorax with ventral mat of long silky setae. Pleonites 2–4 unarmed. Eyestalk a dorsally flattened projection; cornea not pigmented. Maxilliped 1 exopod flagellum absent. Maxilliped 3 with rudimentary epipod. Maximum cl. 53 mm. Slope (600–1500 m). Temperate Northern W Pacific (active hydrothermal vents in the Okinawa Trough and elsewhere). 1 species (Chan et al. 2000; Tsuchida et al. 2003). Filamentous bacteria attach to silky setae on the ventral surface and are a source of food (Tsuchida et al. 2011; Watsuji et al. 2010). References Ahyong ST (2014) Deep-sea squat lobsters of the Munidopsis serricornis complex in the Indo-West Pacific, with descriptions of six new species (Crustacea: Decapoda: Munidopsidae). Records of the Australian Museum 66, 197–216. doi:10.3853/j.2201-4349.66.2014.1630 Ahyong ST, Schnabel KE, Maas EW (2009) Anomuran phylogeny: new insights from molecular data. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 399–414. CRC Press, Boca Raton. Ahyong ST, Andreakis N, Taylor J (2011a) Mitochondrial phylogeny of the deep-sea squat lobsters, Munidopsidae (Galatheoidea). Zoologischer Anzeiger 250, 367–377. doi:10.1016/j.jcz.2011.06.005 Ahyong ST, Schnabel KE, Macpherson E (2011b) Chapter 3. Phylogeny and fossil record of the marine squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 73–104. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Bracken-Grissom HD, Cannon ME, Cabezas P, Feldmann RM, Schweitzer CE, et  al. (2013) A comprehensive and integrative reconstruction of evolutionary history for Anomura (Crustacea: Decapoda). BMC Evolutionary Biology 13, 128 doi:10.1186/1471-2148-13-128. Cabezas P, Alda F, Macpherson E, Machordom A (2012) Genetic characterization of the endangered and endemic anchialine squat lobster Munidopsis polymorpha from Lanzarote (Canary Islands): management implications. ICES Journal of Marine Science 69, 1030–1037. doi:10.1093/icesjms/fss062 Chan T-Y, Lee DA, Lee CS (2000) The first deep-sea hydrothermal animal reported from Taiwan: Shinkaia crosnieri Baba and Williams, 1998 (Crustacea: Decapoda: Galatheidae). Bulletin of Marine Science 67, 799–804.

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Dong D, Li X (2018) New record of Munidopsis taiwanica (Decapoda, Anomura) from a mud volcano field in the Mariana Trench, with in-situ observations on habitat. Crustaceana 91, 363–373. doi:10.1163/15685403-00003773 Macpherson E (2007) Species of the genus Munidopsis Whiteaves, 1784 from the Indian and Pacific Oceans and reestablishment of the genus Galacantha A. Milne-Edwards, 1880 (Crustacea, Decapoda, Galatheidae). Zootaxa 1417, 1–135. doi:10.11646/zootaxa.1417.1.1 Macpherson E, Rodríguez-Flores PC, Machordom A (2020) Squat lobsters of the families Munididae and Munidopsidae from Papua New Guinea. In: Corbari, L., Chan, T-Y and Ahyong, ST (eds) Papua New Guinea. Tropical Deep-Sea Benthos vol. 31. Mémoires du Muséum National d’Histoire Naturelle, Paris 213, 11–120. Marin I (2020) Northern unicorns of the depths: diversity of the genus Munidopsis Whiteaves, 1874 (Decapoda: Anomura: Munidopsidae) in the northwestern Pacific Ocean, with descriptions of three new species along the Russian coast. Progress in Oceanography 183, 102263. doi:10.1016/j.pocean.2020.102263 Osawa M, Lin C-W, Chan T-Y (2008) Species of Galacantha and Munidopsis (Crustacea: Decapoda: Anomura: Galatheidae) from the deep-waters off Taiwan, with the description of two new species. Scientia Marina 72, 35–57. Osawa M, Lin CW, Chan TY (2013) Munidopsidae Ortmann, 1898 (Crustacea, Decapoda, Anomura) collected by the PANGLAO 2005 and AURORA expeditions to the Philippines, with descriptions of four new species from the Philippines and one new species from Taiwan. In: Ahyong, ST, Chan, TY, Corbari, L. and Ng PKL (eds), Tropical Deep-Sea Benthos vol. 27. Mémoires du Muséum National d’Histoire Naturelle 204, 231–286. Palero F, Abelló P, Macpherson E, Matthee CA, Pascual M (2010) Genetic diversity levels in fishery-exploited spiny lobsters of the genus Palinurus (Decapoda: Achelata). Journal of Crustacean Biology 30, 658–663. doi:10.1651/09-3192.1 Poore GCB (2014) Three new American species of Munidopsis (Crustacea: Anomura: Munidopsidae). Nauplius 22, 53–62. doi:10.1590/S010464972014000100006 Rodríguez-Flores PC, Macpherson E, Machordom A (2018) Three new species of squat lobsters of the genus Munidopsis Whiteaves, 1874, from Guadeloupe Island, Caribbean Sea (Crustacea, Decapoda, Munidopsidae). Zootaxa 4422, 569–580. doi:10.11646/zootaxa.​ 4422.4.7 Rodríguez-Flores PC, Macpherson E, Machordom A (2019) Revision of the squat lobsters of the genus Leiogalathea Baba, 1969 (Crustacea, Decapoda, Munidopsidae) with the description of 15 new species. Zootaxa 4560, 201–256. doi:10.11646/zootaxa.4560.2.1 Tsuchida S, Fujiwara Y, Fujikura K (2003) Distribution and population structure of the galatheid crab Shinkaia crosnieri (Decapoda: Anomura: Galatheidae) in the southern Okinawa Trough. Japanese Journal of Benthology 58, 84–88[Japanese]. doi:10.5179/benthos.58.84 Tsuchida S, Suzuki Y, Fujiwara Y, Kawato M, Uematsu K, et  al. (2011) Epibiotic association between filamentous bacteria and the vent-associated galatheid crab, Shinkaia crosnieri (Decapoda: Anomura). Journal of the Marine Biological Association of the United Kingdom 91, 23–32. doi:10.1017/S0025315410001827 Watsuji T, Nakagawa S, Tsuchida S, Toki T, Hirota A, et al. (2010) Diversity and function of epibiotic microbial communities on the galatheid crab, Shinkaia crosnieri. Microbes and Environments 25, 288–294. doi:10.1264/jsme2.ME10135 Wilkens H, Parzefall J, Ribowski A (1990) Population biology and larvae of the anchialine crab Munidopsis polymorpha (Galatheidae) from Lanzarote (Canary Islands). Journal of Crustacean Biology 10, 667–675. doi:10.2307/1548411

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Porcellanidae Haworth, 1825 porcelain crabs, half-crabs Figures 13.10–13.12, Plates 27, 28 Porcellanidae, more commonly known as porcelain crabs or half-crabs, typically have a short carapace, ventrally flexed pleon, and large, robust chelipeds usually held across the front of the body, making them appear very much like true brachyuran crabs. Porcellanids are most closely related to the galatheoid squat lobsters, from which they are readily distinguished by the operculiform rather than pediform maxilliped 3 and smaller, ventrally flexed pleon. Unlike galatheids, porcellanids are primarily filter feeders, using their strongly setose maxilliped 3 to strain food particles from the water. The flattened body of most porcellanids enables them to live in tight crevices in and under rocks and coral. Most are free-living but many are commensal with other invertebrates including gorgonians, soft and hard corals, sea anemones, echinoderms and tubicolous polychaete worms. The Australian Polyonyx transversus (Haswell, 1882) is possibly also associated with the bivalve mollusc, Brechites (as Aspergillum) (Haig 1965). Descriptions of larval stages were reviewed by Vela and González-Gordillo (2016). More than 300 species are known, and all occur at shelf depths, mostly on shallow reefs. The eastern Pacific and Atlantic and porcellanids were substantially treated by Haig (1960, 1965) and Chace (1956); Rodríguez et  al. (2005) reviewed references to the western Atlantic fauna. The IndoWest Pacific porcellanids have not been treated synoptically, but important regional works include Haig (1965), Miyake (1978), Tirmizi et al. (1989), Poore (2004), Osawa (2007) and Osawa and Chan (2010). Most porcellanid genera are small, particularly following partial revision of Porcellana by Haig (1978), but larger

genera remain taxonomically challenging. The large cosmopolitan genera Pachycheles and Petrolisthes and several others are probably not monophyletic (Bracken-Grissom et al. 2013; Werding et al. 2001) and subject to ongoing revision. To date, however, phylogenetic studies have been largely focused on the eastern Pacific and Atlantic fauna (Baeza 2016; Rodríguez et al. 2006; Stillman and Reeb 2001), and none has sufficiently accounted for Indo-West Pacific diversity. Baeza (2016) proposed sinking Liopetrolisthes and Allopetrolisthes into Petrolisthes, but we recognise all three as separate. Although listed as valid by (Osawa and Chan 2010) and Osawa and McLaughlin (2010), we follow Rodríguez et al. (2005; 2006) in treating Madarateuchis Harvey, 1999 as a junior synonym of Clastotoechus Haig, 1960 based on results of phylogenetic analysis. Twenty-nine genera are recognised here. Note that Pisidia appears twice in the key, to accommodate the atypical species, P. dispar (Stimpson, 1858), which may ultimately belong in a separate genus (Osawa and Chan 2010). In some genera the movable articles (2–4) of the antennal peduncle are excluded from orbit by anterior projection of immovable article 1 appressed to anterior margin of carapace (Fig. 13.12g, h, j) while in others these articles are freely accessible to orbit (Fig. 13.12i). Diagnosis. Galatheoidea. Body crab-like. Rostrum usually obsolete. Supraocular spines present, or absent. Carapace smooth or with faint striae or tubercles. Tailfan well developed, folded under preceding pleonites. Maxilliped 1 exopod flagellum well developed. Maxilliped 3 operculiform; ischium and merus broad, expanded mesially. Maxilliped 3 without crista dentata; without epipod. Cheliped usually flattened. Implicit generic attribute. Chelipeds held strongly bent at propodus-carpus articulation. Pereopods 2–4 dactyli curved, stout. Telson of 7 plates.

Key to genera of Porcellanidae 1. – 2. – 3. – 4. – 5.

Chelipeds held directed straight forwards (Figs 13.11a, 13.12n)�����������������������������������������������������������������������������������������������2 Chelipeds held strongly bent at propodus-carpus articulation�������������������������������������������������������������������������������������������������3 Cheliped carpus length at least twice width. Pereopods 2–4 dactyli terminating in single claw (Fig. 13.11a)Orthochela Cheliped carpus wider than long (Fig. 13.12n). Pereopods 2–4 dactyli triunguiculate (Fig. 13.12v)�Pseudoporcellanella Carapace at least 1.5 times as long as wide or less (Fig. 13.10g)������������������������������������������������������������������������������ Euceramus Carapace 1.3 times as long as wide������������������������������������������������������������������������������������������������������������������������������������������������4 Antennal peduncle with movable articles (2–4) freely accessible to orbit (Fig. 13.12h, i), or only partially excluded from orbit (Allopetrolisthes only)�����������������������������������������������������������������������������������������������������������������������������5 Antennal peduncle with movable articles excluded from orbit orbit by anterior projection of immovable article 1 appressed to anterior margin of carapace (Fig. 13.12g, j)������������������������������������������������������������������������������������14 Carapace pterygostomial flap incomplete, either consisting of one large anterior piece and 1 or more posterior pieces separated from each other by membranous interspaces (Fig. 13.12a, f), or comprising single calcified anterior plate extending approximately to level of pereopod 2 followed posteriorly by membranous portion (Fig. 13.12d)�������������������������������������������������������������������������������������������������������������������������������������������6

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters



287

Carapace pterygostomial flap entire, a single, undivided plate, extending posteriorly as far as pereopod 4 (Fig. 13.12e)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������8 6. Chelipeds flattened, subequal. Carapace front strongly produced, distinctly trilobed in dorsal view (Fig. 13.10e)����������������������������������������������������������������������������������������������������������������������������������������������������������� Clastotoechus – Chelipeds thick, robust, distinctly unequal. Carapace front not prominent, sinuous to transverse or broadly rounded in dorsal view (Figs 13.10p, 13.11b, c)��������������������������������������������������������������������������������������������������������7 7. Carapace pterygostomial flap composed of single calcified anterior plate extending approximately to level of pereopod 2 followed posteriorly by entirely membranous portion (Fig. 13.12d)���������������������������Neopisosoma – Carapace pterygostomial flap composed of 2 or more pieces separated from anterior part by membranous interspaces (Fig. 13.12f)������������������������������������������������������������������������������������������������������������������������������������������� Pachycheles 8. Telson composed of 5 plates (Fig. 13.12o, q)��������������������������������������������������������������������������������������������������������������������������������9 – Telson composed of 7 plates (Fig. 13.12p, r)�������������������������������������������������������������������������������������������������������������������������������11 9. Cheliped carpus anterior and posterior margins without spines, at most with angular lobe on anterior margin (Fig. 13.10b). Antennal peduncle with movable articles (2–4) partially excluded from orbit by anterior projection of immovable article 1 narrowly meeting anterior margin of carapace������������������Allopetrolisthes – Cheliped carpus anterior and posterior margins with row of spines. Antennal peduncle with movable articles (2–4) freely accessible to orbit�����������������������������������������������������������������������������������������������������������������������������������10 10. Carapace lateral margins spinose. Cheliped carpus dorsally spinose (Fig. 13.11e)���������������������������������������������� Petrocheles – Carapace lateral margins unarmed. Cheliped carpus dorsally granulate (Fig. 13.10k)�������������������������������� Liopetrolisthes 11. Carapace gastric region strongly elevated (Figs 13.10o, 13.12c). Pereopods 2–4 subcylindrical (Fig. 13.12t)����������������������������������������������������������������������������������������������������������������������������������������������������������Neopetrolisthes – Carapace dorsal surface flattish or weakly convex. Pereopods 2–4 flattened��������������������������������������������������������������������� 12 12. Cheliped fingers distinctly gaping, tips hollowed, scoop-like (Fig. 13.11d)��������������������������������������������������Parapetrolisthes – Cheliped fingers without gape, occluding along length of opposable margin, tips simple������������������������������������������������13 13. Carapace dorsal surface uneven, regions well defined; median frontal lobe with pair of distal elevations separated by deep longitudinal groove. Eyestalk distinctly wider than cornea in dorsal view (Fig. 13.10q)��������������������������������������������������������������������������������������������������������������������������������������������������� Novorostrum – Carapace dorsal surface generally even, regions weakly or undefined; median frontal lobe without lateral elevations, at most with shallow longitudinal groove. Eyestalk as wide as cornea in dorsal view (Fig. 13.11f)�����������������������������������������������������������������������������������������������������������������������������������������������������������������Petrolisthes 14. Carapace distinctly elongate, ~1.3 times as long as wide (Figs 13.10n, 13.11k, n)����������������������������������������������������������������15 – Carapace less than 1.1 times as long as wide������������������������������������������������������������������������������������������������������������������������������17 15. Pereopods 2–4 dactyli quadriunguiculate (Fig. 13.12v). Carapace median frontal lobe strongly produced in advance of lateral lobes (Fig. 13.11k)������������������������������������������������������������������������������������������������������������������� Porcellanella – Pereopods 2–4 dactyli terminating in single claw. Carapace frontal lobes in essentially transverse row������������������������16 16. Carapace dorsal surface covered with rugose protuberances; front quadridentate in dorsal view; lateral margin without anterior branchial spine, with granular lobes or blunt teeth (Fig. 13.11n)���������������������������������Ulloaia – Carapace dorsal surface largely smooth, with weak transverse striations; front tridentate in dorsal view; lateral margins unarmed behind anterior branchial spine (Fig. 13.10n)��������������������������������������������������������� Minyocerus 17. Pereopods 2–4 dactyli straight, slender, terminating in simple sharp point, without spines on flexor margin (Fig. 13.12w)��������������������������������������������������������������������������������������������������������������������������������������������������������������Raphidopus – Pereopods 2–4 dactyli curved, stout, terminating in simple or biunguiculate claw, usually with 1 or more spines on flexor margin�����������������������������������������������������������������������������������������������������������������������������������������������������������18 18. Carapace up to ~1.5 times as wide as long, front strongly curled ventrally, in dorsal view appearing straight or weakly sinuous����������������������������������������������������������������������������������������������������������������������������������������������������������������������19 – Carapace little wider than long to little longer than wide, front largely horizontal or down-tilted as a result of broad furrow on anteromedian surface, in dorsal view appearing multilobed or broadly angular-rounded���������������� 22 19. Carapace cervical groove with margins forming pair of transverse ridges (Fig. 13.10i). Telson of 5 plates (Fig. 13.12o)������������������������������������������������������������������������������������������������������������������������� Heteropolyonyx

288

Marine Decapod Crustacea

– Carapace cervical grooves weak or distinct but without transverse ridges. Telson of 7 plates����������������������������������������� 20 20. Carapace regions well defined by grooves; cervical and gastric grooves formed by thinning and gradual decalcification of cuticle (Fig. 13.10h)�����������������������������������������������������������������������������������������������������������������������Eulenaios – Carapace regions not distinctly defined by grooves (Fig. 13.11i)��������������������������������������������������������������������������������Polyonyx 21. Carapace front broadly angular-rounded in dorsal view (Figs 13.10a, c, m, 13.11h)���������������������������������������������������������� 22 – Carapace front distinctly multilobed in dorsal view (Figs 13.10d, f, l, 13.11h, j)����������������������������������������������������������������� 25 22. Pereopods 2–4 dactyli biunguiculate (Fig. 13.12s)����������������������������������������������������������������������������������������������Aliaporcellana – Pereopods 2–4 dactyli simple������������������������������������������������������������������������������������������������������������������������������������������������������ 23

Fig. 13.10.  Porcellanidae. Habitus, dorsal carapace, some with chelipeds: a, Aliaporcellana; b, Allopetrolisthes; c, Ancylocheles gravelei (Sankolli, 1963); d, Capilliporcellana murakamii (Miyake, 1942); e, Clastotoechus; f, Enosteoides philippinensis Dolorosa & Werding, 2014; g, Euceramus transversilineatus (Lockington, 1878); h, Eulenaios cometes (Walker, 1887); i, Heteropolyonyx biforma Osawa, 2001; j, Heteroporcellana corbicola (Haig, 1960); k, Liopetrolisthes mitra (Dana, 1852); l, Lissoporcellana; m, Megalobrachium; n, Minyocerus kirki Glassell, 1938; o, Neopetrolisthes; p, Neopisosoma; q, Novorostrum indicum (De Man, 1893).

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

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23. Chelipeds subequal; fingers not twisted out of plane with palm (Fig. 13.10m)������������������������������������������ Megalobrachium – Chelipeds usually distinctly unequal; fingers of smaller cheliped in large adults twisted out of plane with palm���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 24 24. Orbit ill-defined in dorsal view, exorbital angle obtuse, blunt, unarmed (Fig. 13.11g)�����������������������������������������������������������������������������������������������������������������������������Pisidia dispar (Stimpson, 1858) – Orbit well defined, exorbital angle forming a small spine (Fig. 13.10c)�������������������������������������������������������������� Ancylocheles 25. Carapace subquadrate, lateral margins largely straight, subparallel (Fig. 13.10j). Telson of 5 plates������Heteroporcellana – Carapace rounded-ovate, branchial margins curved. Telson of 7 plates������������������������������������������������������������������������������ 26 26. Carapace dorsal surface areolate, regions well defined����������������������������������������������������������������������������������������������������������� 27 – Carapace dorsal surface largely smooth or with fine striations��������������������������������������������������������������������������������������������� 28 27. Carapace lateral margin smooth, unarmed (Fig. 13.10d)�������������������������������������������������������������������������������Capilliporcellana – Carapace lateral margin with 2–6 small anteriorly directed spines (Fig. 13.10f)������������������������������������������������ Enosteoides

Fig. 13.11.  Porcellanidae. Habitus, dorsal carapace, some with chelipeds: a, Orthochela pumila Glassell, 1936; b, Pachycheles granti Haig, 1965; c, Pachycheles tuerkayi Werding & Hiller, 2017; d, Parapetrolisthes tortugensis (Glassell, 1945); e, Petrocheles australiensis (Miers, 1876); f, Petrolisthes elongatus (Miers, 1876); g, Pisidia dispar (Stimpson, 1858); h, Pisidia serratifrons (Stimpson, 1858); i, Polyonyx transversus (Haswell, 1882); j, Porcellana lillyae Lemaitre & Campos, 2000; k, Porcellanella; l, Pseudoporcellanella; m, Raphidopus indicus Henderson, 1893; n, Ulloaia perpusillia Glassell, 1938.

290

Marine Decapod Crustacea

Fig. 13.12.  Porcellanidae. Left lateral view, pterygostomial flap: a, Clastotoechus; b, Heteropolyonyx; c, Neopetrolisthes; d, Neopisosoma; e, Novorostrum; f, Pachycheles. Left orbit, antenna: g, Heteropolyonyx; h, Neopisosoma; i, Pachycheles; j, Raphidopus. Cheliped pair: k, l, Lissoporcellana. Right cheliped: m, Neopisosoma; n, Pseudoporcellanella. Telson: o, Heteropolyonyx; p, Neopetrolisthes; q, Petrocheles; r, Petrolisthes. Pereopod 2: s, Aliaporcellana; t, Neopetrolisthes; u, Porcellanella; v, Pseudoporcellanella; w, Raphidopus.

28. Carapace hepatic spine absent; branchial margin unarmed behind epibranchial angle. Chelipeds subequal; dactylus in line with plane of palm, not twisted (Fig. 13.11j)������������������������������������������������������������������������������ Porcellana – Carapace hepatic spine usually present; branchial margin usually with 1 or more spines behind epibranchial angle (Figs 13.10l, 13.11h). Chelipeds usually unequal; dactylus of smaller cheliped often twisted out of plane with palm in large adults (Fig. 13.11h)����������������������������������������������������������������������������������������������������������������������� 29 29. Carapace median frontal lobe usually with median notch (Fig. 13.10l). Chelipeds with similar spination in both sexes��������������������������������������������������������������������������������������������������������������������������������������Lissoporcellana – Carapace median frontal lobe without median notch (Fig. 13.11h). Chelipeds more spinose in females than males������������������������������������������������������������������������������������������������������������������������������������������� Pisidia (part) Aliaporcellana Nakasone & Miyake, 1969 Diagnosis. Carapace rounded, as long as wide or wider than long; lateral margins spinose; front down-tilted as result of broad furrow on anteromedian surface, appearing broadly angular in dorsal view; exorbital angle forming small spine. Antennal peduncle with movable articles excluded from orbit. Chelipeds usually unequal. Pereopods 2–4 dactyli terminating in biunguiculate claw, with or without small spine on flexor margin. Maximum cl. 7 mm (Pl. 27a). Intertidal–shelf (0–180 m; associated with sponges and cnidarians). Western and Central Indo-Pacific. 6 species (Dong et  al. 2011: key to 5 species; Hiller and Werding 2018; Osawa 2007: key to 4 species).

Allopetrolisthes Haig, 1960 Diagnosis. Carapace rounded, about as long as wide; lateral margins unarmed; front weakly projecting, with 3 or 5 lobes in dorsal view;

pterygostomial flap a single, undivided plate, extending posteriorly as far as pereopod 4. Antennal peduncle with movable articles partially excluded from orbit. Chelipeds subequal; carpus anterior and posterior margins without spines, at most with angular lobe on anterior margin. Telson of 5 plates. Maximum cl. 38 mm (Pl. 27b). Intertidal, subtidal (0–13 m). Eastern Indo-Pacific, Temperate South America (Pacific coast). 3 species (Baeza 2016: polyphyly; Haig 1960: key to 3 species).

Ancylocheles Haig, 1978 Diagnosis. Carapace rounded, as long as wide; lateral margins spinose; front largely horizontal or slightly down-tilted, appearing broadly angular in dorsal view; exorbital angle forming small spine. Antennal peduncle with movable articles excluded from orbit. Chelipeds usually unequal. Smaller cheliped fingers in large adults twisted out of plane with palm. Pereopods 2–4 dactyli terminating in simple claw, with small spines on flexor margin. Maximum cl. 7 mm.

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

Intertidal, subtidal (0–17 m). Western and Central IndoPacific. 2 species (Haig 1978: diagnosis; Sankolli 1963: figure of type species; Trivedi et al. 2017: second species).

Capilliporcellana Haig, 1978 Diagnosis. Carapace cordiform, wider than long; lateral margins unarmed; dorsal surface areolate, regions well defined, front horizontal, trilobed in dorsal view, median lobe wider and longer than lateral lobes. Antennal peduncle with movable articles excluded from orbit. Pereopods 2–4 dactyli terminating in simple claw, with small spines on flexor margin. Maximum cl. 7 mm. Intertidal, shelf (0–31 m). Central Indo-Pacific. 2 species (Haig 1978: diagnosis; Haig 1981: second species; Osawa 2007: key to species).

Clastotoechus Haig, 1960 Diagnosis. Carapace rounded, about as long as wide; front strongly produced, distinctly trilobed in dorsal view; pterygostomial flap incomplete, posterior portion composed of many small pieces separated by membranous interspaces. Antennal peduncle with movable articles freely accessible to orbit. Chelipeds subequal. Telson of 5 plates. Maximum cl. 11 mm. Intertidal, subtidal (0–18 m). Tropical W Atlantic, Tropical Eastern Pacific. 6 species (Haig 1960: diagnosis; Harvey 1999: key to species; Madarateuchis is a synonym).

Enosteoides Johnson, 1970 Diagnosis. Carapace ovate, about as long as wide; lateral margins with 2–6 small anteriorly directed spines or serrations; dorsal surface areolate, regions well defined, front largely horizontal or down-tilted as a result of broad furrow on anteromedian surface, appearing trilobed, quadrilobed or broadly angular in dorsal view. Antennal peduncle with movable articles excluded from orbit. Chelipeds subequal. Pereopods 2–4 dactyli terminating in simple claw, with small spines on flexor margin. Maximum cl. 10 mm. Intertidal–shelf (0–150 m). Western and Central Indo-Pacific. 6 species (Dolorosa and Werding 2014: comparison of 5 species; Johnson 1970: diagnosis; Osawa 2007: key to 3 species; Osawa 2016: diagnosis, redescriptions).

Euceramus Stimpson, 1860 Diagnosis. Carapace longer than wide; lateral margins unarmed behind cervical groove. Pereopods 2–4 dactyli terminating in simple claw. Maximum cl. 16 mm (Pl. 27c). Intertidal–shelf (0–40 m). Tropical W Atlantic, Tropical Eastern Pacific. 3 species (Glassell 1938: figure of type species; Haig 1960: rediagnosis, key to 2 species).

Eulenaios Ng & Nakasone, 1993 Diagnosis. Carapace transversely ovate, wider than long; dorsal surface with regions well defined, cervical and gastric grooves

291

formed by thinning and gradual decalcification of cuticle, without transverse ridges, front curled ventrally, appearing weakly curved in dorsal view; exorbital angle unarmed. Antennal peduncle with movable articles excluded from orbit. Chelipeds unequal. Pereopods 2–4 dactyli terminating in biunguiculate claw, with 0–2 small spines on flexor margin. Maximum cl. 8 mm. Intertidal, subtidal (0–20 m; commensal with Chaetopterus spp., Polychaeta). Central Indo-Pacific. 1 species (Ng and Nakasone 1993).

Heteropolyonyx Osawa, 2001 Diagnosis. Carapace subquadrate, wider than long; lateral margins largely straight, subparallel; dorsal surface with cervical groove margins forming pair of transverse ridges, front curled ventrally, appearing straight or weakly sinuous in dorsal view; exorbital angle unarmed. Antennal peduncle with movable articles excluded from orbit. Chelipeds unequal. Pereopods 2–4 dactyli terminating in biunguiculate claw, with 2 or 3 spines on flexor margin. Telson of 5 plates. Maximum cl. 3 mm. Subtidal (commensal with Chaetopterus spp., Polychaeta). Central Indo-Pacific. 1 species (Osawa 2001).

Heteroporcellana Haig, 1978 Diagnosis. Carapace rounded, as long as wide; lateral margins unarmed behind anterior branchial spine; front horizontal, trilobed, median lobe wider and longer than lateral lobes. Antennal peduncle with movable articles excluded from orbit. Chelipeds subequal; fingers opening vertically. Pereopods 2–4 dactyli terminating in simple claw. Telson of 5 plates. Maximum cl. 6 mm. Subtidal, shelf (18–46 m). Tropical Eastern Pacific. 1 species (Haig 1960: figure of type species; 1978: diagnosis).

Liopetrolisthes Haig, 1960 Diagnosis. Carapace ovate or subcircular, as long as wide; lateral margins unarmed; front projecting, trilobed, median lobe appearing triangular in dorsal view; pterygostomial flap a single, undivided plate, extending posteriorly as far as pereopod 4. Antennal peduncle with movable articles freely accessible to orbit. Chelipeds subequal; carpus anterior margin spinose, upper surface granular. Telson of 5 plates. Maximum cl. 12 mm (Pl. 27d). Intertidal, subtidal (0–20 m). Temperate South America (Pacific coast). 2 species (Baeza 2016: phylogeny, figure; Haig 1960: diagnosis).

Lissoporcellana Haig, 1978 Diagnosis. Carapace subcircular, as long as wide or slightly longer than wide; lateral margins curved, usually with 1 or more spines behind epibranchial angle (absent in L. nitida); dorsal surface smooth or with fine striations, front horizontal, distinctly multilobed in dorsal view, median frontal lobe usually with median notch; exorbital angle forming small spine; hepatic spine usually present. Antennal peduncle with movable articles excluded from

292

Marine Decapod Crustacea

orbit. Chelipeds usually unequal. Smaller cheliped fingers often twisted out of plane with palm in large adults. Maximum cl. 9 mm (Pl. 27e). Intertidal–shelf (0–128 m; associated with sponges and cnidarians). Western and Central Indo-Pacific. 10 species (Haig 1978: diagnosis; Osawa 2007: key to species).

Megalobrachium Stimpson, 1858

Antennal peduncle with movable articles freely accessible to orbit. Chelipeds thick, robust, distinctly unequal. Telson of 5 or 7 plates. Maximum cl. 8 mm (Pl. 27i). Intertidal, subtidal (0–18 m). Tropical W Atlantic, Tropical Eastern Pacific. 7 species (Haig 1960: diagnosis, key to 3 E Pacific species; Miranda and Mantelatto 2016: probable polyphyly; Rodríguez et al. 2005: 2 groups recognised; Werding 1986: key to 4 W Atlantic species).

Diagnosis. Carapace cordiform or subcircular, about as long as wide; lateral margins unarmed, or with 5–6 stout, pointed, laterally directed tubercles; dorsal surface smooth or with fine striations or areolate, front largely horizontal or slightly down-tilted, appearing broadly angular or rounded in dorsal view. Antennal peduncle with movable articles excluded from orbit. Chelipeds subequal. Pereopods 2–4 dactyli terminating in simple claw, with small spines on flexor margin. Telson of 5 or 7 plates. Maximum cl. 9 mm (Pl. 27f, g). Intertidal–shelf (0–50 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific, Temperate South America (Atlantic coast). 13 species (Ferreira and Tavares 2017: descriptions of 2 species; Haig 1960: rediagnosis, key to 8 E Pacific species).

Diagnosis. Carapace cordiform or subcircular, about as long as wide or slightly wider than long; lateral margins unarmed; dorsal surface flattish or weakly convex but distinctly uneven, regions well defined, front produced, broad, trilobed, median lobe with deep longitudinal groove separating pair of distal elevations; pterygostomial flap a single, undivided plate, extending posteriorly as far as pereopod 4. Antennal peduncle with movable articles freely accessible to orbit. Eyestalk distinctly wider than cornea in dorsal view. Chelipeds subequal. Pereopods 2–4 flattened. Maximum cl. 8 mm (Pl. 27j). Intertidal. Central Indo-Pacific. 4 species (Ferreira and Tavares 2020: redescription; Osawa 1998: diagnosis, 3 species).

Minyocerus Stimpson, 1858

Orthochela Glassell, 1936

Diagnosis. Carapace ~1.3 times as long as wide; lateral margins unarmed behind anterior branchial spine; dorsal surface with fine striations, front tridentate in dorsal view, lobes in transverse row. Antennal peduncle with movable articles excluded from orbit. Chelipeds subequal. Pereopods 2–4 dactyli terminating in simple claw. Maximum cl. 4 mm. Intertidal–shelf (0–59 m; associated with asteroids, Luidia spp., and ophiuroids). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Glassell 1938: figure).

Diagnosis. Carapace ~1.5 times as long as wide; lateral margins with 12–15 close-set spinules. Chelipeds held directed straight forwards; carpus twice as long as wide. Pereopods 2–4 dactyli terminating in simple claw. Maximum cl. 4 mm. Intertidal, subtidal (0–9 m; commensal with gorgonian corals). 1 species (Glassell 1936: figure of type species; Haig 1960: diagnosis).

Novorostrum Osawa, 1998

Pachycheles Stimpson, 1858

Diagnosis. Carapace ovate, up to ~1.3 times as long as wide; lateral margins unarmed behind anterior branchial spine; dorsal surface gastric region strongly elevated, front strongly produced, broad, weakly trilobed; pterygostomial flap a single, undivided plate, extending posteriorly as far as pereopod 4. Antennal peduncle with movable articles freely accessible to orbit. Chelipeds subequal. Pereopods 2–4 subcylindrical. Maximum cl. 15 mm (Pl. 27h). Intertidal, subtidal (0–20 m; associated with large sea anemones). Indo-West Pacific. 3 species (Miyake 1937: diagnosis; Osawa and Fujita 2001: description of third species).

Diagnosis. Carapace rounded or subquadrate, about as long as wide or slightly wider than long; front not prominent, appearing sinuous to transverse or rounded in dorsal view; pterygostomial flap composed of 1 or more pieces separated from anterior part by membranous interspaces. Antennal peduncle with movable articles freely accessible to orbit. Chelipeds thick, robust, distinctly unequal. Telson of 5 or 7 plates. Maximum cl. 18 mm (Pl. 27k, l). Intertidal–shelf (0–180 m). Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific. 48 species (Haig 1960: rediagnosis, key to 18 E Pacific species; Miranda and Mantelatto 2016: molecular relationships suggest two groups, one with setose carapace and chelipeds, the other without setae; Osawa 2007: key to 13 Indo-West Pacific species; Werding and Hiller 2017: figures).

Neopisosoma Haig, 1960

Parapetrolisthes Haig, 1962

Diagnosis. Carapace subquadrate to subcircular, as long as wide or slightly wider than long; front not prominent, appearing broadly rounded in dorsal view; pterygostomial flap composed of single calcified anterior plate extending approximately to level of pereopod 2 followed posteriorly by entirely membranous portion.

Diagnosis. Carapace cordiform, slightly wider than long; lateral margins spinose; dorsal surface weakly convex, with numerous transverse striae, regions weakly defined, front weakly to strongly produced, obtusely triangular, spinose; pterygostomial flap a single, undivided plate, extending posteriorly as far as

Neopetrolisthes Miyake, 1937

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

pereopod 4. Antennal peduncle with movable articles freely accessible to orbit. Chelipeds unequal, slender; fingers distinctly gaping, tips hollowed, scoop-like. Pereopods 2–4 flattened. Maximum cl. 7 mm. Intertidal–shelf (0–54 m). Tropical W Atlantic. 1 species (Glassell 1945: figure of type species; Haig 1962: diagnosis; Rodríguez et al. 2006: comment on relationships; Werding 1982: figure).

Petrocheles Miers, 1876 Diagnosis. Carapace cordiform, about as long as wide; lateral margins spinose; front projecting, dentate; pterygostomial flap a single, undivided plate, extending posteriorly as far as pereopod 4. Antennal peduncle with movable articles freely accessible to orbit. Chelipeds subequal; carpus anterior margin and upper surface spinose. Telson of 5 plates. Maximum cl. 23 mm. Intertidal–shelf (0–100 m). Temperate Australasia. 2 species (McLay 1988: New Zealand species; Poore 2004: Australian species).

Petrolisthes Stimpson, 1858 Diagnosis. Carapace cordiform or subcircular, about as long as wide; lateral margins unarmed; dorsal surface flattish or weakly convex, surface evenly ornamented, regions undefined or weakly defined, front weakly to strongly produced, broad, sinuously triangular to trilobed, at most with shallow longitudinal groove, without pair of distal elevations; pterygostomial flap a single, undivided plate, extending posteriorly as far as pereopod 4. Antennal peduncle with movable articles freely accessible to orbit. Eyestalk as wide as cornea in dorsal view. Chelipeds subequal; fingers without gape, occluding along length of opposable margin, tips simple. Pereopods 2–4 flattened. Maximum cl. 33 mm (Pl. 28a–e). Intertidal–shelf (0–183 m). Cosmopolitan. 113 species (Osawa 2007: key to 14 species from New Caledonia; Osawa and McLaughlin 2010: rediagnosis, key to 19 species from Taiwan). Petrolisthes elongatus (H. Milne Edwards, 1837) is an exotic species in Tasmania, Australia (Poore 2004).

Pisidia Leach, 1820 Diagnosis. Carapace rounded or ovate, as long as wide or slightly longer than wide; lateral margins curved, with 1 or more spines behind epibranchial angle; dorsal surface smooth or areolate, front horizontal, in dorsal view appearing distinctly trilobed with median lobe entire, without median notch, or broadly angular to weakly sinuous (P. dispar); exorbital angle usually forming spine (obtuse, blunt in P. dispar); hepatic spine usually present. Antennal peduncle with movable articles excluded from orbit. Chelipeds usually unequal, more spinose in female than male. Smaller cheliped fingers often twisted out of plane with palm in large adults. Pereopods 2–4 dactyli terminating in simple claw, with small spines on flexor margin. Maximum cl. 11 mm. Intertidal–shelf (0–188 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Temperate Southern Africa, Temperate Australasia. 14 species

293

(Koukouras et al. 2002: key to 3, now 2, species of Atlantic-Mediterranean; Osawa 2007: key to 9 Indo-West Pacific species; Yang and Sun 1990: descriptions of 5 species). The rostrum and exorbital angle of Pisidia dispar (Stimpson, 1858) differ from congeners; it belongs in a separate genus (Osawa and Chan 2010). It is isolated in the key above.

Polyonyx Stimpson, 1858 Diagnosis. Carapace transversely ovate or subhexagonal, as long as wide or wider than long; lateral margins unarmed; dorsal surface with regions indistinct, without transverse ridges, front curled ventrally, appearing broadly straight to weakly sinuous in dorsal view; exorbital angle unarmed. Antennal peduncle with movable articles excluded from orbit. Chelipeds unequal. Pereopods 2–4 dactyli terminating in bi- or triunguiculate claw, with 1 or 2 small spines on flexor margin. Maximum cl. 10 mm (Pl. 28f, g). Intertidal–shelf (0–152 m; associated with corals, sponges, tubicolous polychaetes, Chaetopterus, possibly bivalve molluscs, Brechites). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Tropical Eastern Pacific, Temperate Southern Africa, Temperate Australasia. 32 species (Chace 1956: key to 3 W African species; Haig 1960: key to 3 E Pacific species, redescriptions; Osawa 2007: key to 6 species from New Caledonia; Osawa and Sato 2022: key to 19 Indo-West Pacific species of the Polyonyx sinensis Stimpson, 1858 group).

Porcellana Lamarck, 1801 Diagnosis. Carapace cordiform, slightly wider than long; lateral margins unarmed behind anterior branchial spine; dorsal surface smooth or with fine striations, front horizontal, with 3 or 5 lobes in dorsal view; hepatic spine absent. Antennal peduncle with movable articles excluded from orbit. Chelipeds subequal. Pereopods 2–4 dactyli terminating in simple claw, with small spines on flexor margin. Maximum cl. 24 mm (Pl. 28h). Intertidal–slope (0–393 m; usually free-living but P. curvifrons Yang & Sun, 1990 commensal with Pagurus sp. (Yang and Sun 1990)). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Central Indo-Pacific, Tropical Eastern Pacific. 15 species (Chace 1956: 3 W African species; Haig 1966; Lemaitre and Campos 2000: key to 4 W Atlantic species; Osawa 2007: key to 4 Indo-West Pacific species).

Porcellanella White, 1852 Diagnosis. Carapace ~1.3 times as long as wide; lateral margins unarmed; front median lobe strongly produced in advance of lateral lobes. Antennal peduncle with movable articles excluded from orbit. Chelipeds subequal. Pereopods 2–4 dactyli quadriunguiculate. Maximum cl. 12 mm (Pl. 28i). Intertidal–shelf (0–72 m; commensal with pennatulaceans). Western and Central Indo-Pacific. 2 species (Nakasone and Miyake 1972: figures; Sankarankutty 1963: species description; Sivasubramanian et al. 2014: figures).

294

Marine Decapod Crustacea

Pseudoporcellanella Sankarankutty, 1962 Diagnosis. Carapace ~1.8 times as long as wide; lateral margins unarmed. Chelipeds held directed straight forwards; carpus wider than long. Pereopods 2–4 dactyli triunguiculate. Maximum cl. 20 mm. Subtidal, shelf (5–40 m; commensal with pennatulaceans). Western and Central Indo-Pacific. 1 species (Ng and Nakasone 1994: rediagnosis, figures; Sankarankutty 1962).

Raphidopus Stimpson, 1858 Diagnosis. Carapace transversely ovate, wider than long. Antennal peduncle with movable articles excluded from orbit. Chelipeds unequal. Pereopods 2–4 dactyli straight, slender, terminating in simple sharp point, without spines on flexor margin. Maximum cl. 7 mm. Intertidal, subtidal (0–11 m; free living or commensal with tubicolous polychaetes, Loimia). Western and Central IndoPacific. 5 species (Ng and Nakasone 1994: rediagnosis, comparison of 3 species; Osawa and Ng 2018: key to species).

Ulloaia Glassell, 1938 Diagnosis. Carapace ~1.3 times as long as wide; lateral margins without anterior branchial spine, with granular lobes or blunt teeth; dorsal surface covered with rugose protuberances, front quadridentate in dorsal view, lobes in transverse row. Antennal peduncle with movable articles excluded from orbit. Chelipeds subequal. Pereopods 2–4 dactyli terminating in simple claw. Maximum cl. 4 mm. Intertidal, subtidal (0–15 m). Tropical Eastern Pacific. 1 species (Glassell 1938: figure of type species; Haig 1960: rediagnosis). References Baeza JA (2016) Molecular phylogeny of porcelain crabs (Porcellanidae: Petrolisthes and allies) from the south eastern Pacific: the genera Allopetrolisthes and Liopetrolisthes are not natural entities. PeerJ 4, e1805. doi:10.7717/peerj.1805 Bracken-Grissom HD, Cannon ME, Cabezas P, Feldmann RM, Schweitzer CE, et  al. (2013) A comprehensive and integrative reconstruction of evolutionary history for Anomura (Crustacea: Decapoda). BMC Evolutionary Biology 13, 128 doi:10.1186/1471-2148-13-128. Chace FA (1956) Porcellanid crabs. Expédition océanographique Belge dans les eaux côtières Africaines de l’Atlantique sud 1948–1949). Résultats Scientifiques 3, 1–54. Dolorosa RG, Werding B (2014) A new mangrove-inhabiting porcelain crab of the genus Enosteoides (Crustacea: Decapoda: Anomura) from Puerto Princesa Bay, Palawan, the Philippines. Bulletin of Marine Science 90, 865–872. doi:10.5343/bms.2013.1079 Dong D, Li X, Chan T-Y (2011) A new species of Aliaporcellana Nakasone and Miyake, (Crustacea: Decapoda: Anomura: Porcellanidae) from Taiwan, with redescription of Aliaporcellana suluensis (Dana, 1852). Bulletin of Marine Science 87, 485–499. doi:10.5343/bms.2010.1096 Ferreira LAdA, Tavares M (2017) A new species of Megalobrachium Stimpson, 1858 (Crustacea: Anomura: Porcellanidae) from the eastern Pacific coasts of Panama and Colombia, with re-description of

M. erosum. Journal of the Marine Biological Association of the United Kingdom 97, 1407–1415. doi:10.1017/S0025315416000709 Ferreira LAdA, Tavares M (2020) The rare porcelain crab Novorostrum decorocrus Osawa, 1998 (Anomura: Porcellanidae) from Indonesia. Nauplius 28, e2020007. doi:10.1590/2358-2936e2020007 Glassell SA (1936) New porcellanids and pinnotherids from tropical North American waters. Transactions of the San Diego Society of Natural History 8, 277–304. doi:10.5962/bhl.part.14900 Glassell SA (1938) New and obscure decapod Crustacea from the West American coasts. Transactions of the San Diego Society of Natural History 8, 411–454, pls 27–36. Glassell SA (1945) Four new species of North American crabs of the genus Petrolisthes. Journal of the Washington Academy of Sciences 35, 223–229. Haig J (1960) The Porcellanidae (Crustacea Anomura) of the eastern Pacific. Allan Hancock Pacific Expeditions 24, 1–440, pls 1–41. Haig J (1962) Papers from Dr. Th. Mortensen’s Pacific Expedition 1914– 1916. LXXIX. Porcellanid crabs from eastern and western America. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i Kjøbenhavn 124, 171–192. Haig J (1965) The Porcellanidae (Crustacea, Anomura) of Western Australia), with descriptions of four new Australian species. Journal of the Royal Society of Western Australia 48(97), 118. Haig J (1966) The Porcellanidae (Crustacea Anomura) of the Iranian Gulf and Gulf of Oman. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i Kjøbenhavn 129, 49–65. Haig J (1978) Contribution toward a revision of the porcellanid genus Porcellana (Crustacea: Decapoda: Anomura). Proceedings of the Biological Society of Washington 91, 706–714. Haig J (1981) Porcellanid crabs from the Indo-west Pacific, Part II. Steenstrupia (Copenhagen) 7, 269–291. Harvey AW (1999) A review of the genus Clastotoechus Haig, with descriptions of a new genus and two new species (Decapoda, Anomura, Porcellanidae). American Museum Novitates 3255, 1–32. Hiller A, Werding B (2018) On a new commensal species of Aliaporcellana from the western Pacific (Crustacea, Decapoda, Porcellanidae). ZooKeys 780, 1–9. doi:10.3897/zookeys.780.26388 Johnson DS (1970) The Galatheidae (Crustacea: Decapoda) of Singapore and adjacent waters. Bulletin of the National Museum (Singapore) 35, 1–44. Koukouras A, Mavidis M, Noël PY (2002) The genus Pisidia Leach (Decapoda, Anomura) in the Northeastern Atlantic Ocean and the Mediterranean Sea.Crustaceana75, 451–463.doi:10.1163/156854002760095516 Lemaitre R, Campos NH (2000) Porcellana lillyae, new species (Decapoda, Anomura, Porcellanidae), from the Caribbean Sea, with a key to the Western Atlantic species of the genus. Journal of Crustacean Biology 20, 259–265. doi:10.1163/1937240X-90000028 McLay CL (1988) Brachyura and crab-like Anomura of New Zealand [Crabs of New Zealand]. Leigh Laboratory Bulletin 22, 1–463. Miranda I, Mantelatto FL (2016) Porcelain crabs of the genera Pachycheles Stimpson and Neopisosoma Haig (Decapoda : Anomura : Porcellanidae): new premises based on molecular data and comments on phylogenetic relationships in the family. Invertebrate Systematics 30, 509–520. doi:10.1071/IS16025 Miyake S (1937) A new crab-shaped anomuran living commensally with a gigantic sea-anemone (Neopetrolisthes ohshimai gen. et sp. nov.). Zoological Magazine, Tokyo 49, 34–36. Miyake S (1978) The crustacean Anomura of Sagami Bay collected by His Majesty the Emperor of Japan. Biological Laboratory, Imperial Household, Tokyo. Nakasone Y, Miyake S (1972) Four unrecorded porcellanid crabs (Anomura: Porcellanidae) from Japan. Bulletin of Science & Engineering Division,

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

University of the Ryukyus. Mathematics and Natural Sciences 15, 136–147. Ng PKL, Nakasone Y (1993) Taxonomy and ecology of the porcellanid crab Polyonyx cometes Walker, 1887 (Crustacea: Decapoda), with description of a new genus. Journal of Natural History 27, 1103–1117. doi:10.1080/00222939300770681 Ng PKL, Nakasone Y (1994) On the porcellanid genera Raphidopus Stimpson, 1858, and Pseudoporcellanella Sankarankutty, 1961, with description of a new mangrove species, Raphidopus johnsoni from Singapore (Decapoda, Anomura). Crustaceana 66, 1–21. doi:10.1163/156854094X00116 Osawa M (1998) Novorostrum, new genus (Decapoda: Anomura: Porcellanidae), with descriptions of three related species. Journal of Crustacean Biology 18, 161–176. doi:10.2307/1549531 Osawa M (2001) Heteropolyonyx biforma new genus and species, from Japan, and redescription of Polyonyx utinomii (Decapoda: Porcellanidae). Journal of Crustacean Biology 21, 506–520. doi:10.1163/2002197599990152 Osawa M (2007) Porcellanidae (Crustacea: Decapoda: Anomura) from New Caledonia and the Loyalty Islands. Zootaxa 1548, 1–49. doi:10.11646/zootaxa.1548.1.1 Osawa M (2016) Enosteoides Johnson, 1970 (Crustacea: Decapoda: Anomura: Porcellanidae) from the Central Philippines, with description of a new species of the genus. Nauplius 24, e2016013. doi:10.1590/​ 2358-2936e2016013 Osawa M, Chan T-Y (2010) Part III. Porcellanidae (porcelain crabs). In Crustacean fauna of Taiwan: Crab-like anomurans (Hippoidea, Lithodoidea and Porcellanidae). (Ed. Chan TY) pp. 67–181. National Taiwan Ocean University, Keelung. Osawa M, Fujita Y (2001) A new species of the genus Neopetrolisthes Miyake 1937 (Crustacea: Decapoda: Porcellanidae) from the Ryukyu Islands, southwestern Japan. Proceedings of the Biological Society of Washington 114, 162–171. Osawa M, McLaughlin PA (2010) Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea) Part II – Porcellanidae. Raffles Bulletin of Zoology Supplement 23, 109–129. Osawa M, Ng PKL (2018) A new species of the genus Raphidopus Stimpson, 1858 (Crustacea: Decapoda: Anomura: Porcellanidae) from Peninsular Malaysia, with additional records of R. johnsoni Ng & Nakasone, 1994 from Southeast Asia, and a key to species in the genus. Zootaxa 4433, 111–126. doi:10.11646/zootaxa.4433.1.6 Osawa M, Sato T (2022) A distinctive new species of the genus Polyonyx Stimpson, 1858 (Crustacea: Decapoda: Anomura: Porcellanidae) from Okinawa, southwestern Japan. Zootaxa 5091, 587–597. doi:10.11646/ zootaxa.5091.4.6 Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Rodríguez IT, Hernández G, Felder DL (2005) Review of the Western Atlantic Porcellanidae (Crustacea: Decapoda: Anomura) with new records, systematic observations, and comments on biogeography. Caribbean Journal of Science 41, 44–582. Rodríguez IT, Hernández G, Felder DL (2006) Phylogenetic relationships among western Atlantic Porcellanidae (Decapoda: Anomura), based on partial sequences of the mitochondrial 16S rRNA gene, with comments on morphology. Crustacean Research Special Number 6, 151–166. doi:10.18353/crustacea.Special2006.6_151 Sankarankutty C (1962) On a new genus of Porcellanidae (Crustacea  – Anomura). Journal of the Marine Biological Association of India 3, 92–95. Sankarankutty KN (1963) On three species of porcellanids (Crustacea Anomura) from the Gulf of Mannar. Journal of the Marine Biological Association of India 5, 273–279.

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Sankolli KN (1963) On a new species of porcellanid crab (Decapoda, Anomura) from India. Journal of the Marine Biological Association of India 5, 280–283. Sivasubramanian K, Ravichandran S, Anbuchezhian R (2014) First discovery of porcellanid crab, Porcellanella picta (Crustacea: Decapoda: Porcellanidae), from south-east coast of India. Journal of Asia-Pacific Biodiversity 7, 248–251. doi:10.1016/j.japb.2014.05.001 Stillman JH, Reeb CA (2001) Molecular phylogeny of eastern Pacific porcelain crabs, genera Petrolisthes and Pachycheles, based on the mtDNA 16S rDNA sequence: phylogeographic and systematic implications. Molecular Phylogenetics and Evolution 19, 236–245. doi:10.1006/mpev.2001.0924 Tirmizi NM, Yaqoob M, Siddiqui FA (1989) Marine Fauna of Pakistan: 3. Porcellanid crabs (Crustacea, Anomura). Centre of Excellence in Marine Biology, University of Karachi, Karachi. Trivedi JN, Osawa M, Vachhrajani KD (2017) A new species of the genus Ancylocheles Haig, 1978 (Crustacea: Decapoda: Anomura: Porcellanidae) from Gujarat, northwestern India. Zootaxa 4299, 384–390. doi:10.11646/zootaxa.4299.3.4 Vela MJ, González-Gordillo JI (2016) Larval descriptions of the family Porcellanidae: a worldwide annotated compilation of the literature (Crustacea, Decapoda). ZooKeys 564, 47–70. doi:10.3897/zookeys.564.7018 Werding B (1982) Porcellanid crabs of the Islas del Rosario, Caribbean coast of Colombia, with a description of Petrolisthes rosariensis new species (Crustacea: Anomura). Bulletin of Marine Science 32, 439–447. Werding B (1986) Die Gattung Neopisosoma Haig, 1960 im tropischen Westatlantik, mit der Beschreibung von Neopisosoma neglectum spec. nov. und Neopisosoma orientale spec. nov. (Crustacea: Anomura: Porcellanidae). Zoölogische Mededeelingen 16, 159–179. Werding B, Hiller A (2017) Description of a new species of Pachycheles (Decapoda, Anomura, Porcellanidae) from the southern Caribbean Sea. Crustaceana 90, 1279–1288. doi:10.1163/15685403-00003684 Werding B, Hiller A, Misof B (2001) Evidence of paraphyly in the neotropical porcellanid genus Neopisosoma (Crustacea: Anomura: Porcellanidae) based on molecular characters. Hydrobiologia 449, 105–110. doi:10.1023/A:1017545120296 Yang S, Sun X (1990) On the porcellanid crabs from the coast of Fujian Province, China. Memoirs of Beijing Natural History Museum 45, 1–15[in Chinese].

Hippoidea Latreille, 1825 sand crabs, mole crabs Hippoids are burrowing crabs living in sandy habitats, predominantly from the tropics and subtropics. Many species are abundant on beaches and accessible for study  – their ecology is well studied (e.g. Lastra et al. 2016; Petracco et al. 2017). Their longitudinally streamlined body and flattened walking legs are key adaptations allowing them to rapidly burrow backwards into the substrate. The taxonomy of all families, genera and species of ­A lbuneidae and Blepharipodidae was documented in detail by (Boyko 2002). Boyko and Harvey (2009) reviewed this study adding an analysis of phylogeny and biogeography. Few species have been added since the compilation of species by Boyko and McLaughlin (2010). Most phylogenetic studies place the hippoids as the sister group to the remaining anomurans (Ahyong et al. 2009; Bracken-Grissom et al. 2013; Tsang et al. 2011).

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Diagnosis. Sternal plastron consisting of thoracic sternites 3–7, thoracic sternite 8 independent. Pleon symmetrical; pleonites separate, calcified and distinct. Ocular scales absent. Antennal peduncle of 5 articulating articles.

Pereopods 1 symmetrical, simple or flat, subchelate. Pereopods 2–4 with flattened dactyli. Pereopod 4 ambulatory, like 2 and 3. Uropods narrow, symmetrical. Telson longer than wide, not forming tail-fan with uropods.

Key to families of Hippoidea 1. – 2. –

Pereopod 1 dactylus simple (Fig. 13.15d–f)������������������������������������������������������������������������������������������������������������������� Hippidae Pereopod 1 dactylus chelate (Figs 13.13s, 13.14c)������������������������������������������������������������������������������������������������������������������������2 Pereopod 1 propodal finger spinose (Fig. 13.14c). Gills trichobranch (Fig. 13.14d)���������������������������������� Blepharipodidae Pereopod 1 propodal finger smooth or with blunt teeth (Fig. 13.13s). Gills phyllobranch (Fig. 13.13t)���������� Albuneidae

References Ahyong ST, Schnabel KE, Maas EW (2009) Anomuran phylogeny: new insights from molecular data. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 399–414. CRC Press, Boca Raton. Boyko CB (2002) A worldwide revision of the Recent and fossil sand crabs of the Albuneidae Stimpson and Blepharipodidae, new family (Crustacea: Decapoda: Anomura: Hippoidea). Bulletin of the American Museum of Natural History 272, 1–396. doi:10.1206/00030090(2002)2722.0.CO;2 Boyko CB, Harvey AW (2009) Phylogenetic systematics and biogeography of the sand crab families Albuneidae and Blepharipodidae (Crustacea: Anomura: Hippoidea). Invertebrate Systematics 23, 1–18. doi:10.1071/ IS06053 Boyko CB, McLaughlin PA (2010) Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea) Part IV – Hippoidea. Raffles Bulletin of Zoology Supplement 23, 139–151. Bracken-Grissom HD, Cannon ME, Cabezas P, Feldmann RM, Schweitzer CE, et  al. (2013) A comprehensive and integrative reconstruction of evolutionary history for Anomura (Crustacea: Decapoda). BMC Evolutionary Biology 13, 128 doi:10.1186/1471-2148-13-128. Lastra M, López J, Troncoso J, Hubbard DM, Dugan JE (2016) Scavenger and burrowing features of Hippa pacifica (Dana 1852) on a range of tropical sandy beaches. Marine Biology 163, 212. doi:10.1007/ s00227-016-2985-9 Petracco M, Cardoso RS, Martinelli Filho JE, Turra A (2017) Effects of beach morphodynamic features on production and P/B ratio of the crab Emerita brasiliensis Schmitt, 1935 (Decapoda: Hippidae) in sandy beaches of South America. Journal of the Marine Biological Association of the United Kingdom 97, 1215–1221. doi:10.1017/S0025315416000576

Tsang LM, Chan T-Y, Ahyong ST, Chu KH (2011) Hermit to king, or hermit to all: multiple transitions to crab-like forms from hermit crab ancestors. Systematic Biology 60, 616–629. doi:10.1093/sysbio/syr063

Albuneidae Stimpson, 1858 mole crabs, sand crabs Figure 13.13, Plate 29a–e Mole crabs or sand crabs are obligate diggers in fine sandy beaches and similar subtidal sediments. They conceal themselves completely in sand to be invisible from the surface (Faulkes 2017). Boyko (2002) reviewed the taxonomy of Albuneidae in detail, recognising two subfamilies, Albuneinae and Lepidopinae. Two genera, however, were treated as incertae sedis in a later review of the family phylogeny and biogeography (Boyko and Harvey 2009). Boyko (2010) updated taxonomic and distributional data. Diagnosis. Hippoidea. Pereopod 1 chelate, propodal finger smooth or with blunt teeth (Fig.  13.13s). Gills phyllobranchiate (Fig. 13.13t). Implicit generic attributes. Carapace branchiostegite with spine. Distal ocular peduncular segment flat, much shorter than carapace. Antennal article 1 without dorsodistal spine; flagellum with more than 2 articles. Maxilliped 2 exopod with flagellum.

Key to genera of Albuneidae 1. – 2. – 3. – 4. –

Pleonite 5 with pleura (Fig. 13.13j). Antennal acicle short (shorter than article 3) (Fig. 13.13n, o)�������������������������������������2 Pleonite 5 without pleura (Fig. 13.13i). Antennal acicle long (longer or equal to article 3) (Fig. 13.13m, p)����������������������5 Antenna with 3 flagellar articles (Fig. 13.13n)����������������������������������������������������������������������������������������������������� Leucolepidopa Antenna with more than 3 flagellar articles (Fig. 13.13o)����������������������������������������������������������������������������������������������������������3 Rostral area anterior margin truncate. Eyes narrow, separated by eye-width (Fig. 13.13b)����������������������������� Austrolepidopa Rostral area anterior margin rounded or concave. Eyes about as wide as long, separated by less than eye-width (Fig. 13.13c, e)������������������������������������������������������������������������������������������������������������������������������������������������������������4 Antennular dorsal flagellum with ~60 articles. Maxilliped 2 exopod without flagellum (Fig. 13.13r)�������� Paraleucolepidopa Antennular dorsal flagellum with 80–250 articles. Maxilliped 2 exopod with flagellum (Fig. 13.13q)�������������� Lepidopa

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

297

Fig. 13.13.  Albuneidae. Dorsal carapace, or anterior margin, with ocular peduncles: a, Albunea; b, Austrolepidopa; c, Lepidopa; d, Paralbunea; e, Paraleucolepidopa; f, Squillalbunea; g, Stemonopa; h, Zygopa. Pleon: i, Albunea; j, Paraleucolepidopa. Telson: k, Squillalbunea; l, Stemonopa. Antenna, lateral view: m, Albunea; n, Leucolepidopa; o, Paraleucolepidopa; p, Squillalbunea. Maxilliped 2: q, Lepidopa; r, Paraleucolepidopa. Pereopod 1: s, Albunea. t, phyllobranch gill.

5. – 6. – 7. – 8. –

Ocular peduncles reduced, fused. Carapace branchiostegite without spine (not visible in dorsal view) (Fig. 13.13h)����������������������������������������������������������������������������������������������������������������������������������������������������������������������� Zygopa Ocular peduncles normal, separate. Carapace branchiostegite with spine (Fig. 13.13a, d–g)���������������������������������������������6 Antennal article 1 with dorsodistal spine (Fig. 13.13m)������������������������������������������������������������������������������������������������Albunea Antennal article 1 without dorsodistal spine (Fig. 13.13p)��������������������������������������������������������������������������������������������������������7 Telson triangular (Fig. 13.13k). Carapace covered in setose lines (Fig. 13.13f)����������������������������������������������� Squillalbunea Telson ovate (Fig. 13.13l). Carapace setal lines restricted to major grooves (Fig. 13.13d, g)�������������������������������������������������8 Eyes longer than carapace (Fig. 13.13g)���������������������������������������������������������������������������������������������������������������������� Stemonopa Eyes shorter than carapace (Fig. 13.13d)�������������������������������������������������������������������������������������������������������������������� Paralbunea Albunea Weber, 1795

Diagnosis. Carapace with setal lines restricted to grooves. Pleonite 5 without pleura. Telson triangular. Antennal article 1 with dorsodistal spine; acicle long (equal or greater than length of antennal segment 3). Maximum cl. 27 mm (Pl. 29a, b). Intertidal–shelf (0–152 m). Temperate Northern and Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific. 20 species (Boyko 2002: key to species).

Austrolepidopa Efford & Haig, 1968 Diagnosis. Rostral area anterior margin truncate. Pleonite 5 with pleura. Eyes narrow, separated by eye length-width. Antennular

dorsal flagellum with 50–90 articles. Antenna with more than 3 flagellar articles; antennal acicle short (less than length of antennal segment 3). Maximum cl. 13.4 mm (Pl. 29c). Subtidal–slope (11–225 m). Central Indo-Pacific, Temperate Australasia. 3 species (Boyko 2002: key to species; Efford and Haig 1968: genus, descriptions of two species).

Paralbunea Serène, 1977 Diagnosis. Carapace with setal lines restricted to grooves; rostral area concave. Pleonite 5 with pleura. Telson ovate. Eyes as wide as long, contiguous. Antennular dorsal flagellum with 70–145 articles. Antenna with more than 3 flagellar articles; antennal acicle

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short (less than length of antennal segment 3). Maximum cl. 15 mm (Pl. 29e). Subtidal–shelf (5–82 m). Tropical Atlantic, Indo-West Pacific, 5 species (Boyko 2002: key to species; Osawa and Fujita 2012: fifth species).

Paraleucolepidopa Calado, 1996 Diagnosis. Rostral area anterior margin rounded. Pleonite 5 with pleura. Eyes as wide as long, separated by less than eye-width. Antennular dorsal flagellum with ~60 articles. Antenna with more than 3 flagellar articles; antennal acicle short (less than length of antennal segment 3). Maxilliped 2 exopod without flagellum. Maximum cl. 9.2 mm. Subtidal, shelf (1–73 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Boyko 2002: key to species).

Lepidopa Stimpson, 1858 Diagnosis. Rostral area anterior margin rounded. Pleonite 5 with pleura. Eyes as wide as long, separated by less than eyewidth. Antennular dorsal f lagellum with 80–250 articles. Antenna with more than 3 f lagellar articles; antennal acicle short (less than length of antennal segment 3). Maximum cl. 36 mm (Pl. 29d). Intertidal–shelf (0–139 m). Temperate Northern and Tropical Atlantic, Tropical Eastern Pacific, Temperate South America (Chile). 15 species (Boyko 2002: key to 14 species; Ortiz et al. 2018: key to 7 species from Gulf of Mexico and Caribbean).

Leucolepidopa Efford, 1969 Diagnosis. Rostral area anterior margin rounded. Pleonite 5 with pleura. Eyes narrow, separated by twice eye-width. Antennular dorsal flagellum with more than 20 articles. Antenna with 3 flagellar articles; antennal flagellum with 3 articles; antennal acicle short (less than length of antennal segment 3). Maxilliped 2 exopod without flagellum. Maximum cl. 7.2 mm. Shelf (40–76 m). Central Indo-Pacific. 1 species (Efford 1969).

Squillalbunea Boyko, 2002 Diagnosis. Carapace covered in setose lines. Pleonite 5 without pleura. Telson triangular. Antennal acicle long (equal or greater than length of antennal segment 3). Maximum cl. 22 mm. Shelf (27–82 m). Central Indo-Pacific. 1 species (Boyko 2002).

Stemonopa Efford & Haig, 1968 Diagnosis. Carapace with setal lines restricted to grooves. Pleonite 5 without pleura. Telson ovate. Eyes cylindrical, longer than carapace. Antennal article 1 without dorsodistal spine; acicle long (equal or greater than length of antennal segment 3). Maximum cl. 10.4 mm. Shelf (44–83 m). Central Indo-Pacific. 1 species (Efford and Haig 1968).

Zygopa Holthuis, 1961 Diagnosis. Carapace with setal lines restricted to grooves; branchiostegite without spine. Pleonite 5 without pleura. Telson triangular. Antennal acicle long (equal or greater than length of antennal segment 3). Maximum cl. 11.5 mm. Subtidal, shelf (4–73 m). Tropical W Atlantic, Central IndoPacific. 3 species (Ortiz 2015: key to species). References Boyko CB (2002) A worldwide revision of the Recent and fossil sand crabs of the Albuneidae Stimpson and Blepharipodidae, new family (Crustacea: Decapoda: Anomura: Hippoidea). Bulletin of the American Museum of Natural History 272, 1–396. doi:10.1206/0003-0090(2002)272​2.0.CO;2 Boyko CB (2010) New records and taxonomic data for 14 species of sand crabs (Crustacea: Anomura: Albuneidae) from localities worldwide. Zootaxa 2555, 49–61. doi:10.11646/zootaxa.2555.1.3 Boyko CB, Harvey AW (2009) Phylogenetic systematics and biogeography of the sand crab families Albuneidae and Blepharipodidae (Crustacea: Anomura: Hippoidea). Invertebrate Systematics 23, 1–18. doi:10.1071/ IS06053 Efford IE (1969) Leucolepidopa sunda gen. nov., sp. nov. (Decapoda: Albuneidae), a new Indo-Pacific sand crab. Breviora 318, 1–9. Efford IE, Haig J (1968) Two new genera and three new species of crabs (Decapoda: Anomura: Albuneidae) from Australia. Australian Journal of Zoology 16, 897–914. doi:10.1071/ZO9680897 Faulkes Z (2017) The phenology of sand crabs, Lepidopa benedicti (Decapoda: Albuneidae). Journal of Coastal Research 33, 1095–1101. doi:10.2112/JCOASTRES-D-16-00125.1 Ortiz M (2015) Una nueva especie de cangrejo topo ciego del género Zygopa Holthuis, 1961 (Crustacea, Decapoda, Anomura) de Cuba. Novitates Caribaea 8, 82–94. doi:10.33800/nc.v0i8.45 Ortiz MJ, Tello-Musi L, Morales-Moreno A (2018) Especie nueva de cangrejo de arena del género Lepidopa en Veracruz, México, y una clave illustrada de las especies citadas para el Golfo de México el Mar Caribe. Novitates Caribaea 12, 49–62. doi:10.33800/nc.v0i12.84 Osawa M, Fujita Y (2012) New records of Albuneidae (Decapoda, Anomura) from Japan, with description of a new species of Paralbunea. In: Komatsu H, Okuno I, Fukuoka K (eds) Studies in Malacostraca: a homage to Masatsune Takeda. Crustaceana Monographs 17, 245–262.

Blepharipodidae Boyko, 2002 spiny mole crabs Figure 13.14, Plate 29f Blepharipodids include the largest of the hippoids, and like albuneids and hippids, dig in sandy beaches and similar subtidal sediments (Dugan et al. 2000). One species, Blepharipoda liberata Shen, 1949 is a commercially valuable seafood species popular in Shandong Province, China (Zhou et  al. 2019). The family was introduced by Boyko (2002) who redescribed its genera and species, and provided keys to distinguish them. Phylogeny and biogeography were reviewed by Boyko and Harvey (2009). Diagnosis. Pereopod 1 chelate, fixed finger spinose (Fig. 13.14c). Gills trichobranchiate (Fig. 13.14d).

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

299

Subtidal, shelf (5–68 m). Temperate Northern W Pacific, Tropical Eastern Pacific. 2 species (Boyko 2002: rediagnosis, key to species). References

Fig. 13.14.  Blepharipodidae. Dorsal carapace, with ocular peduncles: a, Blepharipoda; b, Lophomastix. c, pereopod 1, Blepharipoda. d, trichobranch gill.

Blepharipoda Randall, 1840 Diagnosis. Carapace with 2 hepatic lateral spines. Ocular peduncle distal segment subdivided into 2 pseudosegments (Fig. 13.14a). Maximum cl. 70 mm (Pl. 29f). Intertidal–shelf (0–90 m). Temperate Northern W Pacific, Tropical Eastern Pacific, Temperate South America. 4 species (Boyko 2002: rediagnosis, key to species).

Lophomastix Benedict, 1904 Diagnosis. Carapace with 1 hepatic lateral spine. Ocular peduncle distal segment entire (Fig. 13.14b). Maximum cl. 33 mm.

Boyko CB (2002) A worldwide revision of the Recent and fossil sand crabs of the Albuneidae Stimpson and Blepharipodidae, new family (Crustacea: Decapoda: Anomura: Hippoidea). Bulletin of the American Museum of Natural History 272, 1–396. doi:10.1206/0003-0090(2002)2722.0.CO;2 Boyko CB, Harvey AW (2009) Phylogenetic systematics and biogeography of the sand crab families Albuneidae and Blepharipodidae (Crustacea: Anomura: Hippoidea). Invertebrate Systematics 23, 1–18. doi:10.1071/ IS06053 Dugan JE, Hubbard DM, Lastra M (2000) Burrowing abilities and swash behavior of three crabs, Emerita analoga Stimpson, Blepharipoda occidentalis Randall, and Lepidopa californica Efford (Anomura, Hippoidea), of exposed sandy beaches. Journal of Experimental Marine Biology and Ecology 255, 229–245. doi:10.1016/S0022-0981(00)00294-X Zhou L, Wang X, Wu B, Sun X, Zhao Q, et al. (2019) Karyological analysis of the sea cicada Blepharipoda liberate Shen from the Rizhao intertidal zone, China. Journal of Oceanology and Limnology 37, 169–175. doi:10.1007/s00343-019-7320-0

Hippidae Latreille, 1825 mole crabs Figure 13.15, Plate 30a–c Mole crabs have compact, bullet-shaped bodies and live buried in sandy substrate or fine sediments in the intertidal to subtidal zone (Defeo and Cardoso 2002). The family was rediagnosed by (Osawa et al. 2010) who also distinguished the genera before redescribing species from Taiwan. Boyko and Harvey (1999) provided keys to Indo-West Pacific species. Diagnosis. Hippoidea. Pereopod 1 simple (Fig. 13.15d–f).

Key to genera of Hippidae 1. – 2. –

Antennal flagellum at least as long as carapace, densely covered ventrally with long plumose setae. Pereopod 1 dactylus lamellate, oval-triangular (Fig. 13.15d)�������������������������������������������������������������������������������������������� Emerita Antennal flagellum considerably shorter than carapace. Pereopod 1 dactylus subcylindrical�������������������������������������������2 Pereopod 1 dactylus l less than third length of carapace, non-articulated (Fig. 13.15e)����������������������������������������������Hippa Pereopod 1 dactylus l nearly as long as carapace, multiarticulate (Fig. 13.15f)�����������������������������������������������Mastigochirus Emerita Scopoli, 1777

Hippa Fabricius, 1787

Diagnosis. Antennal flagellum at least as long as carapace, densely covered ventrally with long plumose setae. Pereopod 1 dactylus lamellate, oval-triangular. Maximum cl. 36 mm (Pl. 30a). Intertidal, subtidal (0–13 m). Temperate and Tropical W Atlantic, Temperate Northern W Pacific, Indo-West Pacific, Temperate South America. 9 species (Haye et al. 2002: phylogenetics; Hsueh 2015: key to species; Tam et al. 1996: zoogeography).

Diagnosis. Antennal flagellum considerably shorter than carapace. Pereopod 1 dactylus subcylindrical, slender, less than third length of carapace, non-articulated. Maximum cl. 35 mm (Pl. 30c). Intertidal, subtidal (0–2 m). Tropical W Atlantic, Indo-West Pacific, Tropical Eastern Pacific, Temperate Australia. 15 species (Haig et  al. 1986: key to 6 species of one of two groups; Osawa et al. 2010: key to 5 species from Taiwan).

300

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Fig. 13.15.  Hippidae. Dorsal carapace, with ocular peduncles: a, Emerita; b, Hippa; c, Mastigochirus. Pereopod 1, propodus, dactylus: d, Emerita; e, Hippa; f, Mastigochirus (all articles).

Mastigochirus Miers, 1878

Lomisoidea Bouvier, 1895

Diagnosis. Antennal flagellum considerably shorter than carapace. Pereopod 1 dactylus subcylindrical, elongate, nearly as long as carapace, multiarticulate. Maximum cl. 13 mm (Pl. 30b). Subtidal–shelf (13–37 m). Western and Central Indo-Pacific. 2 species (Haig 1974: redescribed M. quadrilobatus Miers, 1878; Osawa et al. 2010).

Lomisoidea is a superfamily with one family, one genus and one species. Lomisoids are crab-like and represent one of the several instances of evolution of crab-like form, ‘carcinisation’, in the Anomura. Despite having the habitus of a crab, there seems little doubt that the ‘hairy stone crab’ diverged from a squat lobster-like ancestor (Bracken-Grissom et  al. 2013); McLaughlin and Lemaitre (1997) and Ahyong et  al. (2011) reviewed the history of phylogenetic considerations of Lomis among the squat lobsters and other anomurans. Diagnosis. Sternal plastron consisting of thoracic sternites 3–7, thoracic sternite 8 independent. Pleon symmetrical; pleonites separate, calcified and distinct. Ocular scales absent. Antennal peduncle of 5 articulating articles. Pereopods 1 symmetrical, flat, chelate. Pereopods 2–4 with claw-like dactyli. Pereopod 2–4 ambulatory, similar; pereopod 5 vestigial. Uropods absent (or invisible in natural view and pleopod-like only in female). Telson undivided, symmetrical.

References Boyko CB, Harvey AW (1999) Crustacea Decapoda: Albuneidae and Hippidae of the tropical Indo-West Pacific region. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 20. Mémoires du Muséum National d’Histoire Naturelle, Paris 180, 379–406. Defeo O, Cardoso RS (2002) Macroecology of population dynamics and life history traits of the mole crab Emerita brasiliensis in Atlantic sandy beaches of South America. Marine Ecology Progress Series 239, 169–179. doi:10.3354/meps239169 Haig J (1974) A review of the Australian crabs of family Hippidae (Crustacea, Decapoda, Anomura). Memoirs of the Queensland Museum 17, 175–189. Haig J, Murugan T, Nair NB (1986) Hippa indica, a new species of mole crab (Decapoda, Anomura, Hippidae) from the south west coast of India. Crustaceana 51, 286–292. doi:10.1163/156854086X00449 Haye PA, Tam YK, Kornfield I (2002) Molecular phylogenetics of mole crabs (Hippidae: Emerita). Journal of Crustacean Biology 22, 903–915. doi:10.1163/20021975-99990302 Hsueh P-W (2015) A new species of Emerita (Decapoda, Anomura, Hippidae) from Taiwan, with a key to species of the genus. Crustaceana 88, 247–258. doi:10.1163/15685403-00003413 Osawa M, Boyko CB, Chan T-Y (2010) Part I. Hippoidea (mole crabs). In Crustacean fauna of Taiwan: Crab-like anomurans (Hippoidea, Lithodoidea and Porcellanidae). (Ed. Chan TY) pp. 1–41. National Taiwan Ocean University, Keelung. Tam YK, Kornfield I, Ojeda FP (1996) Divergence and zoogeography of mole crabs, Emerita spp. (Decapoda: Hippidae), in the Americas. Marine Biology 125, 489–497.

Lomisidae Bouvier, 1895 Figure 13.16, Plate 30d The family is monotypic and confined to southern ­Australia. The species’ systematic position has long been enigmatic and after a history of being shifted from one anomuran group to another was thought to be so different from all others that it deserves its own superfamily. McLaughlin (1983) and Tudge (1997) discussed the matter and decided on the classification adopted here (BrackenGrissom et al. 2013).

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

301

Tudge CC (1997) Spermatological evidence supports the taxonomic placement of the Australian endemic hairy stone crab, Lomis hirta (Decapoda: Anomura: Lomidae). Memoirs of the Museum of Victoria 56, 235–244. doi:10.24199/j.mmv.1997.56.09

Paguroidea Latreille, 1802 hermit crabs, king crabs, coconut crabs

Fig. 13.16.  Lomisidae. Lomis hirta (Lamarck, 1818).

Lomis H. Milne Edwards, 1837 hairy stone crab Maximum cl. 16 mm (Pl. 30d). Temperate southern Australia (N to Shark Bay). The hairy stone crab Lomis hirta (Lamarck, 1818) is an unusual false crab common under rocks in the lower intertidal zone of exposed rocky shores of southern Australia. It is best recognised by its flattened form, grey hairy and muddy appearance, with legs and chelipeds so well fitting together that it clings closely to its rocky surface. The antennae are brilliant blue. Lomis, unlike most true brachyuran crabs, possesses vestigial fifth pereopods. It resembles porcellanid crabs more closely but differs in the reduced uropods, presence of female first pleopods, and eyes contiguous and flattened. The first stage zoea larva was described by Cormie (1993) who suspected it may have an abbreviated larval life. References Ahyong ST, Schnabel KE, Macpherson E (2011) Chapter 3. Phylogeny and fossil record of the marine squat lobsters. In The biology of squat lobsters. (Eds Poore GCB, Ahyong ST, Taylor J) pp. 73–104. CSIRO Publishing, Melbourne (also published as Crustacean Issues Vol. 20 by CRC Press, Boca Raton). Bracken-Grissom HD, Cannon ME, Cabezas P, Feldmann RM, Schweitzer CE, et  al. (2013) A comprehensive and integrative reconstruction of evolutionary history for Anomura (Crustacea: Decapoda). BMC Evolutionary Biology 13, 128. doi:10.1186/1471-2148-13-128 Cormie AK (1993) The morphology of the first zoeal stage of Lomis hirta (Lamarck, 1818) (Decapoda, Lomisidae). Crustaceana 64, 249–255. doi:10.1163/156854093X00289 McLaughlin PA (1983) A review of the phylogenetic position of the Lomidae (Crustacea: Decapoda: Anomala). Journal of Crustacean Biology 3, 431–447. doi:10.2307/1548143 McLaughlin PA, Lemaitre R (1997) Carcinization in the Anomura – fact or fiction? I. Evidence from adult morphology. Contributions to Zoology 67, 79–123. doi:10.1163/18759866-06702001

The Paguroidea are the hermit crabs, coconut crab, and king crabs, a heterogeneous assemblage of over 800 species united most obviously by the asymmetry of the pleon. In the rare case of symmetry (Pylochelidae and members of some other families) a reduced fourth pereopod ties the family with other paguroids. Hermit crabs are immediately recognisable by their usual use of gastropod shells for protection of the soft pleon. Some deep-water forms use zoantharian colonies for protection, or do not carry a any protection. Coconut crabs carry a shell as juveniles but fend for themselves as adults. Hermit crabs range from polar to tropical latitudes, semiterrestrial to abyssal habitats, and are often encountered on intertidal rocky shores. The curious relationship between hermit crabs and shells has been investigated many times (Hazlett 1981). Hermit crabs adopt numerous feeding strategies, most species are not especially fussy. Schembri’s (1982) study of fifteen species in New Zealand found deposit-feeding, scavenging, predation and filter-feeding to be options for many species. Hermit crabs also often harbour a wide community of associates on and inside their gastropod housing (Williams and McDermott 2004). King crabs, sometimes called stone crabs, are less commonly met, some in shallow subtidal habitats, some reaching abyssal depths, but most being confined to slope depths. Many are large enough to support major fisheries. The interrelationships of the paguroid families to each other and to other anomurans has been widely debated (Ahyong et  al. 2009; McLaughlin 1983a; McLaughlin and Lemaitre 1997; McLaughlin et  al. 2007). The most recent and comprehensive phylogenetic studies suggest that the paguroids are polyphyletic, with the symmetrical hermit crabs (Pylochelidae) and deep-sea hermit crabs (Parapaguridae) more closely related to clades of squat lobsters than to other hermit crabs (Ahyong et  al. 2009; Bracken-Grissom et al. 2013; Tsang et al. 2011; Wolfe et al. 2019). Just as crablike form has evolved more than once within the anomurans, it appears, so has the propensity to use gastropod shells or other hollow portable objects for protection. Another outcome of recent phylogenetic appraisals of hermit crabs is identification of polyphyly of the three main hermit crab families, Paguridae, Parapaguridae and Diogenidae. This issue has begun to be addressed by recent proposal of four additional families, Calcinidae and

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Paguropsidae (each for several genera taken out of Diogenidae), Xylopaguridae (for two genera removed from Paguridae) and Probeebeidae (for several genera removed from Diogenidae and Parapaguridae) (Fraaije et al. 2017; Gašparič et al. 2016; Fraaije et al. 2022). Here, we include Xylopaguridae, but provisionally retain Diogenidae and Parapaguridae in their traditional sense. Although Calcinus and allies phylogenetically stand separate from the core group of diogenid genera, further study is required to properly determine which extant genera should compose Calcinidae and a revised Diogenidae. Similarly, the respective phylogenetic positions of the diogenids, Paguropsis and Paguropsina, are unclear and warrant further study. Therefore, for the purposes of this guide, we provisionally use Diogenidae and Parapaguridae in their traditional sense. The wider polyphyly of the hermit crabs demonstrated by phylogenetics is yet to be formally reflected in the classification. Definition of the Paguroidea used here is drawn from

three important papers by McLaughlin (McLaughlin 1983a, 1983b, 2003) in which the superfamily Coenobitoidea is no longer recognised (Bracken-Grissom et  al. 2013). In many ways this makes more common sense than earlier classifications that had the two major families of hermit crabs in separate superfamilies, especially with discovery of a family that bridges both (reported in McLaughlin and Lemaitre 2001). Diagnosis. Sternal plastron consisting of thoracic sternites 3–7, thoracic sternite 8 independent. Pleon asymmetrical, or symmetrical (most Pylochelidae); pleonites indistingishable, or separate, calcified and distinct (Pylochelidae, Probeebei, Birgus). Ocular scales present. Antennal peduncle of 5 articulating articles. Pereopods 1 usually asymmetrical, short, stout, chelate. Pereopods 2–3 with claw-like dactyli. Pereopod 4 reduced, smaller than 2 and 3, or ambulatory, like 2 and 3 (Lithodidae only). Uropods often asymmetrical, not forming tail-fan, with rasps, or absent. Telson undivided, usually asymmetrical.

Key families of Paguroidea 1. – 2. – 3. – 4. – 5. – 6. – 7. –

Pereopods 4 developed as normal walking legs; body crab-like. Pleon recurved and carried under cephalothorax (Figs 13.20, 13.21)���������������������������������������������������������������������������������������� Lithodidae (king crabs) Pereopods 4 not developed as normal walking legs; body not crab-like. Pleon usually not recurved and carried under cephalothorax���������������������������������������������������������������������������������������������������������������������������������������������2 Mandible with incisor process mostly corneous, armed with prominent, acute teeth (Fig. 13.30b, c). Thoracic sternites 7 and 8 distinctly separated by membranous area (Fig. 13.30a)����������������������������� Pylojacquesiidae Mandible with incisor process calcareous or at most with only mesial edge corneous, lacking acute teeth. Thoracic sternites 7 and 8 fused�����������������������������������������������������������������������������������������������������������������������������������������������3 Maxillipeds 3 generally widely separated basally (Fig. 13.25g). Chelipeds unequal or less frequently subequal, right largest����������������������������������������������������������������������������������������������������������������������������������������������������������������4 Maxillipeds 3 generally close together basally (Fig. 13.19i). Chelipeds equal, subequal or unequal, left frequently larger��������������������������������������������������������������������������������������������������������������������������������������������������������������������������6 Maxilliped 1 exopod with flagellum (Fig. 13.25f)�����������������������������������������������������������������������������������������������������������������������5 Maxilliped 1 exopod without flagellum (Fig. 13.28l)�����������������������������������������������������������������������������������������Parapaguridae Pleonite 6 operculiform (Fig. 13.31a)���������������������������������������������������������������������������������������������������������������������Xylopaguridae Pleonite 6 not operculiform��������������������������������������������������������������������������������������������������������������������������������������������Paguridae Antennule upper flagellum ending in blunt stick-like article (Fig. 13.17c). Semiterrestrial hermit crabs and coconut crabs���������������������������������������������������������������������������������������������������������������������������������������Coenobitidae Antennule upper flagellum flexible�����������������������������������������������������������������������������������������������������������������������������������������������7 Paired pleopods on pleonites 2–5. Body straight, bilaterally symmetrical (Fig. 13.29). Deep-sea species����������������������������������������������������������������������������������������������������������������������������������������������������� Pylochelidae No paired pleopods on pleonites 4 and 5. Body usually coiled, asymmetrical (sometimes straight, symmetrical: Cancellus). Typical hermit crabs (Figs 13.18, 13.19)�������������������������������������������������������������������� Diogenidae

References Ahyong ST, Schnabel KE, Maas EW (2009) Anomuran phylogeny: new insights from molecular data. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 399–414. CRC Press, Boca Raton.

Bracken-Grissom HD, Cannon ME, Cabezas P, Feldmann RM, Schweitzer CE, et  al. (2013) A comprehensive and integrative reconstruction of evolutionary history for Anomura (Crustacea: Decapoda). BMC Evolutionary Biology 13, 128. doi:10.1186/1471-2148-13-128 Fraaije RHB, Van Bakel BW, Jagt JW (2017) A new paguroid from the type Maastrichtian (upper Cretaceous, the Netherlands) and erection of a

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

new family. Bulletin de la Société Géologique de France 188, 17–20. doi:10.1051/bsgf/2017185 Fraaije RHB, Van Bakel BWM, Jagt JWM, Charbonnier S, Schweigert G, Garcia G, Valentin X (2022) The evolution of hermit crabs (Crustacea, Decapoda, Anomura, Paguroidea) on the basis of carapace morphology: a state-of-the-art-report. Geodiversitas 44, 1–16. Gašparič R, Fraaije RHB, Robin N, de Angeli A (2016) The first record of paguroids from the Eocene of Istria (Croatia) and further phylogenetic refinement of the Paguroidea (Crustacea, Anomura). Bulletin of Geosciences 91, 467–480. doi:10.3140/bull.geosci.1616 Hazlett BA (1981) The behavioural ecology of hermit crabs. Annual Review of Ecology and Systematics 12, 1–22. doi:10.1146/annurev.es.12.​ 110181.000245 McLaughlin PA (1983a) Hermit crabs  – are they really polyphyletic? Journal of Crustacean Biology 3, 608–621. doi:10.2307/1547956 McLaughlin PA (1983b) A review of the phylogenetic position of the Lomidae (Crustacea: Decapoda: Anomala). Journal of Crustacean Biology 3, 431–447. doi:10.2307/1548143 McLaughlin PA (2003) Illustrated keys to families and genera of the superfamily Paguroidea (Crustacea: Decapoda: Anomura), with diagnoses of genera of Paguridae. Memoirs of Museum Victoria 60, 111–144. doi:10.24199/j.mmv.2003.60.16 McLaughlin PA, Lemaitre R (1997) Carcinization in the Anomura – fact or fiction? I. Evidence from adult morphology. Contributions to Zoology 67, 79–123. doi:10.1163/18759866-06702001 McLaughlin PA, Lemaitre R (2001) A new family for a new genus and new species of hermit crab of the superfamily Paguroidea (Decapoda: Anomura) and its phylogenetic implications. Journal of Crustacean Biology 21, 1062–1076. doi:10.1163/20021975-99990198 McLaughlin PA, Lemaitre R, Sorhannus U (2007) Hermit crab phylogeny: a reappraisal and its “fall-out”. Journal of Crustacean Biology 27, 97–115. doi:10.1651/S-2675.1 Schembri PJ (1982) Feeding behaviour of fifteen species of hermit crabs (Crustacea: Decapoda: Anomura) from the Otago region, southeastern New Zealand. Journal of Natural History 16, 859–878. doi:10.1080/​ 00222938200770691 Tsang LM, Chan T-Y, Ahyong ST, Chu KH (2011) Hermit to king, or hermit to all: multiple transitions to crab-like forms from hermit crab ancestors. Systematic Biology 60, 616–629. doi:10.1093/sysbio/syr063 Williams JD, McDermott JJ (2004) Hermit crab biocoenoses: a worldwide review of the diversity and natural history of hermit crab associates. Journal of Experimental Marine Biology and Ecology 305, 1–128. doi:10.1016/j.jembe.2004.02.020 Wolfe JM, Breinholt JW, Crandall KA, Lemmon AR, Lemmon EM, et  al. (2019) A phylogenomic framework, evolutionary timeline and genomic resources for comparative studies of decapod crustaceans. Proceedings. Biological Sciences 286, 20190079. doi:10.1098/rspb.2019.0079

Coenobitidae Dana, 1851 terrestrial hermit crabs Figure 13.17, Plate 30e–g The family comprises two genera of terrestrial or semiterrestrial hermit crabs. Both have marine larval stages and early juveniles that are fully aquatic. Members of the genus Coenobita are typical hermit crabs in carrying a gastropod shell to protect the soft, asymmetrically coiled pleon. Whereas species of Coenobita carry a shell throughout their lives, only

303

Fig. 13.17.  Coenobitidae. a, Coenobita rugosus H. Milne Edwards, 1837; b, antennule, Coenobita.

early juveniles of Birgus carry a protective gastropod shell, which they later abandon with increasing width and calcification of the carapace and pleon (Drew et al. 2010). Dimensions are of the length of the shield (sl.), the anterior harder part of the carapace in front of the cervical groove. Diagnosis. Left cheliped larger than right. Antennule, upper flagellum ending in blunt stick-like article (Fig. 13.17b). Maxillipeds 3 adjacent at base. Semiterrestrial. Birgus Leach, 1816 robber crab, coconut crab Diagnosis. Carapace broad, with expanded gill chambers. Rostrum well developed. Pereopod 4 elongate, chelate. Pleon symmetrical, simply flexed, dorsally protected by large, overlapping, strongly-calcified terga. Maximum sl. Males, 160 mm, females, 120 mm (Pl. 30g). Semiterrestrial, maritime shores on islands. Indo-West Pacific. 1 species. The only species, Birgus latro (Linnaeus, 1767), gets its common names from its so-called liking for coconuts, which it is supposed to steal from the trees. Its general appearance and habits are well known from numerous general accounts and illustrations on the web and published in the popular literature. The coconut crab is a popular delicacy throughout islands of the Indo-West Pacific (Widiyanti et al. 2016). The species previously occurred on the shores of Madagascar, Mauritius and tropical Australia but has been exploited to extinction there now. At up 4 kg it is the largest land-living arthropod species. Males are larger than females. Juveniles carry gastropod shells as do most hermit crabs but at ~15 mm total length long they abandon their protection and migrate to land where they inhabit beaches, coconut groves and damp forests up to 300 m above sea level. Adults are opportunistic omnivores but are

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Marine Decapod Crustacea

commonly found feeding on the fruit and pith of fallen trees. The biology of robber crabs has been reviewed most recently by Drew et al. (2010) and Anagnostou and Schubart (2014).

Coenobita Latreille, 1829 land hermit crabs Diagnosis. Carapace longer than wide. Rostrum obsolete. Pereopod 4 short, semichelate. Pleon asymmetrical, soft, twisted, integument membranous (Fig. 13.17a). Maximum sl. 23 mm (Pl. 30e, f). Semiterrestrial, sandy and muddy shore, maritime. Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific. 17 species. Terrestrial hermit crabs are widely distributed in tropical and subtropical regions where they scavenge terrestrial debris. Several species may occur sympatrically in coastal forests where their ecology is well studied (e.g. Hamasaki et al. 2018; Hsu et al. 2018). Land hermit crabs are sold on the domestic market in many countries as pets. This depends on harvesting in the wild with negative consequences for local populations (Bundhitwongrut 2018). The phylogeography of some species in the north-western Pacific has been elucidated by Hamasaki et  al. (2017). Nakasone (1988) provided a key to seven species from Japan, some more widespread in the Indo-West Pacific. Asakura’s (2004) key, in Japanese, included eight species. Rahayu et al. (2016) were the most recent to describe a new species and discussed possible past misidentifications of some species. References Alcock A (1905) Catalogue of the Indian decapod Crustacea in the collection of the Indian Museum. Part II. Anomura. Fasciculus I. Pagurides. Trustees of the Indian Museum, Calcutta. Anagnostou C, Schubart CD (2014) Morphometric characterisation of a population of adult coconut crabs Birgus latro (Decapoda: Anomura: Coenobitidae) from Christmas Island in the Indian Ocean. Raffles Bulletin of Zoology Supplement 30, 139–149. Asakura A (2004) Recent topics on taxonomy of hermit crabs from Japanese waters–family Coenobitidae. Kaiyo To Seibutsu 26, 83–89[in Japanese with English abstract]. Bundhitwongrut T (2018) Unregulated trade in land hermit crabs in Thailand. The Natural History Bulletin of the Siam Society 63, 27–40. Drew MM, Harzsch S, Stensmyr M, Erland S, Hansson BS (2010) A review of the biology and ecology of the robber crab, Birgus latro (Linnaeus, 1767) (Anomura: Coenobitidae). Zoologischer Anzeiger 249, 45–67. doi:10.1016/j.jcz.2010.03.001 Hamasaki K, Iizuka C, Sanda T, Imai H, Kitada S (2017) Phylogeny and phylogeography of the land hermit crab Coenobita purpureus (Decapoda: Anomura: Coenobitidae) in the Northwestern Pacific Region. Marine Ecology (Berlin) 38, e12369. doi:10.1111/maec.12369 Hamasaki K, Saeki E, Mizuta K, Tanabe M, Yamazaki I, et al. (2018) Tolerance of low salinity by larvae in six terrestrial hermit crab species (Decapoda: Anomura: Coenobitidae). Crustacean Research 47, 101–110. doi:10.18353/crustacea.47.0_101 Hsu C-H, Otte ML, Liu C-C, Chou J-Y, Fang W-T (2018) What are the sympatric mechanisms for three species of terrestrial hermit crab (Coenobita rugosus, C. brevimanus, and C. cavipes) in coastal forests? PLoS One 13, e0207640. doi:10.1371/journal.pone.0207640 Nakasone Y (1988) Land hermit crabs from the Ryukyus, Japan, with a description of a new species from the Philippines (Crustacea, Decapoda, Coenobitidae). Zoological Science 5, 165–178.

Orbigny CDd (1849) Dictionnaire universel d’histoire naturelle: résumant et complétant tous les faits présentés par les encyclopédies, les anciens dictionnaires scientifiques, les Oeuvres complètes de Buffon, et les meilleurs traités spéciaux sur les diverses branches des sciences nautrelles; donnant la description des etres et des divers phénomènes de la nature, l’étymologie et la définition des noms scientifiques, et les principales applications des corps organiques et inorganiques à l’agriculture, à la médecine, aux arts industriels, etc. Vol. Atlas v. 3 (Zoologie-Botanique). MM. Renard & co, Paris. Rahayu DL, Shih HT, Ng PKL (2016) A new species of land hermit crab in the genus Coenobita Latreille, 1829 from Singapore, Malaysia and Indonesia, previously confused with C. cavipes Stimpson, 1858 (Crustacea: Decapoda: Anomura: Coenobitidae). Raffles Bulletin of Zoology Supplement 34, 470–488. Widiyanti SE, Sukoso M, Setyohadi D (2016) Population dynamic of coconut crab (Birgus latro) in Sayafi Island, North Maluku of Indonesia. Journal of Biodiversity and Environmental Sciences 8, 151–161.

Diogenidae Ortmann, 1892 left-handed hermit crabs, equal-handed hermit crabs Figures 13.18, 13.19, Plates 32, 33a–e Diogenid hermit crabs are most easily distinguished from pagurids by having the left cheliped larger than the right, or equal chelipeds. The family includes some of the largest hermit crabs known, such as species of Dardanus and the deep-water Tisea grandis Morgan & Forest, 1991, with a carapace 57 mm long, but many species are very small, 2–4  mm long as adults. The family is most diverse throughout coral reefs and shallow water of the Indo-West Pacific with few Atlantic genera. Many diogenids are colourful and colour patterns are useful to distinguish species. Diogenidae, as currently composed is not a natural group as shown by phylogenetic analyses (Bracken-Grissom et al. 2013; Tsang et al. 2011) although the pattern of non-monophyly is yet to be fully evaluated. A new family, Calcinidae, was recently proposed for several genera taken out of Diogenidae, informed primarily by palaeontological observations (Fraaije et al. 2017). We do not doubt that Calcinus stands phylogenetically apart from the core group of diogenid genera, but further study is required to determine which extant genera should comprise Calcinidae, which should remain in Diogenidae, and which character suites best support such a division, critical for effective keys. Therefore, we provisionally employ Diogenidae in its traditional sense pending further elaboration of Calcinidae. Keys to genera have been offered by the renowned French carcinologist Jacques Forest whose prolific writings on anomurans have contributed so much to knowledge of this difficult group. His key (Forest 1984) was later updated (Forest 1995) and then again (McLaughlin 2003). These keys form the basis of the one below to include all genera now recognised. The first two couplets of the key are

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

difficult and involve careful microscopic investigation under the carapace. Most members of the family have ten arthrobranchs, two on the soft cuticle of each of the five thoracic somites 3–7 (carrying maxilliped 3 to pereopod 4). They also all have a pleurobranch on the hard exoskeleton above the limb on thoracic somites 5–7 (carrying pereopods 2–4) and may have one on the last thoracic somite. Put more simply, in the 14-gilled genera there is a gill above the last leg and in the 13-gilled genera there is not. See McLaughlin (2003) for illustrations. In Pseudopaguristes gills are vestigial and Areopaguristes lacks a further pleurobranch above pereopod 3. Pereopod 1 is the

305

cheliped, pereopods 2 and 3 the two pairs of walking legs, and pereopods 4 and 5 reduced. Dimensions are of the length of the shield (sl.), the anterior harder part of the carapace in front of the cervical groove. Diagnosis. Pleonite 1 distinct from last thoracic somite. Left cheliped larger than right (Fig.  13.18b), or chelipeds subequal (Fig.  13.18a), or right cheliped larger than left (rare). Maxillipeds 3 adjacent at base. Pleopods 3–5 on left side only, or absent in male; pleopods 1 (sometimes 1 and 2) paired (Fig. 13.19p).

Key to genera of Diogenidae 1.

Arthrobranchs vestigial at bases of cheliped and maxilliped 3. Pleurobranch absent above pereopod 2������������������������������������������������������������������������������������������������������������������������������������������������������� Pseudopaguristes – Arthrobranchs well developed at bases of cheliped and maxilliped 3. Pleurobranch present above pereopod 2���������������2 2. Pleurobranch present above pereopod 5 (14 pairs of gills)��������������������������������������������������������������������������������������������������������3 – Pleurobranch absent above pereopod 5 (12 or 13 pairs of gills)����������������������������������������������������������������������������������������������14 3. Maxillule endopod with well developed external lobe (Fig. 13.19h)����������������������������������������������������������������������������������������4 – Maxillule endopod without well developed external lobe (Fig. 13.19g)��������������������������������������������������������������������������������10 4. Maxilliped 3 ischium with well developed crista dentata (Fig. 13.19j)������������������������������������������������������������������������������������5 – Maxilliped 3 ischium without well developed crista dentata (Fig. 13.19k)�����������������������������������������������������������������������������8 5. Chelipeds equal or unequal, each with stridulatory mechanism developed on mesial face of palm (Fig. 13.18h)��������������� 6 – Chelipeds markedly unequal, left largest, without stridulatory mechanism����������������������������������������������������Allodardanus 6. Chelipeds carpi and chelae acute, corneous-tipped spines (Fig. 13.18h). Male pleopod 2 paired���������������� Strigopagurus – Chelipeds carpi and chelae with tubercles or transverse striate. Male pleopod 2 unpaired�������������������������������������������������7 7. Chelipeds equal or left larger; carpus and palm with transverse striae bordered with fine setae (Fig. 13.18e). Ambulatory legs dactyli equal to or longer than propodi. Females pleopods 2–5 unpaired, egg-carrying����������������������������������������������������������������������������������������������������������������������������������������� Ciliopagurus – Chelipeds equal, carpus and palm covered with generally blunt tubercles. Ambulatory legs dactyli much shorter than propodi. Female pleopod 5 non egg-carrying��������������������������������������������������������������������������� Trizopagurus 8. Antennal flagella with microscopic setae������������������������������������������������������������������������������������������������������������ Pseudopagurus – Antennal flagella with paired, long setae�������������������������������������������������������������������������������������������������������������������������������������9 9. Chelipeds equal or slightly subequal, similar; dactyli opening horizontally (Fig. 13.18c)������������������������������������� Isocheles – Chelipeds unequal and dissimilar; dactyli opening almost vertically (Fig. 13.18g)�����������������������������������������Loxopagurus 10. Chelae symmetrical, together forming operculum (Fig. 13.18a). Uropods symmetrical���������������������������������������Cancellus – Chelae symmetrical or asymmetrical, not forming operculum. Uropods asymmetrical��������������������������������������������������11 11. Shield without prominent Y-shaped suture. Chelipeds subequal or right larger���������������������������������������������������������������� 12 – Shield with prominent Y-shaped suture. Left cheliped larger than right (rarely subequal)�����������������������������������������������13 12. Eyes reaching antennal peduncle article 4. Posterior carapace well calcified���������������������������������������������������������������� Tisea – Eyes reaching beyond antennal peduncle. Posterior carapace primarily membranous�������������������������������������Petrochirus 13. Rostrum triangular. Ocular acicle with prominent triangular or subtriangular acicular projection (Fig. 13.19a). Chelipeds and ambulatory legs with ring-like transverse striae (Fig. 13.18i). Female with brood pouch ������������������������������������������������������������������������������������������������������������������������������������������������������������������ Aniculus – Rostrum broadly rounded or obsolete. Ocular acicle with subquadrate acicular projection (Fig. 13.19e). Chelipeds and ambulatory legs without ring-like transverse striae. Female without brood pouch�������������� Dardanus 14. 12 pairs of gills, pleurobranchs absent above pereopods 3 and 5���������������������������������������������������������������������Areopaguristes

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Marine Decapod Crustacea

Fig. 13.18.  Diogenidae. a, Cancellus typus; b, Dardanus setifer. Left chelipeds: c, Isocheles; d, Calcinus; e, Ciliopagurus; f, Dardanus; g, Loxopagurus; h, Strigopagurus. Pereopod 3: i, Aniculus.

– 15. – 16. –

13 pairs of gills, pleurobranchs present above pereopods 3, absent above pereopod 5�������������������������������������������������������15 Male pleopods 1 and/or 2 paired, modified as gonopods. Female pleopod 1 paired, modified or not����������������������������16 Paired pleopods absent in both sexes������������������������������������������������������������������������������������������������������������������������������������������19 Pleon twisted; uropods asymmetrical. Pereopod 4 not chelate (Fig. 13.19m). Pleopods 3–5 unpaired on left side�������������� 17 Pleon curling under but not twisted; uropods symmetrical. Pereopod 4 chelate (Figs 13.22n, o). Pleopods 3–5 unpaired on right or left side�������������������������������������������������������������������������������������������������������������������������������������������18 17. Telson posterior margin with 2 unequal lobes (Fig. 13.19s). Male pleopods 1, 2 (Fig. 13.19p) and female pleopods 1 modified as gonopods����������������������������������������������������������������������������������������������������������������������������Paguristes – Telson posterior margin with 4 lobes, posterior margin divided in 4 lobes, outer lobes unequal, larger than symmetrical inner lobes (Fig. 13.19t). Male pleopods 1 modified as gonopods (Fig. 13.19q), male and female pleopods 2 absent���������������������������������������������������������������������������������������������������������������������������������������Tetralobistes 18. Pereopod 4 dactylus cutting edge armed with row of corneous spines, fixed finger cutting edge with sharp spines claw-like (Fig. 13.19o). Gill lamellae distally divided into filamentous or stub-like extensions (Figs 13.19v). Maxilliped 3 exopod 4 or more times as long as wide������������������������������������������������������������������Paguropsis – Pereopod 4 dactylus cutting edge and fixed finger cutting edges unarmed or with one distinct corneous spinule (Figs 13.19n). Gill lamellae deeply divided distally into finger-like extensions (Figs 13.19u). Maxilliped 3 exopod 2.4 times as long as wide��������������������������������������������������������������������������������������������������Paguropsina 19. Chelipeds subequal����������������������������������������������������������������������������������������������������������������������������������������������������������������������� 20 – Chelipeds unequal, left appreciable larger����������������������������������������������������������������������������������������������������������������������������������21 20. Ocular acicles bi- or multispinose, contiguous or closely set (Fig. 13.19c). Pleonite 6 posterior margin unarmed��������������������������������������������������������������������������������������������������������������������������������������������������������������������Clibanarius – Ocular acicles simple, widely separated (Fig. 13.19b). Pleonite 6 posterior margin spinulose (Fig. 13.19r)��� Bathynarius 21. Rostrum obsolete to broadly rounded; with intercalary rostral process present, well developed, reduced or vestigial (Fig. 13.19f). Cheliped fingers with darks corneous tips�������������������������������������������������������������������������Diogenes – Rostrum moderate to well developed; without intercalary rostral process. Cheliped fingers with calcareous tips (Fig. 13.18d)������������������������������������������������������������������������������������������������������������������������������������������������������������ Calcinus

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

307

Fig. 13.19.  Diogenidae. Shield, eyestalks, ocular oscicles: a, Aniculus; b, Bathynarius; c, Clibanarius; d, Loxopagurus; e, Dardanus; f, Diogenes. Maxillule: g, Aniculus; h, Strigopagurus. Maxilliped 3: i, Clibanarius (pair); j, Ciliopagurus; k, Paguristes. Pereopod 4, propodus, dactylus: m, Paguristes; n, Paguropsina; o, Paguropsis; Male pleopods 1, 2 in situ: p, Paguristes. Male pleopod 1, q, Tetralobistes. Telson: r, Bathynarius; s, Paguristes; t, Tetralobistes. Gills: u, Paguropsina; v, Paguropsis.

Allodardanus Haig & Provenzano, 1965 Diagnosis. Pleurobranch present above pereopod 5. Ocular acicle simple, widely separated from its pair. Maxillule endopod with prominently recurved external lobe. Chelipeds markedly unequal, left larger. Male pleopod 2 unpaired, left side only. Female pleopods unpaired. Maximum sl. 25 mm. Shelf, slope (73–450 m). Tropical W Atlantic, Tropical Eastern Pacific. 3 species (McLaughlin and Gore 1985).

Aniculus Dana, 1852 Diagnosis. Pleurobranch present above pereopod 5. Shield with prominent Y-shaped suture in membranous posterior half. Ocular acicle with prominent triangular or subtriangular projection. Maxillule endopod without well developed external lobe. Chelipeds and ambulatory legs with ring-like transverse striae. Chelipeds markedly unequal, left larger. Male pleopods unpaired. Female pleopods unpaired, with brood pouch. Maximum sl. 52 mm (Pl. 32b). Intertidal–shelf (0–150 m). Temperate Northern W Pacific, Indo-West Pacific, Tropical Eastern Pacific, Temperate Southern Africa. 9 species. (Forest 1984: key to 8 species; McLaughlin and Hoover 1996)

Areopaguristes Rahayu & McLaughlin, 2010 Diagnosis. Pleurobranchs absent above pereopods 3 and 5 (12 pairs of gills). Ocular acicle elongate, spinulose. Maxillule endopod with prominently recurved external lobe. Chelipeds markedly unequal, left larger, or equal or subequal; dactyli opening in horizontal plane

or oblique plane. Male pleopods 1, 2 paired, modified as gonopods, 3–5 unpaired. Female pleopods 1 paired, modified as gonopods, 2–5 unpaired, without brood pouch (rarely) or with brood pouch. Telson posterior margin divided into 2 subequal to markedly unequal lobes. Maximum sl. 14 mm (Pl. 32a). Intertidal–shelf (0–150 m). Tropical Atlantic, Temperate Northern W Pacific, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate Southern Africa, Temperate Australasia. 29 species (Ayón-Parente et al. 2015: list of species, distributions; Komai 2009: as Striatiotes; Rahayu 2005; Rahayu and McLaughlin 2010: replacement name).

Bathynarius Forest, 1989 Diagnosis. Pleurobranch absent above pereopod 5. Pleonite 6 posterior margin spinulose. Ocular acicle simple, widely separated from its pair. Chelipeds equal or subequal. Male and female pleopods unpaired. Maximum sl. 11 mm. Shelf, slope (100–472 m). Tropical Atlantic, Indo-West Pacific. 5 species (Forest 1993: key to 4 species; Komai and Takeda 2004).

Calcinus Dana, 1851 Diagnosis. Pleurobranch absent above pereopod 5. Ocular acicle slender. Chelipeds markedly unequal, left larger; dactyli with calcareous tips; left carpus with prominent tubercle on upper-lateral surface. Male and female pleopods unpaired. Maximum sl. 19 mm (many species much smaller) (Pl. 32c–e). Intertidal–shelf (0–73 m). Temperate Northern W Pacific, Indo-West Pacific, Tropical Eastern Pacific, Temperate Southern

308

Marine Decapod Crustacea

Africa, Temperate Australia. 46 species (Morgan 1991: list of 33 world species, key to 17 Australian species; Poupin and McLaughlin 1998: key to Indo-West Pacfic species). Campos and Lemaitre (1994) compared species from the tropical Western Atlantic, Poupin and Lemaitre (2003) listed all from the Indo-West Pacific by region, and Poupin and Bouchard (2006) reviewed Eastern Pacific species.

Cancellus H. Milne Edwards, 1836 miner hermit crabs Diagnosis. Pleurobranch present above pereopod 5. Pleon spherical or ovoid sac, typically symmetrical. Maxillule endopod without well developed external lobe. Chelipeds symmetrical, together forming operculum with slightly concave surfaces of ambulatory legs. Male pleopods absent. Female pleopods unpaired, without brood pouch. Uropods symmetrical. Maximum sl. 11 mm (Pl. 32f). Intertidal–slope (0–402 m). Tropical Atlantic, Temperate Northern W Pacific, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate Southern Africa, Temperate Australasia (inhabiting cavities in stones, using the chelipeds and second pereopods as an operculum). 17 species (Forest et  al. 2000: rediagnosis; Mayo 1973: key to 9 species).

Ciliopagurus Forest, 1995 Diagnosis. Pleurobranch present above pereopod 5. Maxillule endopod with prominently recurved external lobe. Chelipeds markedly unequal, left larger, equal or subequal; each with stridulatory mechanism on mesial face of palm formed of rows of elongate corneous ridges; carpi and chelae with transverse striae bordered with setae. Male pleopod 2 unpaired, left side only. Female unpaired pleopods 2–5 egg-carrying. Maximum sl. 15 mm (Pl. 32g). Intertidal–slope (0–954 m). Tropical E Atlantic, Temperate Northern W Pacific, Indo-West Pacific (in gastropod shells of Conidae or Cypraeidae). 19 species (Forest 1995: distributions, key to 16 species; Poupin and Malay 2009: S. strigatus species complex). All species have striking red and yellow transverse stripes on the legs. Most species have flattened chelipeds.

Clibanarius Dana, 1852 Diagnosis. Pleurobranch absent above pereopod 5. Ocular acicle bi- or multispinose, contiguous or closely set. Chelipeds equal or subequal. Male and female pleopods unpaired. Maximum sl. 11 mm (Pl. 32h, i). Freshwater, estuarine, marine, intertidal–shelf (0–130 m). Temperate Northern W Pacific, Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific, Temperate Southern Africa, Temperate Australia. 58 species (Osawa and Fujita 2008). Species identification depends principally on colour pattern but keys are few (Malay et al. 2018; Yoshikawa et  al. 2019). Clibanarius is one of few genera of hermit crabs to tolerate brackish water and includes the only freshwater species, C. fonticola McLaughlin & Murray, 1990, in pools in Vanuatu (McLaughlin and Murray 1990).

Dardanus Paul’son, 1875 Diagnosis. Pleurobranch present above pereopod 5. Shield with prominent Y-shaped suture in membranous posterior half. Ocular acicle subrectangular or subquadrate projection. Maxillule endopod without well developed external lobe. Chelipeds subequal (rarely) or left larger. Male and female pleopods unpaired, without brood pouch. Maximum sl. 40 mm (Pl. 32k, l, 33a). Intertidal–slope (0–750 m). Cosmopolitan except Temperate S America and polar seas. 46 species (Ayón-Parente and Hendrickx 2009: 6 species from Eastern Pacific; Malay et al. 2018; McLaughlin 2002).

Diogenes Dana, 1851 Diagnosis. Pleurobranch absent above pereopod 5. Rostrum replaced by independent intercalary spine or scale between ocular acicles, sometimes small or obsolete. Maxilliped 3 crista dentata weak. Chelipeds markedly unequal, left larger. Male and female pleopods unpaired. Maximum sl. 14 mm (many tropical species 2–4 mm). Intertidal–shelf (0–99 m). Temperate Northern E Atlantic, Temperate Northern Pacific, Tropical Atlantic, Indo-West Pacific, Temperate Southern Africa, Temperate Australasia. 80 species (Rahayu and Pratiwi 2022: key to 22 species from Indonesia; Rahayu 2015: literature review).

Isocheles Stimpson, 1858 Diagnosis. Pleurobranch present above pereopod 5. Antennal flagella with paired, moderate to long setae. Ocular acicle bi- or multispinose, contiguous or closely set. Maxillule endopod without well developed external lobe. Maxilliped 3 crista dentata absent. Chelipeds equal or subequal; dactyli opening in horizontal plane. Male and female pleopods unpaired. Maximum sl. 20 mm. Subtidal, shelf (1–10, 188 m). Tropical W Atlantic, Tropical Eastern Pacific. 5 species (Forest and de Saint Laurent 1968; Mantelatto et al. 2006).

Loxopagurus Forest, 1964 Diagnosis. Pleurobranch present above pereopod 5. Antennal flagella with paired, moderate to long setae. Ocular acicle bi- or multispinose, contiguous or closely set. Maxillule endopod without well developed external lobe. Maxilliped 3 crista dentata absent. Chelipeds markedly unequal, left larger; dactyli opening in almost vertical plane. Male pleopods unpaired. Female pleopods unpaired. Maximum sl. 14 mm (Pl. 33c). Intertidal–shelf (0–30 m). Tropical W Atlantic. 1 species (Forest and de Saint Laurent 1968).

Paguristes Dana, 1851 Diagnosis. Pleurobranch absent above pereopod 5. Ocular acicle bi- or multispinose, contiguous or closely set, or elongate, spinulose. Chelipeds equal or subequal; dactyli opening in horizontal plane. Male pleopods 1, 2 paired, modified as gonopods, 3–5

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

unpaired. Female pleopods 1 paired, modified as gonopods, 2–5 unpaired (usually), with or without brood pouch. Telson posterior margin divided into 2 subequal to markedly unequal lobes. Maximum sl. 25 mm, typically much smaller (Pl. 33b, d). Intertidal–slope (0–1600 m depth). Cosmopolitan except Temperate South America. 121 species (Komai 2010; Komai et al. 2015; Rahayu 2006; Rahayu and Forest 2009).

Paguropsina Lemaitre, Rahayu & Komai, 2018 blanket hermit crabs Diagnosis. 13 pairs of gills; gill lamellae deeply divided distally into finger-like extensions. Pleon curling under but not twisted. Ocular acicle small, subtriangular, with small dorsodistal spine. Maxillule endopod with prominently recurved external lobe. Maxilliped 3 exopod 2.4 times as long as wide. Chelipeds equal or subequal. Pereopod 4 chelate, lacking rasp-like surfaces, dactylus and fixed finger cutting edges unarmed or with one distinct corneous spinule. Male pleopods 1, 2 paired, modified as gonopods, 3–5 unpaired, or pleopods absent. Female pleopods 1 paired, modified as gonopods, 2–5 unpaired, with brood pouch. Uropods symmetrical. Maximum sl. 6 mm. Shelf, slope (52–849 m; protected by a colonial sea-anemone having a sheet-like coenosarc that the hermit crab tucks under its telson and stretches over its back like a blanket). Central IndoPacific. 2 species (Lemaitre et al. 2018).

Paguropsis Henderson, 1888 blanket hermit crabs Diagnosis. 13 pairs of gills; gill lamellae distally divided into filamentous or stub-like extensions. Pleon curling under but not twisted. Ocular acicle small, subtriangular, with small dorsodistal spine. Maxillule endopod with prominently recurved external lobe. Maxilliped 3 exopod 4 or more times as long as wide. Chelipeds equal or subequal. Pereopod 4 chelate, lacking rasp-like surfaces, dactylus cutting edge armed with row of corneous spines, fixed finger cutting edge with sharp spines claw-like. Male pleopods 1, 2 paired, modified as gonopods, 3–5 unpaired, or pleopods absent. Female pleopods 1 paired, modified as gonopods, 2–5 unpaired, with brood pouch. Uropods symmetrical. Maximum sl. 23 mm. Shelf, slope (30–1125 m: using sea-anemones as protection). Western and Central Indo-Pacific. Temperate Australia (S Qld). 5 species (Lemaitre et al. 2018).

Petrochirus Stimpson, 1858 giant hermit crabs Diagnosis. Pleurobranch present above pereopod 5. Ocular acicle simple, widely separated from its pair. Eyes reaching beyond antennal peduncle. Maxillule endopod without well developed external lobe. Chelipeds subequal or right slightly larger. Male and female pleopods unpaired. Maximum sl. 45 mm (Pl. 33f).

309

Subtidal, shelf (1–160 m). Tropical Atlantic, Tropical Eastern Pacific. 3 species, well studied in western Atlantic (e.g. Buranelli and Mantelatto 2012; Williams 1984). Ball and Haig (1974) reported in detail on the eastern Pacific species.

Pseudopaguristes McLaughlin, 2002 Diagnosis. 8 pairs of functional gills; arthrobranchs at bases of maxilliped 3 and cheliped vestigial or absent; pleurobranch above pereopod 2 absent. Ocular acicle elongate, spinulose. Maxillule endopod with prominently recurved external lobe. Maxilliped 3 crista dentata weak. Chelipeds markedly unequal, left larger, or equal or subequal; dactyli opening in horizontal plane or oblique plane. Male pleopods unpaired, or pleopods 1, 2 paired, modified as gonopods, 3–5 unpaired. Female pleopods 1 paired, 2–5 unpaired, with or without brood pouch. Maximum sl. 6 mm. Intertidal–slope (0–739 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 15 species (Rahayu 2005; 2008: key to 8 species).

Pseudopagurus Forest, 1952 Diagnosis. Pleurobranch present above pereopod 5. Antennal flagella with microscopic setae. Ocular acicle bi- or multispinose, contiguous or closely set. Maxillule endopod without well developed external lobe. Maxilliped 3 crista dentata absent. Chelipeds markedly unequal, left larger. Male and female pleopods unpaired, without brood pouch. Maximum sl. 21 mm. Intertidal, subtidal (0–10 m). Tropical E Atlantic. 2 species (Forest 1952).

Strigopagurus Forest, 1995 Diagnosis. Pleurobranch present above pereopod 5. Maxillule endopod with prominently recurved external lobe. Chelipeds markedly unequal, left larger; each with stridulatory mechanism on mesial face of palm formed of rows of elongate corneous ridges; carpi and chelae with acute, corneous-tipped spines. Male pleopod 2 paired, 3–5 unpaired. Female pleopods unpaired. Maximum sl. 37 mm. Subtidal–slope (3–580 m). Temperate Northern W Pacific, Central and Eastern Indo-Pacific, Temperate Australasia. 5 species (Forest 1995).

Tetralobistes Ayon-Parente & Hendrickx, 2010 Diagnosis. Pleurobranch absent above pereopod 5. Ocular acicle elongate, spinulose. Maxillule endopod with prominently recurved external lobe. Chelipeds equal or subequal; dactyli opening in horizontal plane. Male pleopods 1 paired, modified as gonopods; pleopods 2 absent. Female pleopods 1 absent, 2–5 unpaired, without brood pouch. Telson posterior margin divided in 4 lobes, outer lobes unequal, larger than symmetrical inner lobes. Maximum sl. 2.1 mm. Intertidal, subtidal (0–18 m). Tropical Eastern Pacific. 2 species (Ayón-Parente and Hendrickx 2010).

310

Marine Decapod Crustacea

Tisea Forest & Morgan, 1991 Diagnosis. Strong calcification and numerous spine-tipped tubercles on posterior carapace. Pleurobranch present above pereopod 5. Ocular acicle small, subtriangular, with small dorsodistal spine. Eyes reaching to antennal article 4 Maxillule endopod without well developed external lobe. Chelipeds equal or subequal. Female pleopods unpaired, without brood pouch. Maximum sl. 59 mm, cl. 130 mm. Slope (265–350 m). Central Indo-Pacific (Timor Sea). 1 species (Morgan and Forest 1991).

Trizopagurus Forest, 1952 Diagnosis. Pleurobranch present above pereopod 5. Ocular acicle uni- or bidentate. Maxillule endopod with prominently recurved external lobe. Chelipeds equal or subequal; each with stridulatory mechanism on mesial face of palm formed of rows of elongate corneous ridges; carpi and chelae covered with blunt tubercles. Male pleopod 2 unpaired, left side only. Female unpaired pleopod 5 not egg-carrying. Maximum sl. 9.7 mm. Intertidal, subtidal (0–15 m). Tropical E Atlantic, Tropical Eastern Pacific (Central America). 3 species (Forest 1995). References Ayón-Parente M, Hendrickx ME (2009) A review of the Dardanus sinistripes (Stimpson, 1859) (Decapoda, Anomura, Diogenidae) species complex with the description of five new species from the Mexican Pacific. Zootaxa 2323, 1–71. doi:10.11646/zootaxa.2323.1.1 Ayón-Parente M, Hendrickx ME (2010) A new genus and new species of hermit crab (Crustacea: Anomura: Paguroidea: Diogenidae) from the eastern tropical Pacific. Zootaxa 2677, 49–59. doi:10.11646/zootaxa.​ 2677.1.5 Ayón-Parente M, Hendrickx ME, Lemaitre R (2015) Redescription and taxonomic status of Paguristes praedator Glassell, 1937 and P. oxyophthalmus Holthuis, 1959 (Anomura: Paguroidea: Diogenidae), with an emendation to the diagnosis of the genus Areopaguristes Rahayu & McLaughlin, 2010. Zootaxa 3915, 491–509. doi:10.11646/zootaxa.3915.4.2 Ball EB, Haig J (1974) Hermit crabs from the tropical Eastern Pacific. I. Distribution, color, and natural history of some common shallow-water species. Bulletin of the Southern California Academy of Sciences 73, 95–104. Bracken-Grissom HD, Cannon ME, Cabezas P, Feldmann RM, Schweitzer CE, et  al. (2013) A comprehensive and integrative reconstruction of evolutionary history for Anomura (Crustacea: Decapoda). BMC Evolutionary Biology 13, 128 doi:10.1186/1471-2148-13-128. Buranelli RC, Mantelatto FL (2012) Reproductive apparatus of the male giant hermit crab Petrochirus diogenes (Anomura, Diogenidae): morphology and phylogenetic implications. Aquatic Biology 16, 241–251. doi:10.3354/ab00453 Campos NH, Lemaitre R (1994) A new Calcinus (Decapoda: Anomura: Diogenidae) from the tropical Western Atlantic, and a comparison with other species of the genus from the region. Proceedings of the Biological Society of Washington 107, 137–150. Forest J (1952) Caractères et affinités de Pseudopagurus, genre nouveau établi pour un Paguridae de la côte occidentale d’Afrique, Pagurus granulimanus Miers. Bulletin de l’Institut Français d’Afrique Noire 14, 799–812. Forest J (1984) Révision du genre Aniculus (Decapoda Diogenidae). Crustaceana 8(Supplement), 1–91.

Forest J (1993) Présence du genre Bathynarius (Crustacea, Decapoda, Diogenidae) en Indonésie et dans le Pacifique central, avec la description de deux espèces nouvelles. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 14, 483–500. Forest J (1995) Crustacea Decapoda Anomura: révision du genre Trizopagurus Forest, 1952 (Diogenidae), avec l’établissement de deux genres nouveaux. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 13. Mémoires du Muséum National d’Histoire Naturelle, Paris 163, 9–149. Forest J, de Saint Laurent M (1968) Résultats scientifiques des campagne de la “Calypso”, VII. Campagne de la “Calypso” au large des côtes atlantiques de l’Amérique du Sud (1961–1962) 6. Crustacés Décapodes: Pagurides. Annales de l’Institut Océanographique de Monaco 45, 47–172. Forest J, de Saint Laurent M, McLaughlin PA, Lemaitre R (2000) The marine fauna of New Zealand: Paguridea (Decapoda: Anomura) exclusive of the Lithodidae. NIWA Biodiversity Memoir 114, 1–250. Fraaije RH, Van Bakel BW, Jagt JW (2017) A new paguroid from the type Maastrichtian (upper Cretaceous, the Netherlands) and erection of a new family. Bulletin de la Société Géologique de France 188, 17–20. doi:10.1051/bsgf/2017185 Komai T (2009) A review of the northwestern Pacific species of the genus Paguristes (Decapoda: Anomura: Diogenidae). II. Species transferred to the genus Stratiotes, with descriptions of two new species. Natural History Research 10, 59–91. Komai T (2010) A review of the northwestern Pacific species of the genus Paguristes (Decapoda: Anomura: Diogenidae). III. Clarification of the identity of a species heretofore referred to Paguristes balanophilus Alcock and descriptions of two new species from Japan. Natural History Research 11, 9–33. Komai T, Takeda M (2004) Two new deepwater species of hermit crabs (Crustacea: Decapoda: Anomura: Paguroidea) from Japan. Bulletin of the National Science Museum, Tokyo (ser. A). Zoology 30, 113–127. Komai T, Reshmi R, Kumar AB (2015) A new species of the hermit crab genus Paguristes Dana, 1851 (Crustacea: Decapoda: Anomura: Diogenidae) from southwestern India. Zootaxa 3937, 517–532. doi:10.11646/ zootaxa.3937.3.5 Lemaitre R, Rahayu DL, Komai T (2018) A revision of “blanket-hermit crabs” of the genus Paguropsis Henderson, 1888, with the description of a new genus and five new species (Crustacea, Anomura, Diogenidae). ZooKeys 752, 17–97. doi:10.3897/zookeys.752.23712 Malay MM, Rahayu DL, Chan T-Y (2018) Hermit crabs of the genera Calcinus Dana, Clibanarius Dana, and Dardanus Paul’son from the PANGLAO 2004 Expedition, with description of a new species and a checklist of the hermit crabs of the Philippines (Crustacea: Anomura). Raffles Bulletin of Zoology 66, 23–65. Mantelatto FL, Robles R, Biagi R, Felder DL (2006) Molecular analysis of the taxonomic and distributional status for the hermit crab genera Loxopagurus Forest, 1964 and Isocheles Stimpson, 1858 (Decapoda, Anomura, Diogenidae). Zoosystema 28, 495–506. Mayo BS (1973) A review of the genus Cancellus (Crustacea: Diogenidae) with the description of a new species from the Caribbean Sea. Smithsonian Contributions to Zoology 150, 1–63. doi:10.5479/si.00810282.150 McLaughlin PA (2002) A review of the hermit-crab (Decapoda: Anomura: Paguridea) fauna of southern Thailand, with particular emphasis on the Andaman Sea, and descriptions of three new species. Phuket Marine Biological Center Special Publication 23, 385–460. McLaughlin PA (2003) Illustrated keys to families and genera of the superfamily Paguroidea (Crustacea: Decapoda: Anomura), with diagnoses of genera of Paguridae. Memoirs of Museum Victoria 60, 111–144. doi:10.24199/j.mmv.2003.60.16

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McLaughlin PA, Gore RH (1985) A new species of Allodardanus (Decapoda, Diogenidae) from the Western Atlantic. Crustaceana 49, 36–41. doi:10.1163/156854085X00189 McLaughlin PA, Hoover JP (1996) A new species of Aniculus Dana (Decapoda: Anomura: Diogenidae) from Hawaii. Proceedings of the Biological Society of Washington 109, 299–305. McLaughlin PA, Murray T (1990) Clibanarius fonticola, new species (Anomura: Paguridae: Diogenidae), from a fresh-water pool on Espiritu Santo, Vanuatu. Journal of Crustacean Biology 10, 695–702. doi:10.2307/1548413 Morgan GJ (1991) A review of the hermit crab genus Calcinus Dana (Crustacea: Decapoda: Diogenidae) from Australia, with descriptions of two new species. Invertebrate Taxonomy 5, 869–913. doi:10.1071/IT9910869 Morgan GJ, Forest J (1991) A new genus and species of hermit crab (Crustacea, Anomura, Diogenidae) from the Timor Sea, north Australia. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 13, 189–202. Osawa M, Fujita Y (2008) Clibanarius ambonensis (Crustacea: Decapoda: Anomura: Diogenidae) from the Ryukyu Islands, south-western Japan. Marine Biodiversity Records 1, e16. doi:10.1017/S1755267206001667 Poupin J, Bouchard JM (2006) The eastern Pacific species of the genus Calcinus Dana, 1851, with description of a new species from Clipperton Atoll (Decapoda, Anomura, Diogenidae). Zoosystema 28, 465–486. Poupin J, Lemaitre R (2003) Hermit crabs of the genus Calcinus Dana, 1851 (Decapoda: Anomura: Diogenidae) from the Austral islands, French Polynesia, with description of a new species. Zootaxa 391, 1–20. doi:10.11646/zootaxa.391.1.1 Poupin J, Malay MC (2009) Identification of a Ciliopagurus strigatus (Herbst, 1804) species-complex, with description of a new species from French Polynesia (Crustacea, Decapoda, Anomura, Diogenidae). Zoosystema 31, 209–232. doi:10.5252/z2009n2a1 Poupin J, McLaughlin PA (1998) Additional records of Calcinus species (Decapoda: Anomura: Diogenidae) from French Polynesia with description of three new species and a key to Indo-West Pacific species of the genus. Crustacean Research 27, 9–27. doi:10.18353/crustacea.27.0_9 Rahayu DL (2005) Additions to the Indonesian fauna of the hermit crab genus Pseudopaguristes McLaughlin and a further division of the genus Paguristes Dana (Crustacea: Decapoda: Paguroidea: Diogenidae). Zootaxa 831, 1–42. doi:10.11646/zootaxa.831.1.1 Rahayu DL (2006) The genus Paguristes (Crustacea, Decapoda, Diogenidae) from Indonesia. In: Richer de Forges, B., and Justine, J.-L. (eds), Tropical Deep-Sea Benthos, Vol. 24. Mémoires du Muséum National d’Histoire Naturelle, Paris 193, 349–374. Rahayu DL (2008) The genus Pseudopaguristes McLaughlin (Crustacea: Decapoda: Anomura: Diogenidae) from the southwestern Pacific, with the descriptions of two new species. Zootaxa 1840, 54–66. doi:10.11646/zootaxa.1840.1.2 Rahayu DL (2015) New record and new species of the hermit crab genus Diogenes Dana, 1851 (Decapoda: Anomura: Diogenidae) from Singapore. Raffles Bulletin of Zoology Supplement 31, 182–192. Rahayu DL, Forest J (2009) Le genre Paguristes Dana aux Philippines avec la description de deux nouvelles espèces (Decapoda, Anomura, Diogenidae) (The genus Paguristes Dana in the Philippines with the description of two new species (Decapoda, Anomura, Diogenidae). Crustaceana 82, 1307–1338. doi:10.1163/001121609X12475745628388 Rahayu DL, McLaughlin PA (2010) Areopaguristes, a generic replacement name for Stratiotes Thomson, 1899 (Crustacea: Decapoda: Paguroidea: Diogenidae). Zootaxa 2509, 67–68. doi:10.11646/zootaxa.2509.1.6 Rahayu DL, Pratiwi R (2022) Diogenes matabiru, a new species of hermit crab from Lombok Island, Indonesia (Crustacea, Decapoda, Anomura, Diogenidae. Zootaxa 5093, 493–500. doi:10.11646/zootaxa.5093.4.7

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Tsang LM, Chan T-Y, Ahyong ST, Chu KH (2011) Hermit to king, or hermit to all: multiple transitions to crab-like forms from hermit crab ancestors. Systematic Biology 60, 616–629. doi:10.1093/sysbio/syr063 Williams AB (1984) Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C. Yoshikawa A, Ikeo K, Imoto J, Jaingam W, Putri LSE, et al. (2019) Molecular phylogeny of Clibanarius Dana, 1852 from the Indo-West Pacific: evolution of pereopod colour pattern and habitat adaptation. Crustaceana 92, 799–839. doi:10.1163/15685403-00003910

Lithodidae Samouelle, 1819 king crabs, stone crabs Figures 13.20–13.22, Plate 31 The king crabs or stone crabs occur worldwide and are largely deep-water species, although several occur in shallow water at high latitudes, especially in the North Pacific. Around 135 species are known of which more than 60 occur in the Pacific Ocean. King crabs are best known from several commercially important northern hemisphere species, especially Paralithodes camtschaticus (Tilesius, 1815) (Red King Crab) and Lithodes aequispinus (Benedict, 1895) (Golden King Crab) (Stevens 2014). In the southern hemisphere, Lithodes santolla (Molina, 1782) (Southern King Crab) and several species of Paralomis are commercially landed off southern South America, and exploratory fisheries for lithodids are active in New Zealand waters (Ahyong 2010). While many favour polar or subpolar regions some are known from close to hydrothermal vents or cool seeps (Chevaldonné and Olu 1996; Macpherson 1994; de Saint Laurent and Macpherson 1997). Lithodids differ from other paguroids in being crab-like, not inhabiting gastropod shells and in having a usually flattened, calcified pleon. Lithodids closely resemble brachyuran crabs but notably differ in having only three pairs of walking legs and an asymmetrical pleon in females. No other decapods more closely approximate the brachyuran form than do lithodids, being the paradigm example of carcinisation, that is, the evolution of a crab-like form from a non-crab ancestor (Keiler et al. 2015; Keiler et al. 2017). Lithodids have long been recognised as being derived from within the asymmetrical hermit crabs (specifically Paguridae), the so-called ‘hermit to king’ hypothesis (Ahyong and O’Meally 2004; Bouvier 1894; Cunningham et al. 1992; Morrison et  al. 2002; Richter and Scholtz 1994) and this is reflected in the historical placement of lithodids among the paguroids. Although several studies proposed an independent derivation of king crabs (McLaughlin and Lemaitre 1997; McLaughlin et  al. 2004), and a separate superfamily, Lithodoidea (with Lithodidae and Hapalogastridae) (McLaughlin et al. 2007), other phylogenetic studies consistently recovered the intrinsic connection between king crabs and pagurid

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hermits crabs (Ahyong et  al. 2009; Bracken-Grissom et  al. 2009; Schnabel et  al. 2011; Tsang et  al. 2008; Wolfe et  al. 2019). As such, we retain the king crabs in a single family, Lithodidae, within Paguroidea. We treat Hapalogastrinae, sometimes treated as a family, as one of two subfamilies. Major references on the taxonomy of lithodids with keys to species and entry to the literature can be found for the Atlantic Ocean (Macpherson 1988), for the eastern Pacific (Macpherson 1992; Macpherson and Wehrtman 2010; Retamal 1994), for the northern Pacific (Makarov 1938; Sakai 1971; Schmitt 1921), and for subantarctic and Australasian regions (Ahyong 2010; Macpherson 2004). The taxonomic literature dealing with the larger genera in the Indo-West

Pacific is more scattered (Ahyong 2020; Ahyong et al. 2010; Macpherson 1990; Macpherson and Chan 2008; Macpherson and Wehrtman 2010). An illustrated key to genera can be found in McLaughlin (2003). All species known at the time were listed by McLaughlin et al. (2010). Diagnosis. Crab-like. Maxilliped 1 exopod with flagellum. Maxillipeds 3 widely separate at base. Epistomial spines absent. Chelipeds equal or unequal, right usually larger. Pereopods 2–4 ambulatory. Pleon asymmetrical in females, symmetrical in males; terga membranous or composed of calcified plates and nodules; pleonite 1 strongly reduced; pleonites 2–6 well developed. Pleopods 2–5 unpaired in females, absent in males. Uropod absent.

Key to genera of Lithodidae 1. – 2. –

Rostrum usually well developed, with basal spine and 1 or more additional spines (Fig. 13.21). Pleon well calcified, consisting of calcified plates or numerous closely set spines (Fig. 13.22i–l)���������������������������Lithodinae … 2 Rostrum short, simple, triangular without additional dorsal or lateral spines (Fig. 13.20). Pleon soft, membranous, somites 3–5 largely membranous but usually with small, thin, well separated plates (Fig. 13.20f, g)���������������������������������������������������������������������������������������������������������������������������������������� Hapalogastrinae … 11 Carapace transversely ovate, without protuberances, granules or spines, nearly smooth; markedly wider than long, laterally strongly expanded entirely concealing pereopods 2–4 when folded. Rostrum wide, subquadrate to subhexagonal, distally truncate (Fig. 13.21a, b)������������������������������������������������������������������ Cryptolithodes Carapace triangular, pyriform or subpentagonal, little wider than long to little longer than wide; with numerous protuberances, granules or spines; lateral margins, at most, weakly expanded over pereopods 2–4, at most covering proximal part of merus. Rostrum variable in shape, but never distally truncated�������������������3

Fig. 13.20.  Lithodidae, Hapalogastrinae. a, Acantholithodes hispidus (Stimpson, 1860); b, Dermaturus mandtii Brandt, 1850; c, Hapalogaster cavicauda Stimpson, 1859; d, Oedignathus inermis (Stimpson, 1860); e, Placetron inermis (Stimpson, 1860). Pleonites 2 and pleonites 3– telson: f, Acantholithodes; g, Placetron.

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

3. – 4. – 5. –

313

Thoracic sternite 5 (pereopods 2) with deep longitudinal median fissure (Fig. 13.22g)������������������������������������������������������4 Thoracic sternite 5 (pereopods 2) without deep median fissure�����������������������������������������������������������������������������������������������6 Rostrum usually distally bifid, with 1 or 2 pairs of dorsal spines, 1 ventral spine (Figs 13.21d, 13.22a, b). Pleonite 2 composed of 3 plates (fused median and submedians, paired marginals; Fig. 13.22h) or fused into single plate���������������������������������������������������������������������������������������������������������������������������������������������������������������Lithodes Rostrum distally undivided, with pair of dorsal spines, without ventral spine. Pleonite 2 composed of 5 plates (median, paired submedians, paired marginals; Fig. 13.22i, j)���������������������������������������������������������������������������������5 Carapace as long as wide (Fig. 13.21f). Pleonites 3–5 in male comprising dense field of small, pointed or spiniform nodules joined by arthrodial membrane (Fig. 13.22i); female left side with well developed plate on each somite, right side membranous. Scaphocerite absent or rudimentary (Fig. 13.22c, d)��������� Neolithodes Carapace slightly wider than long (Fig. 13.21g). Pleonites 3–5 in both sexes membranous, with blunt, calcified nodules medially; submedian and marginal plates well defined on both sides (Fig. 13.22j). Scaphocerite of 1 or more spines (Fig. 13.22f)����������������������������������������������������������������������������������������������������Paralithodes

Fig. 13.21.  Lithodidae, Lithodinae. a, b, Cryptolithodes typicus Brandt, 1848; c, Glyptolithodes cristatipes (Faxon, 1893); d, Lithodes jessica Ahyong, 2010; e, Echidnocerus mandtii (Brandt, 1848); f, Neolithodes flindersi Ahyong, 2010; g, Paralithodes brevipes (H. Milne Edwards & Lucas, 1841); h, Paralomis echidna Ahyong, 2010; i, Phyllolithodes papillosus Brandt, 1848; j, Rhinolithodes wosnessenskii Brandt, 1848; k, Sculptolithodes derjugini Makarov, 1934.

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Marine Decapod Crustacea

Fig. 13.22.  Lithodidae, Lithodinae. Rostrum, dorsal, lateral: a, b, Lithodes; c, d, Neolithodes; e, Phyllolithodes. Antenna, scaphocerite: f, Paralithodes. Thoracic sternum, pereopods 1–4 coxae: g, Lithodes. Pleonite 2: h, Lithodes. Pleonites 2–6, telson: i, Neolithodes (female); j, Paralithodes (female); k, Paralomis (female); l, Phyllolithodes.

6.

Rostral apex divided into 2 rounded lobes (Figs 13.21i, Fig. 13.22e). Pleonites 3–5 submedian plates bearing concave, membranous areas. Pleonite 2 composed of 3 plates (median, paired marginals fused with submedians) (Fig. 13.22l)������������������������������������������������������������������������������������������������������������������������������������ Phyllolithodes – Rostral apex undivided. Pleonites 3–5 submedian plates evenly calcified, without concave, membranous areas. Pleonite 2 composed of 5 separate plates or a single plate�����������������������������������������������������������������������������������������7 7. Carapace subtriangular, wider than long, widest at posterior margin (Fig. 13.21j). Pleonite 2 composed of 5 plates. Pleonites 3–5 marginal and submedian plates fused������������������������������������������������������������������������� Rhinolithodes – Carapace as wide or longer than wide, widest anterior to posterior margin. Pleonite 2 entire, a single plate������������������8 8. Rostrum thick, blunt, linguiform in dorsal view, extending anteriorly beyond eyes by at least length of eye (Fig. 13.21k). Scaphocerite minute, rudimentary. Pleonites 3–5 median plate subdivided������������� Sculptolithodes – Rostrum short, conical, not extending beyond eyes, or extending beyond eyes by at most half eye-length. Scaphocerite well developed, spinose. Pleonite 3 median plate divided or undivided; pleonites 2–5 median plate undivided�������������������������������������������������������������������������������������������������������������������������������������������������������������9 9. Rostrum formed by basal spine and single dorsal spine or granule (Fig. 13.21c)������������������������������������������ Glyptolithodes – Rostrum formed by basal spine and 1 or more pairs of dorsal spines (sometimes small)��������������������������������������������������10 10. Pleonite 3 median plate subdivided into 3 parts. Pereopod 4 0.7 times carapace width (Fig. 13.21e)���������� Echidnocerus – Pleonite 3 median plate undivided (Fig. 13.22k). Pereopod 4 as long as or longer than carapace width (Fig. 13.21h)������������������������������������������������������������������������������������������������������������������������������������������������������������������ Paralomis 11. Carapace dorsal surface densely covered with upright spines (Fig. 13.20a)������������������������������������������������ Acantholithodes – Carapace dorsal surface without upright spines���������������������������������������������������������������������������������������������������������������������� 12 12. Carapace weakly calcified. Pereopods 2–4 merus extensor margins spinose or dentate (Fig. 13.20c, e)�������������������������13 – Carapace hard, well calcified. Pereopods 2–4 merus extensor margins unarmed (Fig. 13.20b, d)�����������������������������������14 13. Carapace surface strongly setose, obscuring uneven, granulate surface. Pereopods 2–4 shorter than twice carapace length (Fig. 13.20c)��������������������������������������������������������������������������������������������������������������������� Hapalogaster – Carapace covered with arcuate scales, scarcely setose. Pereopods 2–4 longer than twice carapace length (Fig. 13.20e)������������������������������������������������������������������������������������������������������������������������������������������������������� Placetron 14. Carapace dorsal surface covered with transverse grooves (Fig. 13.20b)��������������������������������������������������������������Dermaturus – Carapace dorsal surface covered with low, rounded tubercles or swellings (Fig. 13.20d)������������������������������� Oedignathus

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

315

Subfamily Hapalogastrinae Brandt, 1850

Subfamily Lithodinae Samouelle, 1819

Diagnosis. Rostrum short, simple, triangular without additional dorsal or lateral spines. Pleon soft, membranous, somites 3–5 largely membranous but usually with small, thin, well separated plates.

Diagnosis. Rostrum usually well developed, with basal spine and 1 or more additional spines. Pleon well calcified, consisting of calcified plates or numerous closely set spines. Implicit generic attributes. Scaphocerite well developed, spinose. Thoracic sternite 5 (pereopods 2) without deep, longitudinal, median fissure. Pereopods 2–4 fully visible dorsally.

Acantholithodes Holmes, 1895 Diagnosis. Carapace pyriform; dorsal surface densely covered with upright spines. Pereopods 2–4 ~1.5 times carapace length; merus extensor margins with sharp spines. Maximum cl. 62 mm. Intertidal–slope (0–245 m). Temperate Northern E Pacific. 1 species (Schmitt 1921: rediagnosis, description).

Dermaturus Brandt, 1850 Diagnosis. Carapace trapezoid, hard, well calcified; lateral margins unarmed; dorsal surface covered with long transverse grooves, sparsely setose. Pereopods 2–4 ~1.5 times carapace length; merus extensor margins unarmed. Maximum cl. 23 mm. Intertidal–shelf (0–72 m). Temperate Northern Pacific. 1 species (Makarov 1938: rediagnosis, species illustration and description; Schmitt 1921: rediagnosis, species illustration and description).

Hapalogaster Brandt, 1850 Diagnosis. Carapace trapezoid; lateral branchial margins with or without spines; dorsal surface strongly setose, unarmed, irregularly rugose. Pereopods 2–4 ~1.5 times carapace length; merus extensor margins with blunt spines. Maximum cl. 25 mm. Intertidal–shelf (0–180 m). Temperate Northern Pacific. 4 species (Makarov 1938: rediagnosis, key to 3 species; Schmitt 1921: rediagnosis, key to 2 species).

Oedignathus Benedict, 1895 Diagnosis. Carapace trapezoid, hard, well calcified; lateral margins unarmed; dorsal surface covered with low, rounded tubercles or swellings, sparsely setose. Pereopods 2–4 ~1.5 times carapace length; merus extensor margins unarmed. Maximum cl. 25 mm. Intertidal, subtidal (0–15 m). Temperate Northern Pacific. 1 species (Makarov 1938: rediagnosis, species illustration and description; Schmitt 1921: rediagnosis, species illustration and description).

Placetron Schalfeew, 1892 Diagnosis. Carapace trapezoid, weakly calcified; lateral margins unarmed; dorsal surface covered with arcuate scales, sparsely setose. Pereopods 2–4 longer than twice carapace length; merus extensor margins with blunt spines. Maximum cl. 65 mm. Intertidal–shelf (0–110 m). Temperate Northern E Pacific. 1 species (Makarov 1938: rediagnosis, description).

Cryptolithodes Brandt, 1848 Diagnosis. Carapace transversely ovate, without protuberances, granules or spines, nearly smooth; markedly wider than long, laterally strongly expanded. Rostrum wide, subquadrate to subhexagonal, distally truncate. Pereopods 2–4 concealed by carapace when folded. Maximum cl. 65 mm (Pl. 31a). Intertidal, subtidal (0–55 m). Temperate Northern Pacific. 3 species (Makarov 1938: rediagnosis, key to 3 species; Schmitt 1921: rediagnosis, key to 2 species).

Echidnocerus White, 1842 Diagnosis. Carapace subhexagonal, wider than long; with blunt granular dorsal protuberances and blunt marginal tubercles. Rostrum short, conical, extending beyond eyes by at most half eyelength, formed by basal spine and 1 or more pairs of dorsal spines. Pleonite 2 of single plate. Pleonites 3–5 submedian plates evenly calcified, without concave, membranous areas, not fused with marginal plates. Maximum cl. 200 mm. Intertidal–slope (0–547 m). Temperate Northern Pacific. 2 species (Makarov 1938: rediagnosis; Schmitt 1921: rediagnosis, key to 2 species). Echidnocerus has priority over the more commonly used Lopholithodes Brandt, 1848.

Glyptolithodes Faxon, 1895 Diagnosis. Carapace pentagonal, wider than long; with blunt granular dorsal protuberances and blunt marginal tubercles. Rostrum short, conical, not extending beyond eyes, formed by basal spine and single dorsal spine or granule. Thoracic sternite 5 (pereopods 2) without deep, longitudinal, median fissure. Pleonite 2 of single plate. Pleonites 3–5 submedian plates evenly calcified, without concave, membranous areas; median plate undivided. Maximum cl. 90 mm. Shelf, slope (245–800 m). Tropical Eastern Pacific, Temperate South America (Pacific coast). 1 species (Haig 1974: figure; Martin et al. 1997: distribution).

Lithodes Latreille, 1806 Diagnosis. Carapace pyriform, about as long as wide; with numerous granules or spines; branchial margins not expanded laterally. Rostrum comprised of median spine (usually distally bifid), 1 or 2 pairs of anterolaterally directed dorsal spines; prominent ventral spine. Scaphocerite well developed, spinose, or rudimentary. Thoracic sternite 5 (pereopods 2) with deep, longitudinal, median fissure. Pleonite 2 of single plate, or of 3 plates (fused median and

316

Marine Decapod Crustacea

submedians, paired marginals). Pleonites 3–5 in both sexes with blunt, calcified nodules medially; submedian and marginal plates well defined on both sides. Maximum cl. 200 mm (Pl. 31b). Subtidal–slope (16–1821 m). Cosmopolitan. 30 species (Ahyong 2010: key to 8 species from Australia and New Zealand; Macpherson 1988: rediagnosis, key to 8 Atlantic species).

spines. Rostrum apex divided into two rounded lobes. Pleonite 2 of 3 plates (fused median and submedians, paired marginals). Pleonites 3–5 submedian plates with concave, membranous areas. Maximum cl. 90 mm. Intertidal–shelf (0–183 m). Temperate Northern E Pacific. 1 species (Makarov 1938: rediagnosis; Schmitt 1921: rediagnosis).

Neolithodes A. Milne-Edwards & Bouvier, 1894

Rhinolithodes Brandt, 1848

Diagnosis. Carapace pyriform, about as long as wide; with numerous granules or spines; branchial margins not expanded laterally. Rostrum comprised of distally undivided median spine and pair of anterolaterally directed dorsal spines at base. Scaphocerite absent or rudimentary. Thoracic sternite 5 (pereopods 2) with deep, longitudinal, median fissure. Pleonite 2 of 5 plates (one median, paired submedians and marginal). Pleonites 3–5 in male comprising dense field of small, pointed or spiniform nodules joined by arthrodial membrane; female left side with well developed plate on each somite, right side membranous. Maximum cl. 200 mm (Pl. 31c). Shelf–bathyal (70–3207 m). Cosmopolitan. 13 species (Ahyong 2010: key to 4 species from Ross Sea, Australia and New Zealand; Macpherson 1988: rediagnosis, key to 6 Atlantic species).

Diagnosis. Carapace subtriangular, wider than long, widest at posterior margin; with blunt granular dorsal protuberances, deep fissures and short marginal spines. Rostrum apex undivided. Pleonite 2 of 5 plates (one median, paired submedians and marginal). Pleonites 3–5 submedian plates evenly calcified, without concave, membranous areas; marginal and submedian plates fused. Maximum cl. 60 mm (Pl. 31g). Subtidal, shelf (6–102 m). Temperate Northern E Pacific. 1 species (Makarov 1938: rediagnosis; Schmitt 1921: rediagnosis).

Paralithodes Brandt, 1848 Diagnosis. Carapace pyriform to subpentagonal, slightly wider than long; with numerous granules or spines; branchial margins not expanded laterally. Rostrum comprised of distally undivided median spine and pair of dorsal spines at base. Thoracic sternite 5 (pereopods 2) with deep, longitudinal, median fissure. Pleonite 2 of 5 plates (one median, paired submedians and marginal). Pleonites 3–5 in both sexes with blunt, calcified nodules medially; submedian and marginal plates well defined on both sides. Maximum cl. 230 mm (Pl. 31d). Intertidal–slope (0–500 m). Temperate Northern Pacific. 5 species (Makarov 1938: rediagnosis, key to 3 species; Schmitt 1921: rediagnosis, key to 2 species).

Paralomis White, 1856 Diagnosis. Carapace pyriform to subhexagonal, slightly longer than wide to wider than long; surface and margins variously ornamented. Rostrum short, conical, extending beyond eyes by at most half eye-length, formed by basal spine and 1 or more pairs of dorsal spines. Pleonite 2 of single plate. Pleonites 3–5 submedian plates evenly calcified, without concave, membranous areas, not fused with marginal plates. Maximum cl. 140 mm (Pl. 31e, f). Intertidal–abyssal (0–4152 m). Cosmopolitan. 71 species (Ahyong 2010: key to 11 species from Ross Sea, New Zealand and Australia; 2020: 3 new species; Macpherson 1988: rediagnosis, key to 16 Atlantic species; Macpherson 1992: key to 12 species from eastern Pacific; Macpherson 2003: key to 22 species from Western and Central Pacific).

Phyllolithodes Brandt, 1848 Diagnosis. Carapace subtriangular, slightly wider than long; with blunt granular dorsal protuberances and long, blunt marginal

Sculptolithodes Makarov, 1934 Diagnosis. Carapace subtriangular, as wide as or wider than long; with blunt granular dorsal protuberances, deep fissures and short marginal spines. Rostrum thick, blunt, linguiform in dorsal view, extending anteriorly beyond eyes by at least length of eye. Scaphocerite absent or rudimentary. Pleonite 2 of single plate. Pleonites 3–5 submedian plates evenly calcified, without concave, membranous areas; median plate subdivided. Maximum cl. 30 mm. Shelf (20–35 m). Temperate Northern W Pacific. 1 species (Makarov 1938: rediagnosis; Sakai 1971: photograph). References Ahyong ST (2010) The marine fauna of New Zealand: king crabs of New Zealand, Australia and the Ross Sea (Crustacea: Decapoda: Lithodidae). NIWA Biodiversity Memoir 123, 1–194. Ahyong ST (2020) First king crabs from Papua New Guinea (Crustacea: Decapoda: Lithodidae) In: Corbari, L., Chan, T-Y and Ahyong, ST (eds) Papua New Guinea. Tropical Deep-Sea Benthos vol. 31. Mémoires du Muséum National d’Histoire Naturelle, Paris 213, 121–140. Ahyong ST, O’Meally D (2004) Phylogeny of the Decapoda Reptantia: resolution using three molecular loci and morphology. Raffles Bulletin of Zoology 52, 673–693. Ahyong ST, Schnabel KE, Maas EW (2009) Anomuran phylogeny: new insights from molecular data. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 399–414. CRC Press, Boca Raton. Ahyong ST, Macpherson E, Chan T-Y (2010) Part II. Lithodoidea (king crabs). In Crustacean fauna of Taiwan: Crab-like anomurans (Hippoidea, Lithodoidea and Porcellanidae). (Ed. Chan T-Y) pp. 42–66. National Taiwan Ocean University, Keelung. Bouvier EL (1894) Sur la transformation des paguriens en crabes anomures de la sous-famille des Lithodinés. Comptes Rendus Hebdomadaires des Séances de l’Académie des Sciences 119, 350–352. Bracken HD, Toon A, Felder DL, Martin JW, Finley M, et  al. (2009) The decapod tree of life: compiling the data and moving toward a consensus of decapod evolution. Arthropod Systematics & Phylogeny 67, 99–116. Chevaldonné P, Olu K (1996) Occurrence of anomuran crabs (Crustacea: Decapoda) in hydrothermal vent and cold-seep communities: a review. Proceedings of the Biological Society of Washington 109, 286–298.

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

Cunningham CW, Blackstone NW, Buss LW (1992) Evolution of king crabs from hermit crab ancestors. Nature 355, 539–542. doi:10.1038/​ 355539a0 de Saint Laurent M, Macpherson E (1997) Une nouvelle espèce du genre Paralomis White, 1856, des sources hydrothermales du Sud-ouest Pacifique (Crustacea, Decapoda, Lithodidae). Zoosystema 19, 721–727. Haig J (1974) Observations on the lithodid crabs of Peru, with description of two new species. Bulletin of the Southern California Academy of Sciences 73, 152–164. Keiler J, Richter S, Wirkner CS (2015) The anatomy of the king crab Hapalogaster mertensii Brandt, 1850 (Anomura: Paguroidea: Hapalogastridae) – new insights into the evolutionary transformation of hermit crabs into king crabs. Contributions to Zoology 84, 149–165. doi:10.1163/18759866-08402004 Keiler J, Wirkner CS, Richter S (2017) One hundred years of carcinization – the evolution of the crab-like habitus in Anomura (Arthropoda: Crustacea). Biological Journal of the Linnean Society 121, 200–222. doi:10.1093/biolinnean/blw031 Macpherson E (1988) Revision of the family Lithodidae Samouelle, 1819 (Crustacea, Decapoda, Anomura) in the Atlantic Ocean. Monografías de Zoología Marina 2, 9–153. Macpherson E (1990) Crustacea Decapoda: on some species of Lithodidae from the Western Pacific. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 6. Mémoires du Muséum National d’Histoire Naturelle, Paris 145, 217–226. Macpherson E (1992) Paralomis phrixa (Decapoda, Anomura, Lithodidae), a new species from northern Peru, and a key to the eastern Pacific species of the genus. Crustaceana 63, 313–317. doi:10.1163/​156854092X00479 Macpherson E (1994) Occurrence of two Lithodid Crabs (Crustacea: Decapoda: Lithodidae) in the cold seep zone of the south Barbados Accretionary Prism. Proceedings of the Biological Society of Washington 107, 465–468. Macpherson E (2003) Some lithodid crabs (Crustacea: Decapoda: Lithodidae) from the Solomon Islands (SW Pacific Ocean), with the description of a new species. Scientia Marina 67, 413–418. doi:10.3989/ scimar.2003.67n4413 Macpherson E (2004) A new species and new records of lithodid crabs (Crustacea: Decapoda: Lithodidae) from the Crozet and Kerguelen Islands area (Subantarctica). Polar Biology 27, 418–422. doi:10.1007/ s00300-004-0609-1 Macpherson E, Chan T-Y (2008) Some lithodid crabs (Crustacea: Decapoda: Lithodidae) from Taiwan and adjacent waters, with the description of one new species from Guam. Zootaxa 1924, 43–52. doi:10.11646/zootaxa.1924.1.2 Macpherson E, Wehrtman I (2010) Occurence of lithodid crabs (Decapoda, Lithodidae) on the Pacific coast of Costa Rica, Central America. Crustaceana 82, 143–151. Makarov VV (1938) Crustacea. Anomura (Translation by Israel Program for Scientific Translation 1962). Fauna SSSR 10, 1–278, pls 1–5. Martin JW, Sanchez C, Pereyra R (1997) Notes on the distribution of two lithodid crabs (Crustacea: Decapoda: Anomura) from off the coast of Baja California Sur, Mexico. Bulletin of the Southern California Academy of Sciences 96, 78–86. McLaughlin PA (2003) Illustrated keys to families and genera of the superfamily Paguroidea (Crustacea: Decapoda: Anomura), with diagnoses of genera of Paguridae. Memoirs of Museum Victoria 60, 111–144. doi:10.24199/j.mmv.2003.60.16 McLaughlin PA, Lemaitre R (1997) Carcinization in the Anomura – fact or fiction? I. Evidence from adult morphology. Contributions to Zoology 67, 79–123. doi:10.1163/18759866-06702001 McLaughlin PA, Lemaitre R, Tudge CC (2004) Carcinization in the Anomura  – fact or fiction? II. Evidence from larval, megalopal and

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early juvenile morphology. Contributions to Zoology 73, 165–205. doi:10.1163/18759866-07303001 McLaughlin PA, Lemaitre R, Sorhannus U (2007) Hermit crab phylogeny: a reappraisal and its “fall-out”. Journal of Crustacean Biology 27, 97–115. doi:10.1651/S-2675.1 McLaughlin PA, Komai T, Lemaitre R, Rahayu DL (2010) Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea) Part I – Lithodoidea, Lomisoidea and Paguroidea. Raffles Bulletin of Zoology Supplement 23, 5–107. Morrison CL, Harvey AW, Lavery S, Tieu K, Huang Y, et al. (2002) Mitochondrial gene rearrangements confirm the parallel evolution of the crab-like form. Proceedings. Biological Sciences 269, 345–350. doi:10.1098/​rspb.2001.1886 Retamal MA (1994) Los Lithodidae Chilenos The Chilean Lithodidae. Anales del Instituto de la Patagonia, Serie. Ciência e Natura 21, 111–129. Richter S, Scholtz G (1994) Morphological evidence for a hermit crab ancestry of lithodids (Crustacea, Decapoda, Anomala, Paguroidea). Zoologischer Anzeiger 233, 187–210. Sakai T (1971) Illustrations of 15 species of crabs of the family Lithodidae, two of which are new to science. Researches on Crustacea 4.5, 1–49, pls 1–21. Schmitt WL (1921) The marine decapod Crustacea of California with special reference to the decapod Crustacea collected by the United States Bureau of Fisheries Steamer Albatross in connection with the biological survey of San Francisco Bay during the years 1912–1913. University of California Publications in Zoology 23, 1–359, pls 1–50. Schnabel KE, Ahyong ST, Maas EL (2011) Galatheoidea are not monophyletic  – molecular and morphological phylogeny of the squat lobsters (Decapoda: Anomura) with recognition of a new superfamily. Molecular Phylogenetics and Evolution 58, 157–168. doi:10.1016/j.ympev.2010.​ 11.011 Stevens BG (Ed.) (2014) King crabs of the world: biology and fisheries management. CRC Press, Boca Raton. Tsang LM, Ma KY, Ahyong ST, Chan T-Y, Chu KH (2008) Phylogeny of Decapoda using two nuclear protein-coding genes: Origin and evolution of the Repantia. Molecular Phylogenetics and Evolution 48, 359–368. doi:10.1016/j.ympev.2008.04.009 Wolfe JM, Breinholt JW, Crandall KA, Lemmon AR, Lemmon EM, et  al. (2019) A phylogenomic framework, evolutionary timeline and genomic resources for comparative studies of decapod crustaceans. Proceedings. Biological Sciences 286, 20190079. doi:10.1098/rspb.2019.0079

Paguridae Latreille, 1802 right-handed hermit crabs Figures 13.23–13.27, Plates 33 g–l, 34a–g The Paguridae have the reputation for being one of the most difficult of all decapod families for the general taxonomist, and consequently, have attracted few, but very active, specialists in recent years. At family level, recognition is usually straightforward: right-handed hermit crabs from shallow water are most likely to be pagurids. Identifying genera, however, is much less easy, requiring close attention to gill arrangements. The extensive literature and the enormous number of undescribed species, especially in the tropical Indo-West Pacific, are discouraging for the uninitiated. Gordan (1956) compiled a checklist of species and

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comprehensive bibliography but this has been superseded by another by McLaughlin et  al. (2010) who listed ~600 species. Present-day systematics of the family builds on the works of Henderson (1888) and Alcock (1905). More recent work by M. de Saint Laurent, J. Forest, P. McLaughlin, R. Lemaitre and T. Komai especially have improved understanding of the family’s diversity. McLaughlin’s (1997, 2003a) keys and generic diagnoses provided a way into the literature but these dealt with only 71 of the 86 genera now recognised. De Saint Laurent-Dechancé (1966) divided the family into three groups based largely on the number of thoracic gills or branchiae, 10, 11 or 13 on each side, but within some genera gill number varies. The position of gills is sometimes referred to by somite number. Thoracic somites 1–8 carry maxillipeds 1–3 and pereopods 1–5 – limb number is used here (Fig. 13.25a). Gills are referred to as biserial (with two lobes attached to the axial rachis) (Fig. 13.3 3b) or quadriserial (with four lobes) (Fig. 13.25e). Intermediate conditions are referred to as distally quadriserial, as in Goreopagurus (Fig.  13.25c) but similar arrangements may also be called fully quadriserial, as in Propagurus (Fig. 13.25d) (­McLaughlin and de Saint Laurent 1998). The 11-gilled Pagurus-group (each side with two arthrobranchs each on maxilliped 3 to pereopod 4, one pleurobranch above pereopod 4) comprises the majority of genera. A subgroup in which females possess paired first pleopods was reviewed in a series of papers beginning with ­McLaughlin (1981a). Enneobranchus and two similar genera have nine gills, differing from others in this group in lacking arthrobranchs from maxilliped 3. The 10-gilled Ostraconotus-group (each side with two arthrobranchs each on maxilliped 3 to pereopod 4, without pleurobranchs) was revised by de Saint Laurent in a series of six papers concluding with de Saint Laurent (1970). In some species of Decaphyllus the arthrobranchs above maxilliped 3 can be vestigial or absent.

The 13-gilled Pylopaguropsis-group (each side with two arthrobranchs each on maxilliped 3 to pereopod 4, one pleurobranch above pereopods 2–4) comprises few genera (Fig. 13.25a). This simple division into three groups is complicated by the gill loss and reductions mentioned above and others. Both Paguriscus and Paguruncio have eight pairs of gills but not the same pairs (see diagnoses below). More recently considerable emphasis has been placed on the structure of male sexual tubes, protrusions from the gonopore on one or other or both of the coxae of pereopods 5 (Lemaitre and McLaughlin 2003a; Tudge and Lemaitre 2004). Orientation of the sexual tubes is diagnostic at a generic level but so too is size (Fig.  13.26). McLaughlin (2003a) defined descriptive terms to quantify tube length, summarised here as short (< 2 times coxal length), moderate (2–5 times coxal length) and long (> 5 times coxal length). Dimensions are given as shield length (sl.). Pereopod 1 is the cheliped, pereopods 2 and 3 the two pairs of walking legs, and pereopods 4 and 5 are short. The key below was modified from that of McLaughlin (2003a), incorporating the update of a section by Komai (2013b) and was updated further by rearrangement and adding recent genera. Since going to press a new genus, Vounopagurus Komai, Rahayu & Fujita, 2021, similar to Kumepagurus, has been erected (Komai et al. 2021). Diagnosis. Pleonite 1 distinct from last thoracic somite. Right cheliped larger than left (Fig.  13.23a). Maxilliped 1 exopod with flagellum (Fig.  13.25f). Maxillipeds 3 widely separate at base (Fig. 13.25g). Pleopods 3–5 on left only, or pleopods 1 (sometimes pleopods 2) paired. Implicit generic attributes. Gills: biserial, 11 pairs. Ocular acicles simple. Pereopod 4 semichelate, propodal rasp with 1 row of corneous scales, preungual process absent. Male pereopodal coxae 5 symmetrical, without sexual tubes; unpaired pleopods 3–5 present. Female gonopore on both coxae 3; pleopods 1 absent; with unpaired pleopods 2–5. Uropods asymmetrical.

Key to genera of Paguridae 1. – 2. – 3.

Crab-like, shield plus broad rostrum as wide or wider than long. Anterior carapace vaulted, well calcified (Figs 13.23f–i). Pleon reduced, symmetrical. Using bivalve or limpet shells���������������������������������������������������������������������2 Hermit crab-like, shield rarely well calcified, rostrum not prominent. Anterior carapace not vaulted, rarely calcified. Pleon rarely reduced, usually twisted. Using gastropod, scaphopod or pteropod shells, or wood�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������5 Carapace anterior and posterior sections lateral margins with row of 8–10 similar teeth. Rostrum short, rounded (Fig. 13.23f). Pereopod 4 propodus broad, discoid (Fig. 13.27B)��������������������������������������������������� Ostraconotus Carapace anterior and posterior sections usually with few prominent teeth. Rostrum prominent, triangular or truncate. Pereopod 4 propodus linear���������������������������������������������������������������������������������������������������������������������������������3 Carapace lateral margins with 2 blunt lobes or none; cervical groove almost divides posterior carapace in half; posterior carapace lobes prominent, laterally projecting (Fig. 13.23g)������������������������������������������������������Patagurus

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters



319

Carapace lateral margins with 2 subacute lobes; cervical groove anterior of posterior carapace; posterior carapace lobes not or scarcely projecting��������������������������������������������������������������������������������������������������������������������������������4 4. Carapace lateral margins of shield with 3 blunt or spiniform lobes; posterior carapace lobe not laterally expanded. Rostrum broadly triangular (Fig. 13.23i)������������������������������������������������������������������ Solitariopagurus – Carapace lateral margins of shield developed into 2 blunt or spiniform, wing-like projections; posterior carapace lobes prominent. Rostrum triangular or truncate (Fig. 13.23h)������������������������� Porcellanopagurus 5. Pleurobranchs above pereopods 2–4 (13 gills)�����������������������������������������������������������������������������������������������������������������������������6 – Fewer than 3 pleurobranchs (8–11 gills)�������������������������������������������������������������������������������������������������������������������������������������15 6. Pleurobranchs above pereopods 2 and 3 vestigial�����������������������������������������������������������������������������������������������������������������������7 – Pleurobranchs all well developed (rarely pleurobranchs above pereopods 2 and 3 absent)�������������������������������������������������8 7. Cornea fully formed. Chelipeds markedly unequal. Female with paired gonopores����������������������������������������� Propagurus – Cornea reduced, distoventrolateral. Chelipeds subequal. Female with left gonopore only�������������������������Chanopagurus 8. Males without unpaired pleopods. Pleonite 6 strongly calcified������������������������������������������������������������������������ Lithopagurus – Male with unpaired pleopods 2–4 or 2–5. Pleonite 6 membranous�����������������������������������������������������������������������������������������9 9. Males with paired, modified pleopods 1 or 2. Gills quadriserial�������������������������������������������������������������������������������������������10 – Males without paired, modified pleopods. Gills biserial or quadriserial������������������������������������������������������������������������������11 10. Males with pleopods 2 paired, modified (Fig. 13.26F); pleopods 3–5 unpaired�����������������������������������������Tomopaguroides – Males with pleopods 1 paired, modified, or absent; pleopods 2–5 unpaired���������������������������������������������� Tomopaguropsis 11. Right cheliped much larger than left, with massive operculate or semioperculate chela (Fig. 13.24e, n)����������������������� 12 – Cheliped subequal or unequal, chela not massive, not operculate (Fig. 13.24f)�������������������������������������������������������������������14 12. Male with coiled right sexual tube and short left sexual tube (Fig. 13.26j). Maxilliped 3 without accessory tooth on inner face�����������������������������������������������������������������������������������������������������������������������������Kumepagurus – Male without sexual tube on left or right coxae 5 (Fig. 13.26A). Maxilliped 3 with accessory tooth on inner face (Fig. 13.25h)������������������������������������������������������������������������������������������������������������������������������������������������������������������������13 13. Gills biserial (Fig. 13.25b). Female pleopods 1 paired, modified. Male pleopods 3–5 unpaired���������������� Pylopaguropsis – Gills quadriserial (Fig. 13.25e). Female pleopods 1 absent. Male pleopods 3–4 unpaired������������������������Bathypaguropsis 14. Posterior carapace median element fused with lateral lobes, posteromedian and posterolateral plates well calcified at least in anterior halves (Fig. 13.23e). Uropod peduncle without elongate spine. Telson posterior margins oblique, spinose (Fig. 13.27d)�������������������������������������������������������������������������������������������� Bythiopagurus – Carapace not calcified. Uropod peduncle with elongate spine. Telson entire (Fig. 13.27b). Chelipeds exceptionally elongate������������������������������������������������������������������������������������������������������������������������������������� Munidopagurus 15. Without pleurobranchs. Male coxae 5 trapezoidal, extending posteriorly to form short paired sexual tubes (Fig. 13.26q)�����������������������������������������������������������������������������������������������������������������������������������������������������������������Paguriscus – With at least 1 pleurobranch. Male coxae not extended, sexual tube independent if present�������������������������������������������16 16. Pleurobranch absent above pereopod 4 (10 gills, sometimes vestigial)���������������������������������������������������������������������������������17 – Pleurobranch present above pereopod 4 (11 or fewer gills)�����������������������������������������������������������������������������������������������������19 17. Crista dentata comprising 2–4 spines (Fig. 13.25i). Eyestalks tapering (Fig. 13.23r)���������������������������������� Scopaeopagurus – Crista dentata a well developed or reduced ridge. Eyestalks with expanded cornea�����������������������������������������������������������18 18. Male right pereopodal coxa 5 with to long sexual tube, recurved anteriorly under thorax. Left sexual tube usually partially obscured by tufts of sternal setae (Fig. 13.26b). Male with 3 unpaired pleopods�������������������������������������������������������������������������������������������������������������������������������������������� Catapaguroides – Male right pereopodal coxa 5 with sexual tube orientated across ventral body surface. Left sexual tube not obscured by tufts of sternal setae (Fig. 13.26e). Males with 4 unpaired pleopods���������������������������������������� Decaphyllus 19. Arthrobranchs vestigial or absent on maxilliped 3����������������������������������������������������������������������������������������������������������������� 20 – Arthrobranchs well developed on maxilliped 3����������������������������������������������������������������������������������������������������������������������� 28 20. Male right pereopod coxae 5 with slender sexual tubes, of similar or different lengths (Fig. 13.26n, u, y)���������������������21 – Male right pereopod coxa 5 without sexual tube, rarely with papilla; left with short or long sexual tube��������������������� 23 21. Male coxae 5 with pair of subequal, slender, membranous sexual tubes directed posteriorly (Fig. 13.26u)��������� Paguruncio

320

Marine Decapod Crustacea

– Male coxae 5 with pair of unequal slender sexual tubes��������������������������������������������������������������������������������������������������������� 22 22. Male with pair of basally weakly calcified, distally filamentous sexual tubes, longer and coiled on left (Fig. 13.26n). Maxilliped 3 with 1 accessory tooth����������������������������������������������������������������������������������������������� Pagurellus – Male with pair of similar tapering sexual tubes, longer and coiled on right (Fig. 13.26y). Maxilliped 3 without accessory tooth������������������������������������������������������������������������������������������������������������� Pusillopagurus 23. Male left pereopod coxa 5 sexual tube short; without sexual tube on right (Fig. 13.26p)�������������������������������������� Pagurina – Male left pereopod coxa 5 sexual tube long; without or with sexual tube or papilla on right (Fig. 13.26h, i, t, C)�������������� 24 24. Rostrum strongly deflected, with prominent dorsal spine (Fig. 13.23w). Gills biserial���������������������������������� Enneophyllus – Rostrum not strongly deflected, without prominent dorsal spine. Gills quadriserial, at least distally��������������������������� 25 25. Male left pereopod coxa 5 sexual tube without terminal fringe of setae (Fig. 13.26h). Pereopod 4 with preungual process (Fig. 13.27z)������������������������������������������������������������������������������������������������������������������������Enneobranchus – Male left pereopod coxa 5 sexual tube with terminal dense fringe of setae. Pereopod 4 without preungual process�������������������������������������������������������������������������������������������������������������������������������������������������������������������� 26 26. Male left pereopod coxa 5 with moderate to long, weakly spiralled sexual tube (Fig. 13.26D)����������������������������Turleania – Male left pereopod coxa 5 with club-like, stout, short to moderate sexual tube directed towards exterior������������������� 27 27. Male left pereopod coxa 5 with club-like, stout sexual tube directed towards exterior, with terminal tuft of long setae (Fig. 13.26t). Female pleopods 1 paired���������������������������������������������������������Pagurojacquesia – Male left pereopod coxa 5 with moderate, stout sexual tube dorsally-exteriorly-directed, terminally spatulate with fringe of short setae (Fig. 13.26i). Female pleopods 1 absent��������������������������������������������� Enneopagurus 28. Telson posterior lobes with long setae on distal lateral margins (Fig. 13.27j, t)������������������������������������������������������������������ 29 – Telson posterior lobes without long setae on distal lateral margins������������������������������������������������������������������������������������� 30 29. Gills quadriserial. Eyestalk tapering, cornea reduced (Fig. 13.23l)�������������������������������������������������������������� Eutrichopagurus – Gills biserial. Eyestalks not tapering, cornea full width (Fig. 13.23s)�������������������������������������������������������������� Trichopagurus 30. Gills quadriserial����������������������������������������������������������������������������������������������������������������������������������������������������������������������������31 – Gill biserial��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������37 31. Maxilliped 3 crista dentata with 1 or more accessory teeth on inner face (Fig. 13.25h)�����������������������������������������������������32 – Maxilliped 3 crista dentata without accessory tooth���������������������������������������������������������������������������������������������Iridopagurus 32. Chelipeds subequal (Fig. 13.24a, b)���������������������������������������������������������������������������������������������������������������������������������������������33 – Chelipeds distinctly unequal, right larger (Fig. 13.24c, d)������������������������������������������������������������������������������������������������������ 36 33. Females with paired, modified pleopod 1��������������������������������������������������������������������������������������������������������������������������������� 34 – Females without paired, modified pleopod 1����������������������������������������������������������������������������������������������������������������������������35 34. Male with unpaired pleopods 3–5. Telson posterior lobes posterolaterally expanded, separated by narrow spinose median cleft (Fig. 13.27n). Inhabiting gastropod shells������������������������������������������������� Michelopagurus – Male with unpaired pleopods 2–5. Telson posterior lobes parallel-sided, separated by broad spinose V-shaped cleft (Fig. 13.27e). Inhabiting scaphopod shells������������������������������������������������������������������������� Dentalopagurus 35. Rostrum triangular (Fig. 13.23n). Ambulatory legs dactyli almost straight, slightly thickened subdistally (Fig. 13.27u)������������������������������������������������������������������������������������������������������������������������������������������������������������������Pagurodes – Rostrum broadly rounded (Fig. 13.23o). Ambulatory legs dactyli evenly curved, evenly tapering to corneous tip (Fig. 13.27v)������������������������������������������������������������������������������������������������������������������������������ Pseudopagurodes 36. Male with short left sexual tube (Figs 13.26C). Female with pleopods 1paired, modified�������������������������Tarrasopagurus – Males with long right sexual tube (Figs 13.26c). Female without pleopods 1 paired, modified��������������������Cestopagurus 37. Male with 1 or 2 sexual tubes or papillae���������������������������������������������������������������������������������������������������������������������������������� 38 – Males without sexual tubes, rarely with one or pair of papillae�������������������������������������������������������������������������������������������� 63 38. Chelipeds subequal; both slender, with clusters of elongate, capsulate setae (thickened proximally, tapering to fine tip) on lower face of merus��������������������������������������������������������������������������������������������������������������������� Leptopagurus – Chelipeds unequal or subequal, neither with capsulate setae on lower face of merus������������������������������������������������������� 39 39. Females with pleopods 1 paired, modified������������������������������������������������������������������������������������������������������������������������������� 40 – Females without pleopods 1 paired, modified���������������������������������������������������������������������������������������������������������������������������41

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

321

40. Male right sexual tube long, elongate, extremely filiform distally (Fig. 13.26d). Right cheliped carpus lower margin not produced������������������������������������������������������������������������������������������������������������������������������������������Cycetopagurus – Male right sexual tube short. Right cheliped carpus lower margin strongly produced (Fig. 13.24h)�����������������������������������������������������������������������������������������������������������������������������������������������������������Goreopagurus 41. Pereopod 4 propodal rasp with 2 or more rows of scales (Fig. 13.27A)�������������������������������������������������������������������������������� 42 – Pereopod 4 propodal rasp with 1 row of scales (Fig. 13.27C, D) or rudimentary or absent���������������������������������������������� 45 42. Ocular acicles multispinose (Fig. 13.23u). Right gonopore present or absent, without right sexual tube, with moderate to long left sexual tube (Fig. 13.26l)���������������������������������������������������������������������������������������� Micropagurus – Ocular acicles simple or with submarginal spine (Fig. 13.23t). Right sexual tube present, left sexual tube very short at most����������������������������������������������������������������������������������������������������������������������������������������� 43 43. Right sexual tube, straight or slightly curved, laterally-posterolaterally directed, proximal half stout, opaque, distal portion transparent (Fig. 13.26a)���������������������������������������������������������������������������������������������Boninpagurus – Right sexual tube short (Fig. 13.26B)����������������������������������������������������������������������������������������������������������������������������������������� 44 44. Telson posterior lobes transverse or oblique, microspinose (Fig. 13.27p). Coxae 5 symmetrical����������������Parapagurodes – Telson posterior lobes separated by broad median cleft (Fig. 13.27o). Coxae 5 asymmetrical (Fig. 13.26s)������������������������������������������������������������������������������������������������������������������������������������������������������� Pagurodofleinia 45. Chela dorsoventrally flattened, spatulate (Fig. 13.24o). Telson lobes asymmetrical, with dentate margins (Fig. 13.27s)������������������������������������������������������������������������������������������������������������������������������������������ Spathapagurus – Chela not spatulate. Telson margins variously ornamented��������������������������������������������������������������������������������������������������� 46 46. Male with single sexual tube (if on right side, rarely with papilla on left coxa 5)��������������������������������������������������������������� 47 – Male with paired sexual tubes or papillae��������������������������������������������������������������������������������������������������������������������������������� 58 47. Male with left sexual tube������������������������������������������������������������������������������������������������������������������������������������������������������������ 48 – Male with right sexual tube����������������������������������������������������������������������������������������������������������������������������������������������������������52 48. Right chela not markedly larger than left��������������������������������������������������������������������������������������������������������������� Spiropagurus – Right chela markedly larger than left (Fig. 13.24m)���������������������������������������������������������������������������������������������������������������� 49 49. Telson with terminal margins armed with spines (Fig. 13.27q)��������������������������������������������������������������������������������������������� 50 – Telson with terminal margins unarmed�������������������������������������������������������������������������������������������������������������������������������������51 50. Eyestalk expanding to wide cornea (Fig. 13.23j). Eastern Atlantic (except A. japonicus)��������������������������������� Anapagurus – Eyestalk tapering to reduced cornea (Fig. 13.23p). Central Indo-Pacific�������������������������������������������������������Pumilopagurus 51. Telson with terminal margin entire (Fig. 13.27k). Cornea strongly dilated (Fig. 13.23m)��������������������������Forestopagurus – Telson with terminal margin with prominent median cleft (Fig. 13.27r). Cornea reduced (Fig. 13.23q)������������������������������������������������������������������������������������������������������������������������������������������������������Pygmaeopagurus 52. Female with single gonopore on left coxa��������������������������������������������������������������������������������������������������������������� Anapagrides – Female with paired gonopores�����������������������������������������������������������������������������������������������������������������������������������������������������53 53. Males with 4 unpaired pleopods�������������������������������������������������������������������������������������������������������������������������Acanthopagurus – Male with 0–3 unpaired pleopods���������������������������������������������������������������������������������������������������������������������������������������������� 54 54. Right sexual tube up to 5 times coxal length�����������������������������������������������������������������������������������������������������������������������������55 – Right sexual tube 5 or more times coxal length����������������������������������������������������������������������������������������������������������������������� 56 55. Pereopod 4 propodus with rasp in female, without in male (Fig. 13.27w, x). Maxilliped 3 crista dentata reduced, without accessory tooth����������������������������������������������������������������������������������������������������� Catapaguropsis – Pereopod 4 propodal rasp present in both sexes. Maxilliped 3 crista dentata serrate, with 1 accessory tooth�������������������������������������������������������������������������������������������������������������������������������������������Catapagurus 56. Sexual tubes terminating in elongate filament (Fig. 13.26m)���������������������������������������������������������������������Nematopaguroides – Sexual tube not terminating in elongate filament (Fig. 13.26w)���������������������������������������������������������������������������������������������57 57. Left pereopod 3 propodus and dactylus dissimilar from other pereopods, with plumose setae on lateral faces�����������������������������������������������������������������������������������������������������������������������������������������������������������Solenopagurus – No pereopods with plumose setae on lateral faces. Inhabiting pteropod shells����������������������������������������������� Pteropagurus 58. Pleon reduced. Male without pleopods. Female with pleopods 2–4 unpaired, uniramous�����������������������������������������������59

322

Marine Decapod Crustacea



Pleon well developed. Male usually with unpaired pleopods. Female with pleopods 2–4 unpaired, biramous, usually with reduced pleopod 5�������������������������������������������������������������������������������������������������������������������������� 60 59. Rostrum prominent, slender spine (Fig. 13.23c). Pereopod 5 subchelate��������������������������������������������������������Alainopagurus – Rostrum broad, blunt, upturned (Fig. 13.23b). Pereopod 5 weakly chelate�����������������������������������������������Alainopaguroides 60. Females with pleopods 1 paired, modified��������������������������������������������������������������������������������������������������������Nematopagurus – Females without pleopods 1 paired, modified���������������������������������������������������������������������������������������������������������������������������61 61. Antennal scaphocerite with row of spines on mesial face (Fig. 13.23d)�������������������������������������������������������Alloeopagurodes – Antennal scaphocerite without row of spines on mesial face��������������������������������������������������������������������������������������������������62 62. Telson posterior lobes rounded, armed with long, slender, corneous spines (Fig. 13.27m). Antennal flagellar articles with paired long setae armed with setules (Fig. 13.25k)�������������������������������������������������������������������� Icelopagurus – Telson posterior lobes subtriangular, armed with spines or spinules (Fig. 13.27l). Antennal flagellar articles with short setae��������������������������������������������������������������������������������������������������������������������������������������������������Hachijopagurus 63. Female with pleopods 1 paired, modified��������������������������������������������������������������������������������������������������������������������������������� 64 – Female without pleopods 1 paired, modified��������������������������������������������������������������������������������������������������������������������������� 79 64. Chelipeds unequal, right elongate, chela about twice as long as wide (Fig. 13.24g)��������������������������������������Ceratopagurus

Fig. 13.23.  Paguridae. a, Labidochirus splendescens (Owen, 1839). Shield, eyestalks, antennae: b, Alainopaguroides; c, Alainopagurus; d, Alloeopagurodes. Carapace: e, Bythiopagurus. Shield: f, Ostraconotus; g, Patagurus rex Anker & Pauly, 2013.; h, Porcellanopagurus; i, Solitariopagurus. Front, eyestalks, ocular acicles: j, Anapagurus; k, Chanopagurus; l, Eutrichopagurus; m, Forestopagurus; n, Pagurodes; o, Pseudopagurodes; p, Pumilopagurus; q, Pygmaeopagurus; r, Scopaeopagurus; s, Trichopagurus. Front, ocular acicles: t, Boninpagurus; u, Micropagurus; v, Pylopaguridium. Rostrum, dorsal, lateral views: w, Enneophyllus.

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323

Fig. 13.24.  Paguridae. Left and right subequal chelipeds to same scale: a, b, Iridopagurus. Left and right unequal chelipeds to same scale: c, d, Cestopagurus caeruleus Komai & Poupin, 2012. Right chela: e, Bathypaguropsis; f, Bythiopagurus; g, Ceratopagurus; h, Goreopagurus; i, Lithopagurus; j, Lophopagurus (Australeramus); k, Lophopagurus (Lophopagurus); l, Protoniopagurus; m, Pumilopagurus; n, Pylopagurus; o, Spathapagurus; p, Tomopagurus. Left chela: q, Anisopagurus; r, Lophopagurus (Lophopagurus); s, Haigiopagurus; t, Manucomplanus; u, Pliopagurus; v, Tomopagurus. Cheliped merus, with capsulate setae: w, Leptopagurus. Chela, ornamented tubercles: x, Agaricochirus (mushroom-shaped tubercles); y, Rhodochirus (with basal rosettes).

Fig. 13.25.  Paguridae. a, Gills on left side (mp3, c1–c4 denote coxae of maxilliped 3 and pereopods 1–4; 10 arthrobranchs, 2 per somite (dark grey) and 3 pleurobranchs above pereopods 2–4 (light grey) are shown); b, section of biserial gill c, section of distally quadriserial gill (Goreopagurus), d, e, sections of quadriserial gills (Propagurus, Pagurojacquesia). Maxilliped 1: f, Pagurus. Maxilliped 3: g, Pagurus (pair); h, Pylopaguropsis; i, Scopaeopagurus (ischium, merus); j, Kumepagurus (ischium). Antennal flagellum: k, Paguritta.

324

Marine Decapod Crustacea

Fig. 13.26.  Paguridae. Male pereopodal coxae 5 with sexual tubes (anterior uppermost): a, Boninpagurus; b, Catapaguroides; c, Cestopagurus; d, Cycetopagurus; e, Decaphyllus; f, Dentalopagurus; g, Diacanthurus; h, Enneobranchus; i, Enneopagurus; j, Kumepagurus; k, Leptopagurus; l, Micropagurus; m, Nematopaguroides; n, Pagurellus; o, Paguridium; p, Pagurina; q, Paguriscus; r, Pagurixus; s, Pagurodofleinia; t, Pagurojacquesia; u, Paguruncio; v, Pliopagurus; w, Pteropagurus; x, Pumilopagurus; y, Pusillopagurus; z, Pylopaguridium; A, Pylopaguropsis; B, Spathapagurus; C, Tarrasopagurus; D, Turleania. Female pereopodal coxae 5, pleopods 1: E, Chanopagurus. Male pereopodal coxae 5, pleopods 2: F, Tomopaguroides.

– 65.

Chelipeds subequal, right more or less operculate, chela little longer than wide (Figs 13.24l, n)������������������������������������� 65 Pleon short, flexed or straight. Male without unpaired pleopods. Female with unpaired pleopods 2–4����������������������������������������������������������������������������������������������������������������������������������� Protoniopagurus – Pleon coiled. Male with some unpaired pleopods. Female with unpaired pleopods 2–5�������������������������������������������������� 66 66. Uropodal peduncle prominently produced posteriorly (Fig. 13.27a). Right chela dorsal surface covered by mushroom-shaped tubercles (Fig. 13.24x)������������������������������������������������������������������������������������������������������� Agaricochirus – Uropodal peduncle not prominently produced posteriorly. Right chela dorsal surface not covered by mushroom-shaped tubercles�������������������������������������������������������������������������������������������������������������������������������������������������� 67 67. Spines on dorsal surfaces of chelae with basal rosettes (Fig. 13.24z)������������������������������������������������������������������ Rhodochirus

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

– 68. – 69. – 70. – 71. – 72. – 73. – 74. – 75. – 76. – 77. – 78. – 79. – 80. – 81. – 82. – 83. – 84.

325

Spines on dorsal surfaces of chelae without basal rosettes����������������������������������������������������������������������������������������������������� 68 Pereopod 4 propodal rasp with 2 or more rows of corneous scales�������������������������������������������������������������������������������������� 69 Pereopod 4 propodal rasp with 1 row of corneous scales��������������������������������������������������������������������������������������������������������71 Left chela not subtriangular in cross-section, dactylus and fixed finger dorsoventrally flattened (Fig. 13.24t)������������������������������������������������������������������������������������������������������������������������������������������������������� Manucomplanus Left chela subtriangular in cross-section, dactylus and fixed finger not dorsoventrally flattened�����������������������������������70 Telson with lateral indentations (Fig. 13.27c)�������������������������������������������������������������������������������������������������������� Anisopagurus Telson without lateral indentations (Fig. 13.27h)��������������������������������������������������������������������������������������������Enallopaguropsis Ocular acicles multispinose (Fig. 13.23v). Male coxa 5 asymmetrical (Fig. 13.26z)����������������������������������� Pylopaguridium Ocular acicles simple. Male coxa 5 symmetrical���������������������������������������������������������������������������������������������������������������������� 72 Telson without lateral indentations (Fig. 13.27h)������������������������������������������������������������������������������������������������ Enallopagurus Telson with lateral indentations�������������������������������������������������������������������������������������������������������������������������������������������������� 73 Right chela subovate to subcircular, margins unarmed, weakly tuberculate or minutely crenulate and/or serrate, but never armed with prominent, blunt or acute spines (Fig. 13.24n)����������������������������������������������������������������74 Right chela of variable shape, margins armed with prominent, blunt or acute spines or tubercles (Fig. 13.24p)��������������� 75 Pereopod 4 with large, prominent preungual process at base of claw (Fig. 13.27C)����������������������������������������� Phimochirus Pereopod 4 without prominent preungual process at base of claw����������������������������������������������������������������������Pylopagurus Left cheliped dactylus and fixed finger excavated ventrally, spoon-shaped (Fig. 13.24v)������������������������������Tomopagurus Left cheliped dactylus and fixed finger acute, not excavated ventrally, spoon-shaped�������������������������������������������������������76 Left chela midline smooth, convex, lateral margin not expanded (Fig. 13.24u). Male with short, prominent sexual tubes (Fig. 13.26v)��������������������������������������������������������������������������������������Pliopagurus Left chela midline elevated into low crest or prominent keel and/or with lateral margin expanded and carinate, or spinose (Fig. 13.24r, s). Male without sexual tubes or papillae�������������������������������������������������������������������� 77 Right chela circumscribed by row of dorsomesial, dorsoproximal and dorsolateral marginal spines (Fig. 13.24j)������������������������������������������������������������������������������������������������������������������������������ Lophopagurus (Australeremus) Right chela not circumscribed by row of dorsomesial, dorsoproximal and dorsolateral spines (Fig. 13.24k)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������78 Left chela midline elevated into prominent keel or crest, lateral margin at most granulate or minutely tuberculate (Fig. 13.24s)���������������������������������������������������������������������������������������������������������Lophopagurus (Lophopagurus) Left chela midline slightly elevated, not forming prominent keel or crest, lateral margin strongly spinose..... ��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������Haigiopagurus Male pereopod 5 coxae asymmetrical (Figs 13.26o, r)������������������������������������������������������������������������������������������������������������� 80 Male pereopod 5 coxae symmetrical�������������������������������������������������������������������������������������������������������������������������������������������81 Male right pereopod 5 coxa produced, gonopore masked by tuft of long, stiff setae (Fig. 13.26r). Male with 3 unpaired pleopods���������������������������������������������������������������������������������������������������������������������������������������������������Pagurixus Male left pereopod 5 coxa produced, gonopore masked by tuft of long, stiff setae (Fig. 13.26o). Male without pleopods������������������������������������������������������������������������������������������������������������������������������������������������������Paguridium Males pereopod 5 coxae both produced, gonopores each masked by tuft of long, stiff setae. Telson terminal margin markedly concave, outer angles acute, with prominent pair of spines adjacent to median cleft (Fig. 13.27f)����������������������������������������������������������������������������������������������������������������������������������������������������������� Diacanthurus Males without coxae of pereopods 5 produced. Telson without markedly concave terminal margin, outer angles variable, without extremely prominent pair of spines adjacent to median cleft (Fig. 13.27g)������������������������ 82 Telson without or with weak transverse indentation (Fig. 13.27g)����������������������������������������������������������������Discorsopagurus Telson with distinct transverse indentation ��������������������������������������������������������������������������������������������������������������������������������3 Posterior portion of cephalothorax, at least in part calcified. Pleon reduced (Fig. 13.23a)�����������������������������Labidochirus Posterior portion of cephalothorax membranous. Pleon well developed����������������������������������������������������������������������������� 84 Left chela with pronounced anticlockwise rotation. Pereopod 4 dactylus with prominent circular sensory structure on lateral face (Fig. 13.27y)�������������������������������������������������������������������������������������������������������������������Elassochirus

326

Marine Decapod Crustacea

Fig. 13.27.  Paguridae. Uropods, telson: a, Agaricochirus; b, Munidopagurus. Telson: c, Anisopagurus; d, Bythiopagurus; e, Dentalopagurus; f, Diacanthurus; g, Discorsopagurus; h, Enallopaguropsis; i, Enneopagurus; j, Eutrichopagurus; k, Forestopagurus; l, Hachijopagurus; m, Icelopagurus; n, Michelopagurus; o, Pagurodofleinia; p, Parapagurodes; q, Pumilopagurus; r, Pygmaeopagurus; s, Spathapagurus; t, Trichopagurus. Pereopod 3, propodus, dactylus: u, Pagurodes; v, Pseudopagurodes. Pereopod 4, propodus, dactylus: w, x, Catapaguropsis, male, female; y, Elassochirus; z, Enneobranchus; A, Micropagurus; B, Ostraconotus; C, Phimochirus; D, Spathapagurus.



Left chela without anticlockwise rotation. Pereopod 4 dactylus without prominent circular sensory structure on lateral face����������������������������������������������������������������������������������������������������������������������������������������������������������� 85 85. Uropods asymmetrical. Pleon spirally flexed����������������������������������������������������������������������������������������������������������������� Pagurus – Uropods symmetrical. Pleon not spirally flexed���������������������������������������������������������������������������������������������������������������������� 86 86. Antennal flagellar articles with few short setae or naked. Male with 3 unpaired pleopods. Female with 4 unpaired pleopods�����������������������������������������������������������������������������������������������������������������������������������������������Orthopagurus – Antennal flagellar articles with paired long setae armed with setules (Fig. 13.25k). Male without unpaired pleopods. Female with 3 unpaired pleopods����������������������������������������������������������������������������������������������������������� Paguritta Acanthopagurus de Saint Laurent, 1968 Diagnosis. Rostrum obtusely and roundly triangular. Chelipeds unequal; right much stronger. Male pereopodal coxae 5 asymmetrical; right pereopodal coxa 5 with short massive sexual tube, directed obliquely towards midline. Telson posterior margins oblique. Maximum sl. 4.3 mm. Slope (225 m). Temperate Northern Atlantic. 1 species (de Saint Laurent 1969).

Agaricochirus McLaughlin, 1981 Diagnosis. Chelipeds unequal; right ovate, armed with mushroom-shaped tubercles. Ambulatory legs carpi lacking dorsodistal

spine; propodal rasp with several rows of corneous scales, preungual process small. Male right and left pereopodal coxae 5 both without sexual tube, sometimes with papilla on one side or other. Female pleopods 1 paired, modified. Uropods symmetrical; peduncle produced posteriorly. Telson with broadly U-shaped median cleft; posterior margins unarmed. Maximum sl. 8.2 mm. Shelf, slope (35–800 m). Tropical W Atlantic. 8 species (McLaughlin 1982: key to species).

Alainopaguroides McLaughlin, 1997 Diagnosis. Anterior carapace vaulted, well calcified, anterolateral regions slightly depressed, posterior carapace slightly calcified; rostrum rounded. Pleon reduced, pleonal plates 2–5 sometimes

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

faintly delineated. Chelipeds subequal; right stronger, but not longer. Propodal rasp rudimentary, preungual process prominent. Male right pereopodal coxa 5 with moderate, stout sexual tube; left pereopodal coxa 5 without sexual tube, or with short sexual tube; without unpaired pleopods. Female with unpaired pleopods 2–4. Uropods symmetrical. Telson posterior margins oblique. Maximum sl. 6.2 mm. Shelf, slope (49–502 m). Central Indo-Pacific. 4 species (Han et al. 2016a: key to species).

Alainopagurus Lemaitre & McLaughlin, 1995 Diagnosis. Anterior carapace vaulted, well calcified, with anterolateral regions globular. Ocular acicles multispinose. Chelipeds subequal; right stronger, but not longer. Pereopod 4 subchelate. Male right pereopodal coxa 5 with moderate, stout sexual tube; left pereopodal coxa 5 with moderate, stout sexual tube; both with long mesial and terminal long setae; without unpaired pleopods. Female gonopore on left pereopodal coxa 3 only; with unpaired pleopods 2–4. Uropods symmetrical. Telson posterior margin entire. Maximum sl. 2.2 mm. Slope (455–700 m) inhabiting corbulid bivalve shells). Eastern Indo-Pacific. 1 species (Lemaitre and McLaughlin 1995).

Alloeopagurodes Komai, 1998 Diagnosis. Lateral projections reduced. Thoracic sternite 6 anterior lobe subrectangular, with spinose margin. Antennal scaphocerite with row of spines on mesial surface. Chelipeds unequal; right elongate in large males. Male right pereopodal coxa 5 with short, mesially-directed sexual tube; left pereopodal coxa 5 with short sexual tube. Telson posterior margins rounded. Maximum sl. 3.8 mm. Shelf (66–120 m). Temperate Northern W Pacific. 1 species (Komai 1998).

Anapagrides de Saint Laurent-Dechancé, 1966 Diagnosis. Chelipeds unequal; right much stronger. Male pereopodal coxae 5 asymmetrical; right and left pereopodal coxae 5 dissimilar, or both with short, stout sexual tubes posteriorly-directed; left pereopodal coxa 5 with short sexual tube, or with papilla. Female gonopore on left pereopodal coxa 3 only. Telson anterior half narrower than posterior, posterior margins oblique, or margins straight. Maximum sl. 2 mm. Subtidal–slope (3–307 m). Temperate Northern W Pacific, Central and Eastern Indo-Pacific (Japan, Hawaii). 3 species (Komai 1999).

Anapagurus Henderson, 1886 Diagnosis. Rostrum rounded lobe. Ocular acicles simple, or with submarginal spine. Chelipeds grossly unequal; right massive. Preungual process absent. Male right pereopodal coxa 5 without sexual tube, or with short sexual tube; left pereopodal coxa 5 with short to moderate sexual tube exteriorly-directed, often curved over pleon dorsally. Telson posterior lobes oblique, spinose, or margins transverse, spinose. Maximum sl. 7.5 mm.

327

Intertidal–slope (0–1262 m). Tropical E Atlantic, Temperate Northern E Atlantic, Temperate Northern Pacific (A. japonicus Ortmann, 1892 only), Temperate Southern Africa. 19 species (García-Gómez 1994: key to 18 species)

Anisopagurus McLaughlin, 1981 Diagnosis. Thoracic sternite 6 anterior lobe subrectangular. Ocular acicles simple, or multispinose. Chelipeds unequal; right suboperculate. Propodal rasp with 3–4 rows of corneous scales, preungual process well developed. Female pleopods 1 paired, modified. Telson posterior margins rounded. Maximum sl. 7.1 mm. Subtidal–slope (2–1016 m; usually inhabiting gastropod shell but some live symbiotically with actinians that produce a chitinous carcinoecium). Tropical W Atlantic. 5 species (Lemaitre and McLaughlin 1996: key to species).

Bathypaguropsis McLaughlin, 1994 Diagnosis. Gills: quadriserial, 13 pairs. Chelipeds unequal; right massive, chela operculate or nearly so, propodal-carpal articulation ~30° from perpendicular. Propodal rasp 1 or more, sometimes incomplete, rows of corneous scales. Male unpaired pleopods 2–5 present. Telson posterior margins oblique. Maximum sl. 5.4 mm (Pl. 33g). Shelf, slope (100–333 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 7 species (Komai and Lemaitre 2002: key to 5 species; Komai and Takeda 2004).

Boninpagurus Asakura & Tachikawa, 2004 Diagnosis. Ocular acicles with submarginal spine. Chelipeds unequal; right much larger, opening horizontally; left cheliped slender. Propodal rasp with 3–4 rows of corneous scales. Male right pereopodal coxa 5 with moderate sexual tube, straight or slightly curved, laterally-posterolaterally directed, proximal half stout, opaque, distal portion transparent; left pereopodal coxa 5 with papilla. Telson posterior lobes oblique, spinose. Maximum sl. 2.6 mm. Intertidal–shelf (0–25 m). Temperate Northern W Pacific. 1 species (Asakura and Tachikawa 2004; Komai et al. 2011).

Bythiopagurus McLaughlin, 2003 Diagnosis. Gills: quadriserial, 13 pairs. Shield and carapace lateral lobes well calcified; posterior carapace median element fused with lateral carapace lobes, posteromedian and posterolateral plates well calcified at least in anterior halves. Pleon somewhat reduced, dextrally twisted. Maxilliped 3 crista dentata with 1 accessory tooth, or more. Chelipeds subequal; left longer, not stronger; dactyli and fixed fingers opening in horizontal plane. Pereopod 4 semichelate (weakly). Male right and left pereopodal coxae 5 both with sexual tube or papilla. Female pleopods 1 paired, modified; with unpaired pleopods 2–5. Telson posterior lobes oblique, spinose. Maximum sl. 7 mm. Slope (1083–1300 m). Temperate Australasia. 1 species (McLaughlin 2003b).

328

Marine Decapod Crustacea

Catapaguroides A. Milne-Edwards & Bouvier, 1892

Ceratopagurus Yokoya, 1933

Diagnosis. Gills: 10 pairs. Maxilliped 3 crista dentata without accessory tooth. Chelipeds unequal; right much stronger. Male right pereopodal coxa 5 with short to long sexual tube directed from right to left under thorax, recurved anteriorly; left pereopodal coxa 5 without sexual tube, or with short sexual tube; left concealed between 2 thick tufts of sternal setae. Female gonopore on left pereopodal coxa 3 only. Telson without lateral indentations; posterior margins oblique, or margins straight. Maximum sl. 4.5 mm. Subtidal–bathyal (3–2828 m). Temperate Northern Atlantic, Temperate Northern W Pacific, Western and Central Indo-Pacific (C. microps A. Milne-Edwards & Bouvier, 1892 said to be cosmopolitan). 31 species (Komai 2009: review of species by ocean basin; Komai and Rahayu 2013a; McLaughlin 2002: key to 17 species). Catapaguroides umbra Komai, 2009 with a shield length of 0.83 mm may be the smallest hermit crab known.

Diagnosis. Chelipeds subequal, similar, long, slender. Propodal rasp with several rows of corneous scales. Female pleopods 1 paired, modified. Telson posterior lobes oblique, spinose. Maximum sl. 7.3 mm. Shelf (80–219 m). Central Indo-Pacific. 1 species (McLaughlin 1988).

Catapaguropsis Lemaitre & McLaughlin, 2006 Diagnosis. Pleon always reduced posteriorly in male, reduced or not in female. Cornea dilated; ocular acicles with submarginal spine. Maxilliped 3 crista dentata reduced, without accessory tooth. Chelipeds unequal, elongate; right stouter than left. Ambulatory legs sexually dimorphic, pereopod 2 distinctly shorter than pereopod 3 in male, dactyli slender; pereopods 2, 3 approximately equal in female, dactyli blade-shaped, or not sexually dimorphic; pereopod 4 semichelate (female) or simple (male), propodal rasp present in female, absent in male, preungual process long, setose. Male right pereopodal coxa 5 with short, stout right sexual tube externallydirected; left pereopodal coxa 5 with short sexual tube, or with papilla; without unpaired pleopods. Female with unpaired pleopods 2–4. Uropods asymmetrical, or symmetrical. Telson with weak lateral indentations; posterior lobes separated by broad median concavity, unarmed or with few minute spinules. Maximum sl. 2.7 mm. Slope (296–388 m; inhabiting a carcinoecium likely produced by an actinian). Central Indo-Pacific. 2 species (McLaughlin and Lemaitre 2007).

Catapagurus A. Milne-Edwards, 1880 Diagnosis. Rostrum rounded lobe. Thoracic sternite 6 anterior lobe subrectangular, with spinose margin, or subrectangular. Chelipeds unequal, elongate; right stouter than left. Pereopods 2, 3 of female, dactyli blade-shaped; preungual process prominent. Male right pereopodal coxa 5 with moderate-long sexual tube, curving left towards exterior over dorsal or lateral side of pleon; left pereopodal coxa 5 without sexual tube, or with short sexual tube, or with papilla. Female with unpaired pleopods 2–5, or with unpaired pleopods 2–4. Telson posterior margins oblique. Maximum sl. 4.6 mm. Subtidal–slope (5–809 m). Tropical W Atlantic, Temperate Northern W Pacific, Western and Central Indo-Pacific. 27 species (Asakura 2001: key to 15 Indo-West Pacific species, as junior synonym Hemipagurus; Komai and Rahayu 2021: rediagnosis; McLaughlin 2004b: rediagnosis; Nucci and de Melo 2012: SW Atlantic species).

Cestopagurus Bouvier, 1897 Diagnosis. Gills: distally quadriserial. Chelipeds unequal; right much stronger. Male right pereopodal coxa 5 with long sexual tube orientated towards left across ventral body surface; left pereopodal coxa 5 with short sexual tube. Telson posterior margins oblique, or margins straight. Maximum sl. 5 mm. Intertidal–slope (0–613 m). Temperate Northern E Atlantic, Temperate Northern Pacific, Western Indo-Pacific, Temperate Australasia. 5 species (Komai and Poupin 2012: figures; Komai et al. 2014: key to species).

Chanopagurus Lemaitre, 2003 Diagnosis. Gills: quadriserial, 13 pairs (pleurobranchs above pereopods 2, 3 vestigial). Cornea reduced, distoventrolateral; ocular acicles simple (contiguous). Chelipeds subequal. Propodal rasp with 1 or 2 rows of corneous scales. Male right and left pereopodal coxae 5 both without sexual tube; unpaired pleopods 2–5 present. Female gonopore on left pereopodal coxa 3 only; pleopods 1 paired, modified. Telson symmetrical, lateral margins blade-like, lateral indentations distinct; posterior lobes half-moon shaped. Maximum sl. 7.1 mm. Slope (880–1070 m). Central Indo-Pacific. 1 species (Osawa and Chan 2009).

Cycetopagurus McLaughlin, 2004 Diagnosis. Ocular acicles subrectangular. Chelipeds subequal; right stronger, but not longer. Male pereopodal coxae 5 asymmetrical; right pereopodal coxa 5 with long sexual tube, stout proximally, extremely filiform distally, posteriorly-, externallydirected; left pereopodal coxa 5 with short sexual tube. Telson with weak lateral indentations, anterior part with marginal spiniform bristles; posterior margins rounded (with corneous spinules). Maximum sl. 3 mm. Shelf (134–140 m). Western Indo-Pacific (Western Australia). 1 species (McLaughlin 2004a).

Decaphyllus de Saint Laurent, 1968 Diagnosis. Gills: 8–10 pairs (anterior arthrobranchs sometimes reduced; without pleurobranchs). Maxilliped 3 crista dentata reduced, without accessory tooth. Chelipeds subequal; right stronger, but not longer. Pereopod 4 simple, propodal rasp present in female, absent in male. Male right pereopodal coxa 5 with long sexual tube orientated towards left across ventral body surface; left pereopodal coxa 5 with short sexual tube directed from

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

329

left to right; unpaired pleopods 2–5 present. Female gonopore on left pereopodal coxa 3 only. Telson without lateral indentations; posterior margin entire, or margin weakly cleft. Maximum sl. 2.8 mm. Subtidal, shelf (4–289 m). Temperate Northern W Pacific, Central Indo-Pacific. 11 species (Komai and Rahayu 2013b: key to species).

semichelate, dactylus with circular sensory structure on lateral face, propodal rasp with several rows of corneous scales. Male unpaired pleopods 3–5 present (5 sometimes absent). Female pleopods 1 paired, modified. Telson posterior margins oblique. Maximum sl. 23 mm. Intertidal–slope (0–388 m). Temperate Northern W Pacific. 3 species (McLaughlin 1974: key to species).

Dentalopagurus McLaughlin, 2007

Enallopaguropsis McLaughlin, 1981

Diagnosis. Gills: quadriserial. Pleon straight. Ocular acicles with submarginal spine. Maxilliped 3 crista dentata with 1 or 2 accessory teeth. Chelipeds unequal, elongate; right stouter than left. Propodal rasp with 1 or 2 rows of corneous scales. Male right and left pereopodal coxae 5 both with short, stout sexual tubes posteriorly-directed; partially masked by tuft of setae; unpaired pleopods 2–5 present. Female pleopods 1 paired, modified. Uropods symmetrical. Telson posterior lobes parallel-sided, separated by broad spinose V-shaped cleft. Maximum sl. 3.7 mm. Slope (582–700 m; using scaphopod shells). Central IndoPacific. 2 species (Osawa and Chan 2008).

Diagnosis. Thoracic sternite 6 anterior lobe a single capsulate seta. Pleon straight, or slightly flexed. Chelipeds unequal; right suboperculate. Propodal rasp with several rows of corneous scales, preungual process small or well developed. Male right pereopodal coxa 5 without sexual tube, or with short sexual tube, or with papilla (or on left side). Female pleopods 1 paired, modified. Telson without lateral indentations; posterior margin entire, or margin weakly concave, or margins convex. Maximum sl. 3.8 mm. Shelf–slope (20–275 m). Tropical W Atlantic, Tropical Eastern Pacific. 3 species (Lemaitre and McLaughlin 2003b: key to species).

Enallopagurus McLaughlin, 1981 Diacanthurus McLaughlin & Forest, 1997 Diagnosis. Chelipeds unequal. Propodal rasp with several rows of corneous scales. Male pereopodal coxae 5 symmetrical, both produced; both masked by tufts of long, stiff setae; unpaired pleopods 2–5 present. Telson lateral margin blade-like; posterior margin concave, outer angles acute, prominent pair of spines adjacent to median cleft. Maximum sl. 18 mm. Shelf–bathyal (65–2134 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia. 6 species (McLaughlin and Forest 1997: key to 5 species; de Saint Laurent and McLaughlin 2000).

Discorsopagurus McLaughlin, 1974 Diagnosis. Pleon straight or slightly flexed, not twisted; tergites of pleonites 3–4 paired, with incompletely fused chitinous plates; tergite of pleonite 6 strongly calcified. Antennal flagella articles with paired long setae armed with prominent setules. Chelipeds unequal. Propodal rasp with multiple rows of corneous scales. Male right pereopodal coxa 5 without sexual tube, or with papilla (or on left side); unpaired pleopods 2–5 present, or unpaired pleopods 3–5 present. Uropods symmetrical. Telson without lateral indentations, or with weak lateral indentations; posterior margin entire. Maximum sl. 3.7 mm. Intertidal–slope (0–220 m; inhabiting boreholes in rock or coral made by pholadid clams and sabellid polychaetes, using their antennae as a cast-net for feeding). Temperate Northern Atlantic, Temperate Northern Pacific. 4 species (Komai 2003: key to species).

Elassochirus Benedict, 1892 Diagnosis. Chelipeds unequal; right considerably larger, carpus with wing-like expansions or strong spines. Pereopod 4 weakly

Diagnosis. Chelipeds unequal; right chela subovate. Preungual process small. Male right pereopodal coxa 5 without sexual tube, or with short sexual tube, or with papilla. Female pleopods 1 paired, modified. Telson without lateral indentations; posterior margin entire, or margin weakly cleft, or margins convex. Maximum sl. 3.1 mm. Shelf–slope (57–265 m). Tropical W Atlantic, Tropical Eastern Pacific. 4 species (Lemaitre and McLaughlin 2003b: key to species).

Enneobranchus García-Gómez, 1988 Diagnosis. Gills: distally quadriserial, 9 pairs (2 arthrobranchs each on pereopods 1–4, pleurobranch above pereopod 4). Maxilliped 3 crista dentata without accessory tooth. Chelipeds unequal; right much stronger. Pereopod 4 simple, preungual process prominent. Male right pereopodal coxa 5 without sexual tube, or with papilla; left pereopodal coxa 5 with moderate to long, coiled sexual tube; right gonopore obscured by setae. Telson posterior margins oblique, or margins straight. Maximum sl. 2.7 mm. Subtidal, shelf (15–241 m). Tropical W Atlantic. 3 species (García-Gómez 1988).

Enneopagurus McLaughlin, 1997 Diagnosis. Gills: quadriserial, 9 pairs (2 arthrobranchs each on pereopods 1–4, pleurobranch above pereopod 4). Maxilliped 3 crista dentata without accessory tooth. Chelipeds subequal; right stronger, but not longer. Male right pereopodal coxa 5 without sexual tube, or with papilla; left pereopodal coxa 5 with moderate, stout sexual tube dorsally-exteriorly-directed, terminally spatulate with fringe of curved setae. Telson with weak lateral indentations; posterior margins oblique. Maximum sl. 3.6 mm. Slope (356–552 m). Central Indo-Pacific. 1 species (McLaughlin 1997).

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Marine Decapod Crustacea

Enneophyllus McLaughlin, 1997

Haigiopagurus McLaughlin, 2005

Diagnosis. Gills: 9 pairs (2 arthrobranchs each on pereopods 1–4, pleurobranch above pereopod 4). Pleon straight. Maxilliped 3 crista dentata reduced, without accessory tooth. Chelipeds unequal; right much stronger. Male left pereopodal coxa 5 with long, basally stout sexual tube exteriorly-directed, curved dorsally across pleon from left to right. Telson with weak lateral indentations; posterior margins oblique. Maximum sl. 1.6 mm. Shelf, slope (206–408 m; inhabiting scaphopod shells). Temperate Northern W Pacific, Central Indo-Pacific. 2 species (Komai 2017).

Diagnosis. Pleon straight, or slightly flexed. Chelipeds unequal; right chela subquadrate-subrectangular. Female pleopods 1 paired, modified. Telson posterior margins straight, or margins excavate. Maximum sl. 6.4 mm.

Eutrichopagurus Komai, 2015

Diagnosis. Lateral projections prominent. Antennal flagella articles with paired long setae armed with prominent setules. Chelipeds unequal, elongate; right stouter than left. Preungual process prominent. Male right pereopodal coxa 5 with short sexual tube, posteriorly-directed; left pereopodal coxa 5 with short sexual tube (usual); unpaired pleopods 2–5 present. Telson posterior margins rounded, with long, slender, corneous spines. Maximum sl. 5.2 mm. Subtidal–slope (15–1260 m). Central Indo-Pacific. 2 species (McLaughlin 1997).

Diagnosis. Gills: quadriserial. Cornea reduced, on tapering eyestalk. Maxilliped 3 crista dentata with 2 accessory teeth. Chelipeds unequal; right much stronger. Female gonopore on left pereopodal coxa 3 only. Telson posterior lobes oblique, spinose (with row of long setae on lateral margins). Maximum sl. 1.9 mm. Subtidal (3–5 m). Central Indo-Pacific. 1 species (Komai 2015).

Forestopagurus García-Gómez, 1994 Diagnosis. Rostrum rounded lobe. Chelipeds unequal; right elongate in large males. Male left pereopodal coxa 5 with moderate, stout sexual tube; without unpaired pleopods. Female with unpaired pleopods 2–4. Telson posterior margin entire. Maximum sl. 3.8 mm. Shelf (31 m). Tropical E Atlantic. 1 species (García-Gómez 1994).

Goreopagurus McLaughlin, 1988 Diagnosis. Gills: biserial or distally quadriserial. Chelipeds grossly unequal; right elongate with prominently produced lower carpal margin. Preungual process absent or well developed. Male right and left pereopodal coxae 5 both with sexual tube or papilla, or both without sexual tube. Female pleopods 1 paired, modified. Telson posterior margins oblique, or margins straight. Maximum sl. 5 mm (Pl. 33i). Shelf, slope (44–1300 m). Tropical W Atlantic, Tropical Eastern Pacific, Temperate Australasia. 4 species (Lemaitre and McLaughlin 2003a: rediagnosis; Nucci and de Melo 2007: key to species).

Hachijopagurus Osawa & Okuno, 2003 Diagnosis. Ocular acicles multispinose. Chelipeds grossly unequal; right massive. Male pereopodal coxae 5 slightly asymmetrical, right larger; right pereopodal coxa 5 with sexual tube rod-like, distally curved, twice coxal length, anteriorly-directed; left pereopodal coxa 5 with rod-like sexual tube ventromesially-directed, distally curved shorter, more slender than right. Female gonopore on left pereopodal coxa 3 only. Telson posterior lobes oblique, spinose. Maximum sl. 1.6 mm. Shelf (40–45 m). Temperate Northern Pacific. 1 species (Osawa and Okuno 2003).

Subtidal (3–23 m; inhabiting gastropods and serpulid polychaete tubes). Tropical Eastern Pacific. 1 species (McLaughlin 1982; McLaughlin and Lemaitre 2001, as Haigia). Icelopagurus McLaughlin, 1997

Iridopagurus de Saint Laurent-Dechancé, 1966 Diagnosis. Gills: quadriserial. Chelipeds subequal. Pereopod 4 simple, preungual process absent or small. Male right pereopodal coxa 5 with coiled sexual tube; left pereopodal coxa 5 with short sexual tube, or with papilla. Telson posterior margins straight. Maximum sl. 5.2 mm. Subtidal–slope (1–713 m). Temperate Northern W Atlantic, Tropical W Atlantic, Tropical Eastern Pacific. 9 species (GarcíaGómez 1983: key to species).

Kumepagurus Komai & Osawa, 2012 Diagnosis. Gills: 13 pairs. Maxilliped 3 crista dentata without accessory tooth. Chelipeds unequal; right massive, chela operculate or nearly so, propodal-carpal articulation ~30° from perpendicular. Male right pereopodal coxa 5 with medium, coiled, sexual tube, laterally-oriented; left pereopodal coxa 5 with short sexual tube, tapering distally to slender acute tip, posteriorly-directed. Telson posterior margins oblique. Maximum sl. 2.4 mm. Shelf (38–172 m). Temperate Northern W Pacific. 2 species (Komai and Osawa 2012; Komai et  al. 2021: K. kaikata Komai, 2020 transferred to Vounopagurus Komai, Rahayu & Fujita, 2021).

Labidochirus Benedict, 1892 Diagnosis. Carapace except branchiostegites, heavily calcified; posterior carapace broader than shield. Ocular acicles obscured by anterior margin of shield. Chelipeds subequal, or chelipeds unequal. Pereopod 4 simple, propodal rasp with 1 or 2 rows of corneous scales. Male without unpaired pleopods. Telson posterior margins straight. Maximum sl. 17 mm. Subtidal, shelf, slope (1–419 m). Arctic, Temperate Northern Pacific. 2 species (McLaughlin 1974: key to species).

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

Leptopagurus Lemaitre, Felder & Poupin, 2017 Diagnosis. Chelipeds subequal; both slender, with clusters of elongate, capsulate setae on lower face of merus. Male right and left pereopodal coxae 5 both with stout, membranous sexual tube; partially masked by tuft of setae. Telson posterior lobes oblique, spinose. Maximum sl. 1.5 mm. Slope (50–224 m). Tropical W Atlantic. 1 species (Lemaitre et al. 2017).

Lithopagurus Provenzano, 1968 Diagnosis. Gills: 13 pairs. Pleon reduced, straight. Chelipeds unequal; right operculate. Propodal rasp with several rows of corneous scales. Male pleopods 2 paired, pleopods 3–5 absent. Female with unpaired pleopods 2–4. Telson without lateral indentations; posterior margin entire. Maximum sl. 2.8 mm. Shelf, slope (146–540 m). Tropical W Atlantic, Central IndoPacific. 3 species (McLaughlin and Lemaitre 2004).

Lophopagurus McLaughlin, 1981 Diagnosis. Pleon typically spirally twisted, or straight, or slightly flexed. Chelipeds unequal; right chela subrectangular, upper surface of palm ringed by row of marginal spines or with sloping or concave dorsomesial component; left chela with upper-lateral margin elevated, or midline elevated into prominent keel. Preungual process absent or small. Male unpaired pleopods 2–5 present, or unpaired pleopods 3–5 present. Female pleopods 1 paired, modified. Uropods asymmetrical, or symmetrical. Telson posterior margins oblique, or margins rounded, or margins straight. Maximum sl. 12 mm. Subtidal–slope (2–1280 m). Temperate Northern W Pacific, Central Indo-Pacific, Temperate Australasia. 16 species. Two subgenera are commonly recognised, Lophopagurus s.s. and Autraleremus McLaughlin, 1981, differentiated in the key by Lemaitre and McLaughlin (2003b). McLaughlin and Gunn (1992) presented keys to the two subgenera, six species of Lophopagurus and seven species of Australeremus.

Manucomplanus McLaughlin, 1981 Diagnosis. Rostrum triangular, or rounded lobe. Chelipeds unequal (right sexually dimorphic). Propodal rasp with several rows of corneous scales, preungual process well developed. Female pleopods 1 paired, modified. Telson posterior margins oblique, or margins rounded. Maximum sl. 5.7 mm. Shelf–slope (24–225 m). Tropical Atlantic, Tropical Eastern Pacific, Temperate Southern Africa. 5 species (Lemaitre and McLaughlin 1996: key to species).

Michelopagurus McLaughlin, 1997 Diagnosis. Gills: quadriserial. Chelipeds subequal (right stouter). Propodal rasp with 1 or 2 rows of corneous scales. Male right pereopodal coxa 5 with short sexual tube, or on left or on both

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coxae; partially masked by tuft of setae. Female pleopods 1 paired, modified. Uropods asymmetrical, or symmetrical. Telson posterior lobes posterolaterally expanded, separated by narrow spinose median cleft. Maximum sl. 4 mm. Slope, bathyal (209–2165 m; some species inhabiting scaphopod shells). Temperate Northern Atlantic, Tropical Atlantic, Central, Eastern and possibly Western Indo-Pacific. 5 species (Lemaitre 2015: key to species)

Micropagurus McLaughlin, 1986 Diagnosis. Ocular acicles multispinose. Chelipeds unequal. Propodal rasp with 3–4 rows of corneous scales. Male right pereopodal coxa 5 without gonopore and sexual tube, or gonopore present; left pereopodal coxa 5 with moderate to long sexual tube. Telson without lateral indentations; posterior margin entire. Maximum sl. 2.9 mm. Intertidal–slope (0–262 m). Indo-West Pacific, Temperate Australasia. 6 species Asakura (2005: key to species; Lemaitre 2010).

Munidopagurus A. Milne-Edwards & Bouvier, 1893 Diagnosis. Gills: 13 pairs. Chelipeds unequal, elongate; right stouter than left. Pereopod 4 elongate, simple, galatheid-like, propodal rasp replaced by row of setae. Male without unpaired pleopods. Female pleopods 1 paired, modified; with unpaired pleopods 2–4. Uropods symmetrical; peduncle with prominent, posteriorly directed spine. Telson without lateral indentations; posterior margin entire. Maximum sl. 14.6 mm. Slope (201–420 m; carrying sea anemone alone). Tropical W Atlantic. 1 species (Provenzano 1971).

Nematopaguroides Forest & de Saint Laurent, 1968 Diagnosis. Rostrum obtusely and roundly triangular. Chelipeds subequal, or chelipeds unequal. Preungual process absent or well developed. Male right pereopodal coxa 5 with moderate to long sexual tube with terminal filament, obliquely-directed; left pereopodal coxa 5 without sexual tube, or with short sexual tube. Telson posterior margins oblique. Maximum sl. 1.5 mm. Subtidal, shelf (1–91 m). Tropical W Atlantic. 3 species (Lemaitre et al. 2017: species reviewed).

Nematopagurus A. Milne-Edwards & Bouvier, 1892 Diagnosis. Chelipeds subequal; right longer or more robust. Male right pereopodal coxa 5 with moderate to long sexual tube, distally filamentous, oriented left to right; left pereopodal coxa 5 with short sexual tube. Female pleopods 1 paired, modified. ­Telson posterior margins oblique, or margins rounded, or margins straight. Maximum sl. 9 mm (Pl. 33j). Subtidal–bathyal (9–2148 m). Temperate Northern Atlantic, Temperate Northern W Pacific, Indo-West Pacific, Temperate Southern Africa, Temperate Australasia. 29 species (McLaughlin 2004d: species groups, key to species)

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Orthopagurus Stevens, 1927 Diagnosis. Pleon straight or slightly flexed, tergites chitinous as lateral plates, tergite 5 entire, tergite 6 strongly calcified. Chelipeds unequal; right suboperculate. Propodal rasp with several rows of corneous scales. Telson posterior margins straight. Maximum sl. 5.6 mm. Subtidal, shelf (11–64 m). Temperate Northern E Pacific. 1 species (McLaughlin 1974).

Ostraconotus A. Milne-Edwards, 1880 Diagnosis. Gills: 10 pairs. Carapace nearly completely calcified, lateral margins evenly serrated; rostrum broadly rounded. Pleon reduced, straight. Maxilliped 3 crista dentata reduced, without accessory tooth. Chelipeds unequal. Pereopod 4 with broad discoid propodus, wider in female; pereopod 4 propodus expanded, flat, dactylus long, simple, propodal rasp present in female, absent in male. Male right pereopodal coxa 5 with moderate to long sexual tube; left pereopodal coxa 5 without sexual tube, or with papilla; without unpaired pleopods. Female with unpaired pleopods 2–4. Uropods symmetrical. Telson posterior margin entire. Maximum sl. 5 mm. Shelf–slope (137–256 m). Tropical Atlantic. 1 species. While oft-mentioned as one of the crab-like, non-shell carrying pagurids (e.g. Poupin and McLaughlin 1996), the only species O. spatulipes A. Milne-Edwards & Bouvier, 1893 seems not to have been illustrated since shortly after its first description (Milne-Edwards and Bouvier 1893).

Pagurellus Lemaitre, Felder & Poupin, 2017 Diagnosis. Gills: 9 pairs (2 arthrobranchs each on pereopods 1–4, pleurobranch above pereopod 4). Chelipeds unequal; right much stronger. Male right pereopodal coxa 5 with basally weakly calcified, distally filamentous sexual tube; left pereopodal coxa 5 with sexual tube similar to right, longer, coiled. Uropods symmetrical. Telson posterior lobes oblique, spinose. Maximum sl. 1.3 mm. Slope (224–247 m). Tropical W Atlantic1 species (Lemaitre et al. 2017).

Paguridium Forest, 1961 Diagnosis. Rostrum rounded lobe. Chelipeds unequal. Male pereopodal coxae 5 asymmetrical, left produced; left pereopodal coxa 5 with short sexual tube, or with papilla; left gonopore masked by tuft of long, stiff setae directed towards right; without unpaired pleopods. Telson posterior margins straight. Maximum sl. 3 mm. Intertidal–shelf (0–51 m). Temperate Northern W Atlantic. 1 species (Forest 1961).

Pagurina Lemaitre, Felder & Poupin, 2017 Diagnosis. Gills: 9 pairs (2 arthrobranchs each on pereopods 1–4, pleurobranch above pereopod 4). Maxilliped 3 crista dentata reduced, without accessory tooth. Chelipeds subequal. Male left

pereopodal coxa 5 with short sexual tube. Uropods symmetrical. Telson posterior lobes oblique, spinose. Maximum sl. 0.8 mm. Subtidal (10 m). Tropical W Atlantic. 1 species (Lemaitre et al. 2017).

Paguriscus Lemaitre, Felder & Poupin, 2017 Diagnosis. Gills: 8 pairs (2 arthrobranchs each on maxilliped 3, pereopods 1, 2, 1 arthrobranch each on pereopods 3, 4, without pleurobranchs). Chelipeds unequal. Male pereopodal coxae 5 slightly asymmetrical, trapezoidal, extending posteriorly to form paired sexual tubes. Uropods almost symmetrical. Telson posterior margins rounded. Maximum sl. 1.3 mm. Subtidal (2–19 m). Tropical W Atlantic. 1 species (Lemaitre et al. 2017).

Paguritta Melin, 1939 Diagnosis. Pleon straight. Ocular acicles simple, or bifid. Antennal flagella articles with paired long setae armed with prominent setules. Chelipeds unequal. Male right and left pereopodal coxae 5 both with sexual tube or papilla; without unpaired pleopods. Female with unpaired pleopods 2–4. Uropods symmetrical. Telson posterior margins straight. Maximum sl. 3 mm (Pl. 33h). Subtidal (1–15 m; inhabiting serpulid or polychaete tubes and worm holes in coral, or vermetid gastropods). Temperate Northern W Pacific, Western Indo-Pacific, Central Indo-Pacific. 7 species (Komai 1996: key to species). McLaughlin and Lemaitre (1993) rediagnosed the genus, described new species and discussed how the cast-net structure of the antennal flagella is used as a filter-feeding mechanism by these largely immobile hermit crabs.

Pagurixus Melin, 1939 Diagnosis. Chelipeds unequal; right elongate in large males. Male pereopodal coxae 5 asymmetrical, right produced; right gonopore obscured by tuft of stiff setae directed towards left. Female gonopore on left pereopodal coxa 3 only, or on both coxae 3. Telson posterior margins oblique, or margins rounded, or margins straight. Maximum sl. 4.9 mm. Intertidal–shelf (0–91 m; inhabiting serpulid polychaete tubes as well as gastropods). Temperate Northern W Pacific, Indo-West Pacific, Temperate Australasia. 42 species (Komai 2010: key to 36 species).

Pagurodes Henderson, 1888 Diagnosis. Gills: quadriserial. Chelipeds subequal, elongate; right more robust. Ambulatory legs dactyli elongate, almost straight, distally slightly thickened and twisted, lower margins with row of corneous spinules. Male right pereopodal coxa 5 with short sexual tube, posteriorly-directed; left pereopodal coxa 5 with short sexual tube, or with papilla. Telson posterior margins oblique, or margins nearly perpendicular. Maximum sl. 18 mm (Pl. 33k). Slope, bathyal (1165–3250 m). Western Indo-Pacific, Temperate Australasia. GOODS bathyal provinces: New Zealand/Kermadec, Indian Ocean. 1 species (de Saint Laurent and McLaughlin 2000).

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

Pagurodofleinia Asakura, 2005 Diagnosis. Chelipeds subequal; right stronger, but not longer. Left pereopod 2 left with row of closely-spaced comb-like teeth on lower margin, right with lower margin unarmed; propodal rasp with 3–4 rows of corneous scales. Male pereopodal coxae 5 asymmetrical; right pereopodal coxa 5 with short sexual tube; left pereopodal coxa 5 without sexual tube, or with papilla. Telson with broadly U-shaped median cleft; posterior angles bearing 1–3 (right) and 2–5 (left) strong corneous spines. Maximum sl. 9.5 mm. Shelf, slope (90–400 m). Temperate Northern W Pacific. 1 species (Asakura 2001: redescribed the only species as Parapagurodes doederleini).

Pagurojacquesia de Saint Laurent & McLaughlin, 2000 Diagnosis. Gills: quadriserial. Maxilliped 3 crista dentata without accessory tooth. Chelipeds subequal. Pereopod 4 semichelate or subchelate. Pereopod 5 subchelate. Male left pereopodal coxa 5 with club-like, stout, short to moderate sexual tube directed towards exterior, with terminal tuft of long setae. Female pleopods 1 paired, modified. Telson posterior margins oblique. Maximum sl. 6 mm. Shelf, slope (207–660 m). Central Indo-Pacific, Temperate Australasia. 1 species (de Saint Laurent and McLaughlin 1999: P. polymorpha described as member of Jacquesia).

Paguruncio Lemaitre, Felder & Poupin, 2017 Diagnosis. Gills: 8 pairs (1 arthrobranch on pereopod 1, 2 arthrobranchs each on pereopods 2–4, pleurobranch above pereopod 4). Chelipeds subequal. Male right and left pereopodal coxae 5 both with slender, membranous sexual tube posteriorly-directed. ­Telson posterior lobes oblique, spinose. Maximum sl. 0.6 mm. Shelf–slope (91–224 m). Tropical W Atlantic. 1 species (Lemaitre et al. 2017).

Pagurus Fabricius, 1775 Diagnosis. Pleon typically spirally twisted, or straight. Ocular acicles simple, or multispinose, or bifid. Maxilliped 3 crista dentata with 1 accessory tooth, or more. Chelipeds subequal. Propodal rasp with 1 row of corneous scales or with several rows of corneous scales, preungual process well developed. Male right and left pereopodal coxae 5 both without sexual tube, or rarely with papilla; without unpaired pleopods. Female gonopore on left pereopodal coxa 3 only (rarely), or on both coxae 3; with unpaired pleopods 2–5, or with unpaired pleopods 2–4 (rarely). Uropods asymmetrical, or symmetrical (rarely). Telson posterior margins oblique, or margins rounded, or margins straight, or margin entire. Maximum sl. 18.6 mm (Pl. 34a). Intertidal–slope (0–1800 m). Cosmopolitan. 181 species. Pagurus is the ‘catch-aIl’ genus for any hermit crab having 11 pairs of biserial phyllobranchiate gills, but lacking secondary sexual modifications or other special characters. As a consequence it the

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most heterogeneous of all pagurid genera. Thirty per cent of > 600 pagurid species belong to Pagurus. No single work has reviewed the genus, which was diagnosed by McLaughlin (1974) with a key to 25 species from the north-east Pacific, and by (McLaughlin 1997). Mantelatto et al. (2009) did not find Pagurus polyphyletic but their study was based on few species. The genus was been divided into eight groups by McLaughlin (1974) on morphological criteria. Some groups have received more recent specific attention: for example, the provenzanoi-group (Lemaitre et  al. 1982), the anachoretus-group (Komai and Rahayu 2004), the bernhardusgroup (Lemaitre and Watabe 2005) and capillatus-group (Komai et al. 2006). At least three of these groups have molecular support (Olguín and Mantelatto 2013).

Parapagurodes McLaughlin & Haig, 1973 Diagnosis. Rostrum acute. Chelipeds unequal; right elongate in large males. Propodal rasp with 3–4 rows of corneous scales. Male right pereopodal coxa 5 with short sexual tube; left pereopodal coxa 5 without sexual tube, or with short sexual tube. Telson posterior lobes oblique, spinose, or margins transverse, spinose. Maximum sl. 9.2 mm. Subtidal–slope (10–475 m). Temperate Northern Pacific, Tropical Eastern Pacific. 2 species (McLaughlin and Asakura 2004).

Patagurus Anker & Paulay, 2013 Diagnosis. Anterior carapace vaulted, well calcified, gastric region slightly elevated; lateral margins each 2 small, subacute lobes; posterior carapace lobes prominent, laterally projecting, calcified except for narrow, posterior, membranous band tucked under calcified portion; branchiostegite with 2 large and 2 small calcified plates; rostrum prominent, subtriangular. Pleon reduced, narrow, elongate, segmentation of anterior pleonites not discernible. Ocular acicles vestigial. Chelipeds unequal; right much stronger. Male right pereopodal coxa 5 with sexual tube calcified, with long tuft of distal setae, mesially-directed; left pereopodal coxa 5 similar but shorter than right; without unpaired pleopods. Uropods symmetrical. Telson posterior margin entire. Maximum sl. 5.1 mm (Pl. 33l). Slope (400 m; using bivalve shells). Eastern Indo-Pacific. 1 species (Anker and Pauly 2013).

Phimochirus McLaughlin, 1981 Diagnosis. Maxilliped 3 crista dentata with 1 accessory tooth, or more. Chelipeds markedly unequal; right chela subovate to subcircular. Preungual process prominent. Female pleopods 1 paired, modified. Telson posterior margins oblique. Maximum sl. 7 mm (Pl. 34b). Intertidal–slope (0–640 m). Tropical W Atlantic, Tropical Eastern Pacific. 8 species (Lemaitre and McLaughlin 2003b: key to species).

Pliopagurus Komai, 2013 Diagnosis. Ocular acicles triangular. Chelipeds unequal; right markedly longer, chela suboperculiform, with dorsomesial margin

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Marine Decapod Crustacea

delimited by row of small spines; left with chela uncompressed dorsoventrally, propodal-carpal articulation 45°. Male right and left pereopodal coxae 5 both with very short, strongly tapering sexual tube. Female pleopods 1 paired, modified. Telson with weak lateral indentations; posterior margins oblique. Maximum sl. 2.5 mm. Shelf (80–90 m). Central Indo-Pacific. 1 species (Komai 2013b).

Shelf–slope (88–300 m). Central Indo-Pacific. 4 species (McLaughlin and Rahayu 2007; Rahayu and Komai 2013).

Pteropagurus McLaughlin & Rahayu, 2006

Diagnosis. Anterior carapace vaulted, well calcified; lateral margins of shield with 2 blunt or spiniform, wing-like projections; posterior carapace lobes prominent; rostrum prominent, broad, triangular or truncate. Pleon reduced, usually globular, with tergites at least faintly delineated. Chelipeds unequal; right much stronger. Male pereopodal coxae 5 symmetrical, sometimes expanded posteriorly; without unpaired pleopods. Female with unpaired pleopods 2–4. Uropods symmetrical. Telson posterior margin entire, or margin weakly cleft. Maximum sl. 15 mm. Subtidal–slope (3–1430 m; using bivalve or limpet shells). Temperate Northern W Pacific, Central Indo-Pacific, Tropical Eastern Pacific, Temperate Southern Africa, Temperate Australasia. 14 species (McLaughlin 2000: key to 11 Indo-West Pacific species).

Diagnosis. Pleon reduced, straight; pleonite 1 partially fused with last thoracic somites, tergites 2–5 weakly delineated or not; tergite 6 chitinous. Ocular acicles with submarginal spine. Maxilliped 3 crista dentata reduced, without accessory tooth. Chelipeds unequal, elongate; right stouter than left. Pereopod 4 semichelate or minutely chelate, propodal rasp rudimentary. Male right pereopodal coxa 5 with long, stout sexual tube posteriorly directed, curving over anterior portion of pleon; left pereopodal coxa 5 without sexual tube, or with short sexual tube. Female gonopore on left pereopodal coxa 3 only, or on both coxae 3. Uropods symmetrical; peduncle with or without posteriorly-directed spine, exopod elongate, endopod reduced, both with 1 or 2 marginal rows of scales. Telson without lateral indentations; posterior margins oblique, or margins rounded. Maximum sl. 1.6 mm. Slope (285–700 m; inhabiting shells of pelagic pteropod molluscs). Central Indo-Pacific. 3 species (McLaughlin 2007).

Propagurus McLaughlin & de Saint Laurent, 1998

Pumilopagurus McLaughlin & Rahayu, 2008

Diagnosis. Gills: quadriserial, 13 pairs (pleurobranchs above pereopods 2, 3 vestigial). Thoracic sternite 6 anterior lobe subsemicircular, or subrectangular. Chelipeds unequal. Propodal rasp with several rows of corneous scales. Male unpaired pleopods 2–5 present, or unpaired pleopods 3–5 present. Telson posterior margins oblique. Maximum sl. 16 mm (Pl. 34c, d). Shelf, slope (38–913 m). Temperate Northern W Pacific, Central Indo-Pacific, Temperate South America, Temperate Southern Africa, Temperate Australasia. 5 species (Komai and Konishi 2003; McLaughlin and Lemaitre 2000).

Diagnosis. Cornea reduced, on tapering eyestalk; ocular acicles bifid, or with submarginal spine. Chelipeds grossly unequal; right massive. Male right pereopodal coxa 5 with short sexual tube; left pereopodal coxa 5 with short sexual tube, directed from left to right across ventral surface. Female gonopore on left pereopodal coxa 3 only. Telson posterior margins transverse, spinose. Maximum sl. 1.4 mm. Subtidal, shelf (7–36 m). Central Indo-Pacific. 1 species (McLaughlin and Rahayu 2008).

Protoniopagurus Lemaitre & McLaughlin, 1996

Diagnosis. Gills: 8 pairs (1 arthrobranch on pereopod 1, 2 arthrobranchs each on pereopods 2–4, pleurobranch above pereopod 4). Maxilliped 3 crista dentata without accessory tooth. Chelipeds subequal, similar, long, slender. Male right pereopodal coxa 5 with coiled, tapering sexual tube; left pereopodal coxa 5 with tapering sexual tube, shorter than on right. Telson posterior lobes oblique, spinose. Maximum sl. 0.9 mm. Subtidal, shelf (15–116 m). Tropical W Atlantic. 1 species (Lemaitre et al. 2017).

Porcellanopagurus Filhol, 1885

Diagnosis. Pleon reduced, straight. Ocular acicles simple, or bifid. Chelipeds subequal; both suboperculate. Propodal rasp with 10–12 rows of corneous scales. Male without unpaired pleopods. Female pleopods 1 paired, modified; with unpaired pleopods 2–4. Uropods symmetrical. Telson posterior margin entire. Maximum sl. 7.8 mm. Shelf, slope (91–585 m). Tropical W Atlantic. 1 species (Lemaitre and McLaughlin 1996).

Pseudopagurodes McLaughlin, 1997 Diagnosis. Gills: distally quadriserial. Chelipeds subequal. Ambulatory legs dactyli evenly curved, evenly tapering to corneous tip, lower margins usually without corneous spines. Male right pereopodal coxa 5 with moderate to long sexual tube, distally filamentous, oriented left to right; without unpaired pleopods. Telson posterior margins oblique. Maximum sl. 4.8 mm.

Pusillopagurus Lemaitre, Felder & Poupin, 2017

Pygmaeopagurus McLaughlin, 1986 Diagnosis. Chelipeds grossly unequal; right massive. Pereopod 4 semichelate (weakly) or simple. Male right pereopodal coxa 5 without gonopore and sexual tube; left pereopodal coxa 5 with shortmoderate, rod-like sexual tube. Female gonopore on left pereopodal coxa 3 only. Telson posterior margins oblique. Maximum sl. 0.9 mm. Subtidal (5–6 m). Central Indo-Pacific. 1 species (McLaughlin 1986).

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

335

Pylopaguridium McLaughlin & Lemaitre, 2001

Solenopagurus de Saint Laurent, 1968

Diagnosis. Ocular acicles multispinose. Chelipeds unequal; right markedly larger than left, subrectangular, operculate. Male pereopodal coxae 5 asymmetrical. Female pleopods 1 paired, modified. Telson posterior margins straight. Maximum sl. 2.3 mm. Subtidal, shelf (3–36 mm). Tropical Atlantic. 1 species (McLaughlin and Lemaitre 2001).

Diagnosis. Gills: distally quadriserial. Chelipeds subequal; right stronger and longer. Left pereopod 3 propodus and dactylus dissimilar from other pereopods, with plumose setae on lateral faces; preungual process absent or well developed. Male right pereopodal coxa 5 with long, basally stout sexual tube exteriorly-directed, curved dorsally across pleon from left to right; left pereopodal coxa 5 with papilla. Telson posterior margins oblique, or margins straight. Maximum sl. 4 mm. Shelf (45–146 m). Tropical Atlantic, Tropical Eastern Pacific. 2 species (Wass 1963: description as species of Cestopagurus).

Pylopaguropsis Alcock, 1905 Diagnosis. Gills: 13 pairs. Chelipeds unequal; right usually massive, operculate or semioperculate, dactylus frequently articulating obliquely with palm. Pereopods 3 propodi and dactyl frequently dissimilar; propodal rasp with 1–3 rows of corneous scales, preungual process absent or well developed. Female pleopods 1 paired, modified. Telson posterior margins oblique, or margins straight, or margins excavate. Maximum sl. 4.6 mm (Pl. 34e). Intertidal–slope (0–610 m). Temperate Northern Pacific, Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific. 20 species (Komai and Osawa 2004: key to 14 species). Lemaitre (2017) discussed three groups of species, tabulated them all with geographic distributions, depth ranges and habitats. He speculated that P. mollymullerae Lemaitre, 2017 engages in cleaner activities or is a den commensal with moray eels.

Pylopagurus A. Milne-Edwards & Bouvier, 1893 Diagnosis. Pleon straight, or slightly flexed. Chelipeds unequal; right markedly larger than left, chela subcircular to subrectangular, operculate. Male right and left pereopodal coxae 5 each without sexual tube or rarely with papilla. Female pleopods 1 paired, modified. Uropods symmetrical. Telson posterior margins oblique, or margins excavate. Maximum sl. 7 mm. Subtidal–slope (11–1020 m; inhabiting scaphopods or semitransparent onuphid polychaete tubes). Tropical W Atlantic, Tropical Eastern Pacific. 5 species (McLaughlin and Lemaitre 2001: redescriptions, key to species).

Rhodochirus McLaughlin, 1981 Diagnosis. Chelipeds unequal; right chela subovate to subquadrate, with spines or tubercles with basal rosettes. Preungual process well developed. Female pleopods 1 paired, modified. Telson posterior margins oblique. Maximum sl. 7 mm. Shelf (95–200 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (McLaughlin 1981b).

Solitariopagurus Türkay, 1986 Diagnosis. Gills: 10 pairs. Anterior carapace vaulted, strongly calcified; lateral margins of shield with 3 blunt or spiniform lobes; posterior carapace lobe not laterally expanded; rostrum prominent, broadly triangular. Pleon reduced, pleonal plate 2 weakly delineated, plates 3–5 clearly defined. Ocular acicles simple (hidden from dorsal view by anterior margin of shield). Chelipeds unequal; right much stronger. Pereopod 4 subchelate. Male right pereopodal coxa 5 with short sexual tube, or with moderate, stout sexual tube; left pereopodal coxa 5 with short sexual tube, or with moderate, stout sexual tube; both with long-terminal and subterminal setae; without unpaired pleopods. Female gonopore on left pereopodal coxa 3 only. Uropods symmetrical; peduncle with prominent, posteriorly directed spine. Telson posterior margin entire. Maximum sl. 5.7 mm (Pl. 34f). Shelf–slope (20–1995 m; using bivalve shells). Indo-West Pacific. 5 species(McLaughlin 2000: key to 4 Indo-West Pacific species; Türkay 2015).

Spathapagurus Lemaitre & Felder, 2011 Diagnosis. Ocular acicles with submarginal spine. Chelipeds grossly unequal; right twice or more length of left, right chela flattened, spatulate, carpus lower margin moderately produced. Male right and left pereopodal coxae 5 both with papilla; both masked by tufts of long, stiff setae. Telson posterior lobes separated by deep median cleft, posterior half of lateral margins and terminal margins armed with spines. Maximum sl. 4.5 mm. Subtidal, shelf (15–69 m). Tropical Atlantic, Tropical Eastern Pacific. 2 species (Lemaitre and Felder 2011).

Scopaeopagurus McLaughlin & Hogarth, 1998

Spiropagurus Stimpson, 1858

Diagnosis. Gills: 10 pairs. Maxilliped 3 crista dentata comprising 2–4 curved, spines, without accessory tooth. Chelipeds grossly unequal; right massive. Male right pereopodal coxa 5 with papilla; left pereopodal coxa 5 with short sexual tube; unpaired pleopods 2–5 present. Female gonopore on left pereopodal coxa 3 only. ­Telson posterior margins oblique. Maximum sl. 1.1 mm. Shelf (46 m). Western Indo-Pacific. 1 species (McLaughlin and Hogarth 1998).

Diagnosis. Chelipeds subequal; right stronger, but not longer. Male right pereopodal coxa 5 without sexual tube, or with papilla; left pereopodal coxa 5 with long, usually coiled, terminally blunt sexual tube. Telson posterior margins acutely triangular. Maximum sl. 9 mm (Pl. 34g). Subtidal, bathyal (5–1500 m). Temperate Northern Atlantic, Western and Central Indo-Pacific. 7 species (Han et al. 2016b: key to species).

336

Marine Decapod Crustacea

Tarrasopagurus McLaughlin, 1997

Turleania McLaughlin, 1997

Diagnosis. Gills: distally quadriserial. Chelipeds unequal. Male right pereopodal coxa 5 with short sexual tube, or with papilla, or without gonopore and sexual tube; left pereopodal coxa 5 with short sexual tube. Female pleopods 1 paired, modified. Telson posterior margins oblique. Maximum sl. 2.7 mm. Slope (205–425 m). Central Indo-Pacific. 1 species (McLaughlin 1997).

Diagnosis. Gills: quadriserial, 9 pairs (2 arthrobranchs each on pereopods 1–4, pleurobranch above pereopod 4) or 11 pairs. Ocular acicles simple, or multispinose. Maxilliped 3 crista dentata without accessory tooth. Chelipeds subequal. Male right pereopodal coxa 5 with papilla; left pereopodal coxa 5 with moderate to long, weakly spiralled sexual tube, with sparse tuft of setae. Telson with weak lateral indentations; posterior margins oblique, or lobes oblique, spinose. Maximum sl. 2.3 mm. Shelf, slope (85–675 m). Central Indo-Pacific. 9 species (Komai 2020b).

Tomopaguroides Balss, 1912 Diagnosis. Gills: quadriserial, 13 pairs. Pleon straight, tergite 5 thickened, possibly calcified plate. Chelipeds grossly unequal; right massive. Propodal rasp with 1 or 2 rows of corneous scales. Male pleopods 2 paired, unpaired pleopods 3–5 present. Uropods symmetrical. Maximum sl. 6.3 mm. Slope (550–1079 m; inhabiting scaphopod shells). Western and Central Indo-Pacific. 1 species (McLaughlin 2004c).

Tomopaguropsis Alcock, 1905 Diagnosis. Gills: quadriserial, 13 pairs. Thoracic sternite 6 anterior lobe with convex median, marginally setose, elevation. Chelipeds subequal. Propodal rasp with several rows of corneous scales. Male unpaired pleopods 2–5 present, or pleopods 1 paired, modified; unpaired pleopods 2–5 present. Telson posterior margins rounded. Maximum sl. 5.4 mm. Subtidal–slope (13–827 m). Tropical W Atlantic, Western and Central Indo-Pacific. 6 species (Jung et  al. 2017: key to species; Lemaitre et al. 2014).

Tomopagurus A. Milne-Edwards & Bouvier, 1893 Diagnosis. Chelipeds unequal; right much stronger. Preungual process prominent. Male unpaired pleopods 2–5 present (vestigial paired pleopods 1 rare). Female pleopods 1 paired, modified, or absent. Telson posterior margins oblique. Maximum sl. 13 mm. Shelf, slope (35–366 m). Tropical W Atlantic, Tropical Eastern Pacific, Temperate South America. 8 species (McLaughlin 1981a: key to species).

Trichopagurus de Saint Laurent, 1968 Diagnosis. Rostrum triangular. Chelipeds unequal. Male pereopodal coxae 5 asymmetrical, left produced; right pereopodal coxa 5 with moderate sexual tube exteriorly-directed, distal part semitransparent, coiled; left pereopodal coxa 5 with short sexual tube. Female gonopore on left pereopodal coxa 3 only. Telson with weak or distinct lateral indentations; posterior lobes oblique, spinose (with row of long setae on lateral margins). Maximum sl. 2.5 mm. Intertidal–shelf (0–100 m). Temperate Northern W Pacific, Indo-West Pacific. 4 species (Komai 2013a: key to species).

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13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

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CHJM, De Grave S, Ng PKL (eds) Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume. Crustaceana Monographs 14, 379–388. doi:10.1163/9789047427759_026 Lemaitre R (2015) A new species of the hermit crab genus Michelopagurus McLaughlin, 1997 (Crustacea: Decapoda: Paguridae) from Moorea, French Polynesia. Zoosystema 37, 363–370. doi:10.5252/z2015n2a5 Lemaitre R (2017) Discovery of a new species of hermit crab of the genus Pylopaguropsis Alcock, 1905 from the Caribbean: “den commensal” or “cleaner”? (Crustacea, Anomura, Paguridae). ZooKeys 646, 139–158. doi:10.3897/zookeys.646.11132 Lemaitre R, Felder DL (2011) A new genus of Paguridae (Crustacea: Decapoda: Anomura) for a new species from the tropical eastern Pacific and Pagurus longimanus Wass, 1963 from the tropical western Atlantic. Zootaxa 3125, 39–50. Lemaitre R, McLaughlin PA (1995) Alainopagurus crosnieri, gen. et sp. nov. (Decapoda, Anomura, Paguridae) from the Western Pacific. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 17, 273–282. Lemaitre R, McLaughlin PA (1996) Revision of Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with the descriptions of new genera and species. Part V. Anisopagurus McLaughlin, Manucomplanus McLaughlin, and Protoniopagurus new genus. Bulletin of Marine Science 59, 89–141. Lemaitre R, McLaughlin PA (2003a) New species of Goreopagurus (Decapoda: Anomura: Paguridae) from Tasmania and reevaluation of sexual tubes in hermit crab systematics. Memoirs of Museum Victoria 60, 221–227. doi:10.24199/j.mmv.2003.60.22 Lemaitre R, McLaughlin PA (2003b) Revision of Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with descriptions of new genera and species. Addendum and taxonomic summary. Proceedings of the Biological Society of Washington 116, 464–486. Lemaitre R, Watabe H (2005) Pagurus ikedai (Crustacea: Anomura: Paguridae), a new hermit crab species of the bernhardus group from Japanese waters. Zootaxa 819, 1–12. doi:10.11646/zootaxa.819.1.1 Lemaitre R, McLaughlin PA, García-Gómez J (1982) The provenzanoi group of hermit crabs (Crustacea, Decapoda, Paguridae) in the Western Atlantic. Part IV. A review of the group, with notes on variations and abnormalities. Bulletin of Marine Science 32, 670–701. Lemaitre R, Vázquez-Bader AR, Gracia A (2014) An unusual new species of paguroid (Crustacea, Anomura, Paguridae) from deep waters of the Gulf of Mexico. ZooKeys 449, 57–67. doi:10.3897/zookeys.449.8541 Lemaitre R, Felder DL, Poupin J (2017) Discovery of a new micro-pagurid fauna (Crustacea: Decapoda: Paguridae) in the Lesser Antilles, Caribbean Sea. Zoosystema 39, 151–195. doi:10.5252/z2017n2a1 Mantelatto FL, Pardo LM, Pileggi LG, Felder DL (2009) Taxonomic reexamination of the hermit crab species Pagurus forceps and Pagurus comptus (Decapoda: Paguridae) by molecular analysis. Zootaxa 2133, 20–32. doi:10.11646/zootaxa.2133.1.2 McLaughlin PA (1974) The hermit crabs (Crustacea Decapoda, Paguridea) of northwestern North America. Zoölogische Verhandelingen 130, 1–396. McLaughlin PA (1981a) Revision of Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with the descriptions of new genera and species. Part I. Ten new genera of the Paguridae and a redescription of Tomopagurus A. Milne Edwards and Bouvier. Bulletin of Marine Science 31, 1–30. McLaughlin PA (1981b) Revision of Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with the descriptions of new genera and species. Part II. Rhodochurus McLaughlin and Phimochurus McLaughlin. Bulletin of Marine Science 31, 329–365. McLaughlin PA (1982) Revision of Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with the descriptions of new genera and species. Part III, Agarochirus McLaughlin, Enallopagurus McLaughlin, and Enalopaguropsis McLaughlin. Bulletin of Marine Science 32, 823–855.

McLaughlin PA (1986) Three new genera and species of hermit crabs (Crustacea: Anomura: Paguridae) from Hawaii. Journal of Crustacean Biology 6, 789–803. doi:10.2307/1548392 McLaughlin PA (1988) The rediscovery of Ceratopagurus Yokoya and a new genus for Pagurus piercei Wass (Decapoda, Paguroidea, Paguridae). Crustaceana 55, 257–267. doi:10.1163/156854088X00357 McLaughlin PA (1997) Crustacea Decapoda: hermit crabs of the family Paguridae from the KARUBAR cruise in Indonesia. In: Crosnier, A and Bouchet, P. (eds), Résultats des Campagnes MUSORSTOM, vol. 16. Mémoires du Muséum National d’Histoire Naturelle, Paris 172, 433–572. McLaughlin PA (2000) Crustacea Decapoda: Porcellanopagurus Filhol and Solitariopagurus Türkay (Paguridae), from New Caledonian area, Vanuatu and the Marquesas: new records, new species. In: Crosnier, A. (ed), Résultats des Campagnes MUSORSTOM, vol. 21. Mémoires du Muséum National d’Histoire Naturelle, Paris 184, 389–414. McLaughlin PA (2002) A new species of the genus Catapaguroides (Decapoda, Anomura, Paguroidea, Paguridae) from Guam, Micronesia. Crustaceana 75, 495–504. doi:10.1163/156854002760095543 McLaughlin PA (2003a) Illustrated keys to families and genera of the superfamily Paguroidea (Crustacea: Decapoda: Anomura), with diagnoses of genera of Paguridae. Memoirs of Museum Victoria 60, 111–144. doi:10.24199/j.mmv.2003.60.16 McLaughlin PA (2003b) A new genus and species of hermit crab (Decapoda: Anomura: Paguridae) from seamounts off south-eastern Tasmania, Australia. Memoirs of Museum Victoria 60, 229–236. doi:10.24199/j.mmv.2003.60.23 McLaughlin PA (2004a) A new genus and species of hermit crabs (Crustacea: Anomura: Paguroidea) from northwestern Australia. Records of the Western Australian Museum 22, 101–108. doi:10.18195/issn.03123162.22(2).​2004.101-108 McLaughlin PA (2004b) A reappraisal of the hermit crab genera Catapagurus A. Milne-Edwards and Hemipagurus Smith (Crustacea: Decapoda: Anomura: Paguridae), with the description of a new species. Zootaxa 433, 1–16. doi:10.11646/zootaxa.433.1.1 McLaughlin PA (2004c) Redescription of Tomopaguroides valdiviae (Balss, 1911) (Crustacea, Decapoda, Anomura, Paguroidea, Paguridae) with notes on variation and female morphology. Zoosystema 26, 469–481. McLaughlin PA (2004d) A review of the hermit crab genus Nematopagurus A. Milne-Edwards and Bouvier, 1892 (Crustacea: Decapoda: Paguridae). In: Marshall, B.A., and Richer de Forges, B. (eds), Tropical Deep-Sea Benthos, Vol. 23. Mémoires du Muséum National d’Histoire Naturelle, Paris 191, 151–229. McLaughlin PA (2007) A new species of Pteropagurus McLaughlin & Rahayu, 2006 and a new genus and species of scaphopod dweller (Decapoda, Anomura, Paguroidea, Paguridae). Zoosystema 29, 503–513. McLaughlin PA, Asakura A (2004) Reevaluation of the hermit crab genus Parapagurodes McLaughlin & Haig, 1973 (Decapoda: Anomura: Paguroidea: Paguridae) and a new genus for Parapagurodes doederleini (Dolflein, 1902). Proceedings of the Biological Society of Washington 117, 42–56. McLaughlin PA, de Saint Laurent M (1998) A new genus for four species of hermit crabs formerly assigned to the genus Pagurus Fabricius (Decapoda: Anomura: Paguridae). Proceedings of the Biological Society of Washington 111, 158–187. McLaughlin PA, Forest J (1997) Crustacea Decapoda: Diacanthurus gen. nov., a new genus of hermit crabs (Paguridae) with both Recent and fossil representation, and the descriptions of two new species. In: Crosnier, A. (ed), Résultats des Campagnes MUSORSTOM, vol. 18. Mémoires du Muséum National d’Histoire Naturelle, Paris 176, 235–259. McLaughlin PA, Gunn SW (1992) Revision of Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with descriptions of new genera and species. Part IV. Lophopagurus McLaughlin and Australeremus

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

McLaughlin. Memoirs of the Museum of Victoria 53, 43–99. doi:10.24199/​ j.mmv.1992.53.03 McLaughlin PA, Hogarth PJ (1998) Hermit crabs (Decapoda: Anomura: Paguridae) from the Seychelles. Zoölogische Verhandelingen 318, 3–48. McLaughlin PA, Lemaitre R (1993) A review of the hermit crab genus Paguritta (Decapoda: Anomura:Paguridae) with descriptions of three new species. Raffles Bulletin of Zoology 41, 1–29. McLaughlin PA, Lemaitre R (2000) Reassignment of Pagurus miyakei Baba, 1986 to Propagurus McLaughlin & de Saint Laurent, 1998 (Decapoda: Anomura: Paguridae) and description of the male. Crustacean Research 29, 58–64. doi:10.18353/crustacea.29.0_58 McLaughlin PA, Lemaitre R (2001) Revision of Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with descriptions of new genera and species. Part VI. Pylopagurus A. Milne-Edwards & Bouvier, 1891, Haigia McLaughlin, 1981, and Pylopaguridium, a new genus. Proceedings of the Biological Society of Washington 114, 444–483. McLaughlin PA, Lemaitre R (2004) The discovery of two new species of Lithopagurus Provenzano, 1968 (Crustacea, Decapoda, Anomura, Paguroidea, Paguridae) and the first records of the genus in the western Pacific. Zoosystema 26, 483–494. McLaughlin PA, Lemaitre R (2007) A new and distinctive species of the hermit crab genus Catapaguropsis (Crustacea: Decapoda: Anomura: Paguroidea: Paguridae) from the South China Sea. Zootaxa 1560, 31–41. doi:10.11646/zootaxa.1560.1.3 McLaughlin PA, Rahayu DL (2007) Pseudopagurodes McLaughlin, 1997 (Crustacea: Anomura: Paguroidea: Paguridae) revisited. Raffles Bulletin of Zoology Supplement 16, 21–27. McLaughlin PA, Rahayu DL (2008) A new genus and species of hermit crab of the family Paguridae (Crustacea: Anomura: Paguroidea) from the Vanuatu Archipelago. Proceedings of the Biological Society of Washington 121, 365–373. doi:10.2988/08-07.1 McLaughlin PA, Komai T, Lemaitre R, Rahayu DL (2010) Annotated checklist of anomuran decapod crustaceans of the world (exclusive of the Kiwaoidea and families Chirostylidae and Galatheidae of the Galatheoidea) Part I – Lithodoidea, Lomisoidea and Paguroidea. Raffles Bulletin of Zoology Supplement 23, 5–107. Milne-Edwards A, Bouvier E-L (1893) Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877–78), in the Caribbean Sea (1878–79), and along the Atlantic coast of the United States (1880), by the U. S. Coast Survey steamer “Blake,” Lieut-Com. S. D. Sigsbee, U. S. N., and Commander J. R. Bartlett, U. S. N., commanding. XXXIII. Description des Crustacés de la famille des paguriens recueillis pendant l’expédition. Memoirs of the Museum of Comparative Zoology at Harvard College 14, 5–172. Nucci PR, de Melo GAS (2007) First record of the genus Goreopagurus McLaughlin, 1998 (Crustacea: Anomura: Paguridae) from Brazil, with description of a new species. Zootaxa 1520, 51–56. doi:10.11646/ zootaxa.1520.1.4 Nucci PR, de Melo GAS (2012) A new species of Catapagurus A. MilneEdwards (Crustacea: Decapoda: Anomura: Paguridae) from Brazil, with a key to the western Atlantic species. Zootaxa 3499, 81–85. doi:10.11646/zootaxa.3499.1.6 Olguín N, Mantelatto FL (2013) Molecular analysis validates of some informal morphological groups of Pagurus (Fabricius, 1775) (Anomura: Paguridae) from South America. Zootaxa 3666, 436–448. doi:10.11646/zootaxa.3666.4.2 Osawa M, Chan T-Y (2008) A new species of Dentalopagurus (Crustacea: Decapoda: Anomura) from deep waters off Taiwan. Bulletin of Marine Science 82, 263–273. Osawa M, Chan T-Y (2009) Additional records of hermit crabs (Crustacea: Decapoda: Anomura: Paguroidea) from Taiwan. Proceedings of the Biological Society of Washington 122, 317–332. doi:10.2988/08-50.1

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Osawa M, Okuno J (2003) A new genus and new species of the family Paguridae (Crustacea: Decapoda: Anomura), from Hachijo-jima Island, Japan, with a list of hermit crab species found in the same collection sites. Proceedings of the Biological Society of Washington 116, 943–955. Poupin J, McLaughlin PA (1996) A new species of Solitariopagurus Türkay (Decapoda, Anomura, Paguridae) from French Polynesia. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 18, 211–224. Provenzano AJ (1971) Biological Results of the University of Miami Deep-Sea Expeditions. 74. Rediscovery of Munidopagurus macrocheles (A. Milne-Edwards, 1880) (Crustacea, Decapoda, Paguridae), with a description of the first zoeal stage. Bulletin of Marine Science 21, 256–266. Rahayu DL, Komai T (2013) Two new species of Pseudopagurodes McLaughlin, 1997 (Crustacea, Decapoda, Anomura, Paguridae) from the Philippines. In: Ahyong, S.T., Chan, T.-Y., Corbari, L. & Ng, P.K.L. (eds), Tropical Deep-Sea Benthos Vol. 27. Mémoires du Muséum National d’Histoire Naturelle 204, 423–435. Tudge C, Lemaitre R (2004) Studies of male sexual tubes in hermit crabs (Crustacea, Decapoda, Anomura, Paguroidea). I. Morphology of the sexual tube in Micropagurus acantholepis (Stimpson, 1858), with comments on function and evolution. Journal of Morphology 259, 106–118. doi:10.1002/jmor.10174 Türkay M (2015) A new species of Solitariopagurus from the Red Sea with notes on S. profundus (Crustacea: Decapoda: Paguridae). Zootaxa 3920, 579–585. doi:10.11646/zootaxa.3920.4.7 Wass ML (1963) New species of hermit crabs (Decapoda, Paguridae) from the Western Atlantic. Crustaceana 6, 133–157. doi:10.1163/156854063X00525

Parapaguridae Smith, 1882 deep-water hermit crabs Figure 13.28, Plate 34h–l Parapagurids are unusual hermit crabs from deep water, recorded from shelf to hadal depths (> 5000 m) where they may be extraordinarily common in trawl samples. They are especially conspicuous on continental slopes. Whereas typical shallow-water hermits sometimes carry a solitary seaanemone on their shell home, parapagurids take this association one step further. Most species carry gastropod shells for protection, but many others live symbiotically with just anthozoans (sea-anemones or zoantharians) instead. The sea anemone settles on a gastropod shell inhabited by the young hermit crab, then covers and extends the shell to produce a chitinous structure termed a carcinoecium (e.g. Crowther et al. 2011). In some other parapagurid genera the home is made only of a cavity in a flat gelatinous colony of a dozen or more zoantharians (Epizoanthus). The protective corona of zoantharians may, as the Reverend Stebbing said, ‘contribute to the commissariat by throwing out its darts as some swift gliding shrimp passes by, and thus reducing it to a condition in which it may be captured by the pagurid.’ The systematics of the Parapaguridae was reviewed by de Saint Laurent (1972) and updated by Lemaitre (1998, 1999, 2004b, 2004c, 2013, 2014) in a series of papers where he reviewed most genera in turn. In these works he provided

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keys and synthesised bathymetry and distributions. Most recently Fraaije et  al. (2022) resurrected the long disused family name, Probeebeidae, for one pagurid, two parapagurids (Labidochirus, Probeebei and Tylaspis) and a diogenid (Tisea); here they are retained in Paguridae, Parapaguridae and Diogenidae, respectively. Pereopod 1 is the cheliped, pereopods 2 and 3 the two pairs of walking legs, and pereopods 4 and 5 reduced. The right cheliped in males tends to become more elongate with increasing size, often markedly so, and the carpus and chela wider than long in females than in males.

Dimensions are of the length of the shield (sl.), the anterior harder part of the carapace in front of the cervical groove. Diagnosis. Right cheliped larger than left. Maxilliped 1 exopod without flagellum (Fig. 13.28l). Maxillipeds 3 widely separate at base (Fig. 13.28m). Implicit generic attributes. Rostrum shorter than ocular peduncle. Shield as wide as long. Posterior carapace mostly membranous. Gills phyllobranchiate; pleurobranch absent above pereopod 5. Cornea present; ocular acicle distinctly developed. Pleopods 3–5 unpaired, on left.

Key to genera of Parapaguridae 1. – 2. – 3. – 4. – 5.

Corneas absent (Fig. 13.28g)�����������������������������������������������������������������������������������������������������������������������������������Typhlopagurus Corneas present��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Carapace prominently spinose overall (Fig. 13.28d). Pleonites separate, well calcified�����������������������������������������Probeebei Carapace shield smooth. Pleonites fused, largely membranous�����������������������������������������������������������������������������������������������3 Ocular acicle minute, unarmed (Fig. 13.28h)������������������������������������������������������������������������������������������������������������������Tylaspis Ocular acicle distinctly developed (Fig. 13.28a–g)����������������������������������������������������������������������������������������������������������������������4 Posterior carapace calcified (Fig. 13.28a). Pleopods 3–5 asymmetrically paired������������������������������������������Bivalvopagurus Posterior carapace mostly membranous. Pleopods 3–5 unpaired, on left������������������������������������������������������������������������������5 Shield longer than wide (Fig. 13.28f). Rostrum acute. Pleon straight����������������������������������������������������������Tsunogaipagurus

Fig. 13.28.  Parapaguridae. Shield, eyestalks, ocular acicles: a, Bivalvopagurus; b, Paragiopagurus; c, Parapagurus; d, Probeebei; e, Strobopagurus; f, Tsunogaipagurus; g, Typhlopagurus. Eyestalks, ocular acicles, dorsolateral view, h, Tylaspis. Epistome and spine, dorsolateral view, i, Oncopagurus. Antennal peduncle: j, Paragiopagurus; k, Parapagurus. Maxilliped 1: l, Oncopagurus. Maxillipeds 3: m, Parapagurus. Pereopod 3 dactylus: n, Paragiopagurus; o, Sympagurus; p, Strobopagurus. Male pleopod 2: q, Parapagurus; r, Strobopagurus. Gill filament: s, Parapagurus. Pereopod 5 with vestigial pleurobranch, t, Sympagurus.

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

– 6. – 7. – 8. – 9. –

341

Shield as wide or wider than long (Fig. 13.28b–c, e). Rostrum blunt or rounded. Pleon twisted����������������������������������������6 Shield wider than long (Fig. 13.28e). Ambulatory legs dactyli straight or nearly so (Fig. 13.28p). Cornea strongly dilated (Fig. 13.28e). Male pleopod 2 with rudimentary exopod, distal segment strongly twisted (Fig. 13.28r)���������������������������������������������������������������������������������������������������������������������������������������������������������� Strobopagurus Shield about as wide as long (Fig. 13.28b). Ambulatory legs dactyli at least slightly curved (Fig. 13.28n, o). Cornea moderately or weakly dilated. Male pleopod without exopod, distal segment not twisted (Fig. 13.28q) (rarely absent)�������������������������������������������������������������������������������������������������������������������������������������������������������7 Pleurobranch above pereopod 5 vestigial (Fig. 13.28t)�������������������������������������������������������������������������������������������Sympagurus Pleurobranch above pereopod 5 absent����������������������������������������������������������������������������������������������������������������������������������������8 Epistomial spine strongly curved upward (Fig. 13.28i)���������������������������������������������������������������������������������������� Oncopagurus Epistomial spine straight or absent�����������������������������������������������������������������������������������������������������������������������������������������������9 Antennal peduncle article 4 with dorsodistal spine (Fig. 13.28j). Ocular peduncle including cornea at least half length of shield. Gills phyllobranchiate, bi- or quadriserial������������������������������������������������� Paragiopagurus Antennal peduncle article 4 unarmed (Fig. 13.28k). Ocular peduncle including cornea usually less than half length of shield (Fig. 13.28c). Gills trichobranchiate, quadriserial (Fig. 13.28s)���������������������������������� Parapagurus Bivalvopagurus Lemaitre, 1993

Diagnosis. Shield wider than long. Posterior carapace calcified. Ocular peduncle including cornea half length of shield, stout. Antennal peduncle article 4 with dorsodistal spine. Ambulatory legs dactyli straight or nearly so. Pleopods 3–5 asymmetrically paired. Maximum sl. 12 mm. Shelf, slope (220–510 m; carrying a bivalve shell with seaanemone for protection. Central Indo-Pacific (South China Sea). 1 species (Lemaitre 1993).

Oncopagurus Lemaitre, 1996 Diagnosis. Epistomial spine strongly curved upward. Ocular peduncle including cornea at least half length of shield; cornea weakly dilated, or reduced, subconical (rarely). Antennal peduncle article 4 with dorsodistal spine. Ambulatory legs dactyli curved. Male pleopod 2 without exopod, distal segment not twisted. Maximum sl. 6 mm, often c. 3 mm (Pl. 34h). Shelf–bathyal (50–2308 m; usually using gastropod shell as adults). Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific, Temperate South America, Temperate Southern Africa, Temperate Australasia. 24 species (Lemaitre 2014: bathymetry, distribution, key to species).

Paragiopagurus Lemaitre, 1996 Diagnosis. Gills phyllobranchiate. Ocular peduncle including cornea at least half length of shield; cornea weakly dilated, or reduced, subconical (rarely). Antennal peduncle article 4 with dorsodistal spine. Ambulatory legs dactyli curved. Male pleopod 2 without exopod, distal segment not twisted. Maximum sl. 12.5 mm. Shelf–bathyal (36–2034 m; protected by gastropod shells with associated sea-anemone, sometimes using scaphopods). Tropical Atlantic, Indo-West Pacific, Temperate Australasia. 25 species (Landschoff and Lemaitre 2017; Lemaitre 2013: key to 24 species). Paragiopagurus ventilatus Lemaitre, 2004 is the only paguroid

associated with hydrothermal vents, at 128–281 m near Taiwan (Lemaitre 2004a).

Parapagurus Smith, 1879 Diagnosis. Gills trichobranchiate, quadriserial. Ocular peduncle including cornea less than half length of shield; cornea weakly dilated. Ambulatory legs dactyli curved. Male pleopod 2 without exopod, distal segment not twisted. Maximum sl. 21 mm (Pl. 34i). Shelf–abyssal (82–5020 m; often living in zoantharians, Epizoanthus sp.). Cosmopolitan except polar seas (not recorded from Eastern Indo-Pacific). 17 species (Lemaitre 1989: reviwed 4 Atlantic species; 1999: bathymetry, distribution, key to 14 Indo-West Pacific species).

Probeebei Boone, 1926 Diagnosis. Rostrum exceeding ocular peduncle, spinose. Carapace anterior and posterior carapace calcified, spinose. Pleon flexed, twisted (female), or straight (male), tergites separate, calcified. Ocular peduncle including cornea less than half length of shield. Ambulatory legs dactyli straight or nearly so. Maximum sl. 43 mm. Slope–abyssal (1145–4775 m; free-living, non-shell carrying). Tropical Eastern Pacific. GOODS bathyal province: Cocos Plate. 1 species (Wolff 1961).

Strobopagurus Lemaitre, 1989 Diagnosis. Shield wider than long. Ocular peduncle including cornea almost as long as shield, stout; cornea strongly dilated. Ambulatory legs dactyli straight or nearly so. Male pleopod 2 with exopod rudimentary or absent, distal segment strongly twisted. Maximum sl. 9 mm. Shelf, slope (40–1200 m). Tropical Atlantic, Indo-West Pacific. 3 species (Lemaitre 2004b: bathymetry, distribution, key to species).

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Marine Decapod Crustacea

Sympagurus Smith, 1883 Diagnosis. Shield as wide as long, or wider than long. Gills trichobranchiate, quadriserial; pleurobranch above pereopod 5 vestigial. Ocular peduncle including cornea at least half length of shield; cornea weakly dilated. Ambulatory legs dactyli curved. Male pleopod 2 without exopod, distal segment not twisted. Maximum sl. 26 mm (Pl. 34j–l). Shelf–bathyal (40–2537 m; often living in zoantharians, Epizoanthus sp.). Tropical Atlantic, Temperate Northern Pacific, IndoWest Pacific, Temperate Australasia. GOODS bathyal province: Nazca Plate. 17 species (Lemaitre 2004c: bathymetry, disctibution, key to species).

Tylaspis Henderson, 1885 Diagnosis. Rostrum acute, unarmed. Ocular peduncle including cornea less than half length of shield; acicle minute, unarmed. Antennal peduncle article 4 with distolateral and distomesial spines. Ambulatory legs dactyli curved. Pleopods 3–5 asymmetrically paired. Maximum sl. 9 mm. Bathyal, abyssal (2075–4344 m; may carry sea anemones). GOODS abyssal provinces: E Pacific basins, S Pacific Ocean, Central Pacific Ocean. 1 species (Lemaitre 1998).

Tsunogaipagurus Osawa, 1995 Diagnosis. Rostrum acute. Shield longer than wide. Pleon straight. Ocular peduncle including cornea at least half length of shield. Ambulatory legs dactyli straight or nearly so. Maximum sl. 5 mm. Slope (912–977 m). Temperate Northern Pacific, Western Indo-Pacific; inhabiting scaphopod shells. 1 species (Osawa 1995, 1996).

Typhlopagurus de Saint Laurent, 1972 Diagnosis. Shield longer than wide (calcified, swollen). Ocular peduncle including cornea less than half length of shield; cornea absent; acicle inner margin spinose. Ambulatory legs dactyli curved. Maximum sl. 6 mm. Slope (1630 m). Central Indo-Pacific (Philippines). 1 species (de Saint Laurent 1972). References Crowther AL, Fautin DG, Wallace CC (2011) Stylobates birtlesi sp. n., a new species of carcinoecium-forming sea anemone (Cnidaria, Actiniaria, Actiniidae) from eastern Australia. ZooKeys 89, 33–48. doi:10.3897/​ zookeys.89.825 de Saint Laurent M (1972) Sur la famille des Parapaguridae Smith, 1882. Description de Typhlopagurus foresti gen. nov., sp. nov., et de quinze espèces ou sous-espèces nouvelles de Parapagurus Smith (Crustacea, Decapoda). Bijdragen tot de Dierkunde 42, 97–123. doi:10.1163/​ 26660644-04202001 Fraaije RHB, Van Bakel BWM, Jagt JWM, Charbonnier S, Schweigert G, Garcia G, Valentin X (2022) The evolution of hermit crabs (Crustacea, Decapoda, Anomura, Paguroidea) on the basis of carapace morphology: a state-of-the-art-report. Geodiversitas 44, 1–16.

Landschoff J, Lemaitre R (2017) Differentiation of three common deepwater hermit crabs (Crustacea, Decapoda, Anomura, Parapaguridae) from the South African demersal abundance surveys, including the description of a new species of Paragiopagurus Lemaitre, 1996. ZooKeys 676, 21–45. doi:10.3897/zookeys.676.12987 Lemaitre R (1989) Revision of the genus Parapagurus (Anomura: Paguroidea: Parapaguridae), including redescriptions of the Western Atlantic species. Zoölogische Verhandelingen 253, 1–106. Lemaitre R (1993) A new genus of Parapaguridae (Decapoda: Anomura). Crustacean Research 22, 11–20. doi:10.18353/crustacea.22.0_11 Lemaitre R (1998) Revisiting Tylaspis anomala Henderson, 1885 (Parapaguridae), with comments on its relationships and evolution. Zoosystema 20, 289–305. Lemaitre R (1999) Crustacea Decapoda: a review of the species of the genus Parapagurus Smith, 1879 (Parapaguridae) from the Pacific and Indian Oceans. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 20. Mémoires du Muséum National d’Histoire Naturelle, Paris 180, 303–378. Lemaitre R (2004a) Discovery of the first hermit crab (Crustacea: Decapoda: Parapaguridae) associated with hydrothermal vents. Cahiers de Biologie Marine 45, 325–334. Lemaitre R (2004b) A review of Strobopagurus Lemaitre, 1989 (Crustacea: Decapoda: Paguroidea: Parapaguridae), with description of a new species. Scientia Marina 68, 355–372. doi:10.3989/scimar.2004.​ 68n3355 Lemaitre R (2004c) A worldwide review of hermit crab species of the genus Sympagurus Smith, 1883 (Crustacea: Decapoda: Parapaguridae). In: Marshall, BA and Richer de Forges, B (eds), Tropical Deep-Sea Benthos, Vol. 23. Mémoires du Muséum National d’Histoire Naturelle, Paris 191, 85–149. Lemaitre R (2013) The genus Paragiopagurus Lemaitre, 1996 (Crustacea, Decapoda, Anomura, Paguroidea, Parapaguridae): a worldwide review and summary, with descriptions of five new species. In: Ahyong, ST, Chan, T-Y, Corbari, L & Ng, PKL (eds), Tropical Deep-Sea Benthos Vol. 27. Mémoires du Muséum National d’Histoire Naturelle, Paris 204, 311–421. Lemaitre R (2014) A worldwide taxonomic and distributional synthesis of the genus Oncopagurus Lemaitre, 1996 (Crustacea: Decapoda: Anomura: Parapaguridae), with descriptions of nine new species. Raffles Bulletin of Zoology 62, 210–301. Osawa M (1995) A new parapagurid genus, Tsunogaipagurus, for Sympagurus chuni (Balss, 1911) (Crustacea: Decapoda: Anomura). Proceedings of the Japanese Society of Systematic Zoology 53, 62–70. Osawa M (1996) Additional note on Tsunogaipagurus chuni (Balss, 1911) (Decapoda, Anomura, Parapaguridae). Crustaceana 69, 878–881. doi:10.1163/156854096X00268 Wolff T (1961) Description of a remarkable deep-sea hermit crab, with notes on the evolution of the Paguridea. Galathea Report 4, 11–32.

Pylochelidae Bate, 1888 symmetrical hermit crabs Figure 13.29, Plate 35a–c Pylochelids differ from all other hermit crabs in having a calcified, segmented pleon, usually symmetrical. Removed from their portable shelter, pylochelids superficially resemble mud shrimp (Axiidea). Instead of gastropod shells, the typical hermit crab home, pylochelids inhabit cavities in small pieces of wood, stone, pumice or sponge, or use tusk

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

shells or bamboo as homes. Members of this family are mostly distributed between 200 and 500 m depth, rarely reaching as shallow as 20 m or exceeding 1500 m. In deep water, the body in one genus of deep-water pagurids, Porcellanopagurus, is also straight and symmetrical, but bivalve shells are used for protection. The straight, calcified pleon of polychelids is quite a departure from that of the stereotypical hermit crab, and indeed, molecular phylogenetic analyses suggest they may be more closely related to clades of squat lobsters than other hermit crabs (e.g. Bracken-Grissom et al. 2013; Tsang et  al. 2008). Further, a study using five nuclear proteincoding genes by Tsang et  al. (2011) suggested that Pylochelidae may even be polyphyletic. Here, we treat the family as a unit. Forest’s (1987a, 1987b) excellent revisions with keys and detailed discussion of systematics and biology set the scene for study of Pylochelidae. The phylogeny and taxonomy of

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the family have been reviewed again in detail more recently (Lemaitre et al. 2009; McLaughlin and Lemaitre 2009) and as a consequence, the taxonomy of Pylochelidae is well understood. Our taxonomy recognises three subfamilies but, for simplicity, not the tribes or subgenera proposed by earlier workers. All genera were diagnosed more completely by Forest (1987b) and McLaughlin and Lemaitre (2009) who provided keys to species. Species continue to be added to the fauna (Komai 2013; Komai and Chan 2016). Dimensions are of the length of the shield (sl.), the anterior harder part of the carapacein front of the cervical groove. Diagnosis. Body hermit crab-like, pleon symmetrical (Fig.  13.29a) (asymmetrical only in Mixtopagurus), pleonites separate, calcified and distinct. Chelipeds subequal. Maxillipeds 3 adjacent at base. With paired pleopods on pleonites 2–5.

Key to subfamilies and genera of Pylochelidae 1. – 2. –

Shield incompletely separated from posterior carapace (Fig. 13.29b–e). Telson divided into anterior and posterior articulating plates (Fig. 13.29j–l)�����������������������������������������������������������������������������������������������Pylochelinae … 2 Shield completely separated from posterior carapace (Fig. 13.29f, g). Telson not divided into anterior and posterior articulating plates (Fig. 13.29m–q)�������������������������������������������������������������������������������������������������������������������������5 Shield distinctly wider than long; anterior margin usually with well developed, rounded rostral lobe. Cornea reduced, conical (Fig. 13.29c)����������������������������������������������������������������������������������������������������������������� Cheiroplatea Shield approximately as wide as long; anterior margin with median concavity, with or without median spinule. Cornea hemispherical (Fig. 13.29b, d–g)������������������������������������������������������������������������������������������������������������������3

Fig. 13.29.  Pylochelidae. Habitus: a, Trizocheles spinosus (Henderson, 1888). Carapace, eyestalks: Pylochelinae: b, Bathycheles; c, Cheiroplatea; d, Pylocheles; e, Xylocheles. Trizochelinae: f, Parapylocheles; g, Trizocheles. Maxilliped 1: h, Trizocheles. Left cheliped: i, Pomatocheles. Telson: Pylochelinae: j, Cheiroplatea; k, Pylocheles; l, Xylocheles. Trizochelinae: m, Cancellocheles; n, Forestocheles; o, Mixtopagurus; p, Parapylocheles; q, Trizocheles.

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3. – 4. – 5. – 6. – 7. – 8. – 9. –

Marine Decapod Crustacea

Chelipeds together forming circular or subcircular operculum; carpus dorsomesial face without cluster of distal tubercles. Telson posterior portion divided into 2 lobes (Fig. 13.29k)�����������������������������������������������������Pylocheles Chelipeds not forming operculum; carpus dorsomesial face with cluster of distal tubercles. Telson posterior portion not divided into 2 lobes������������������������������������������������������������������������������������������������������������������������������������������������4 Ocular peduncles considerably more than half shield length. Cornea large, distinctly pigmented (Fig. 13.29e)������������������������������������������������������������������������������������������������������������������������������������������������������������������Xylocheles Ocular peduncles less than half shield length. Cornea reduced, not distinctly pigmented (Fig. 13.29b)��������������������������������������������������������������������������������������������������������������������������������������������������������������� Bathycheles Maxilliped 2 without epipod. Chelipeds alone forming operculum (Fig. 13.29i)��������Pomatochelinae … Pomatocheles Maxilliped 2 with epipod. Chelipeds not forming operculum alone�������������������������������������������������������Trizochelinae … 6 Posterior carapace distinctly longer than anterior carapace. Ocular peduncles spinose (Fig. 13.29f)����������������������������������������������������������������������������������������������������������������������������������������������������������Parapylocheles Posterior carapace equal to or shorter than anterior carapace. Ocular peduncles not spinose (Fig. 13.29g)����������������������7 Rostrum with subrostral spine. Telson posterior margin entire (Fig. 13.29m)���������������������������������������������� Cancellocheles Rostrum without subrostral spine. Telson posterior margin concave or cleft (Fig. 13.29n–q)��������������������������������������������8 Pleon asymmetrical. Telson posterior lobes usually asymmetrical (Fig. 13.29o)��������������������������������������������Mixtopagurus Pleon symmetrical. Telson with posterior lobes usually symmetrical������������������������������������������������������������������������������������9 Maxilliped 1 exopod without flagellum. Maxilliped 3 crista dentata without accessory tooth. Telson not divided into anterior and posterior portions by lateral indentations (Fig. 13.29n)������������������Forestocheles Maxilliped 1 exopod with flagellum (Fig. 13.29h). Maxilliped 3 crista dentata with accessory tooth. Telson divided into anterior and posterior portions by lateral indentations (Fig. 13.29q)���������������������������� Trizocheles

Subfamily Pomatochelinae Stebbing, 1914 Pomatocheles is the only genus. Diagnosis. Carapace linea transversalis continuous; shield completely separated from posterior carapace. Telson not divided into anterior and posterior articulating plates. Maxilliped 2 without epipod. Chelipeds alone forming operculum (Fig. 13.29i). Pomatocheles Miers, 1879 Shelf, slope (23–450 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. Maximum sl. 6 mm. 3 species (Forest 1987b: diagnosis, key to species).

Subfamily Pylochelinae Bate, 1888 Four genera differ from other pylochelids in having the shield incompletely separated from the posterior carapace and a divided telson. Diagnosis. Carapace linea transversalis interrupted, shield incompletely separated from posterior carapace (Fig.  13.29b–e). Telson divided into anterior and posterior articulating plates (Fig. 13.29j–l). Bathycheles Forest, 1987 Diagnosis. Telson posterior portion not divided into 2 lobes. Ocular peduncles less than half shield length. Cornea hemispherical;

reduced, not distinctly pigmented. Chelipeds not forming operculum; carpus dorsomesial face with cluster of distal tubercles. Maximum sl. 13 mm. Slope, bathyal (283–2217 m). Tropical W Atlantic, Western and Central Indo-Pacific. 6 species (Forest 1987b: diagnosis, descriptions of species).

Cheiroplatea Bate, 1888 Diagnosis. Telson posterior portion not divided into 2 lobes. Ocular peduncles less than half shield length. Cornea reduced, conical; reduced, not distinctly pigmented (if present). Chelipeds chelae together forming operculum, carpi with elevated anterodorsal facet overhanging posterior margin. Maximum sl. 10 mm (Pl. 35c). Slope (291–1156 m). Tropical W Atlantic, Western and Central Indo-Pacific. 7 species (Komai and Chan 2016: key to species).

Pylocheles A. Milne-Edwards, 1880 Diagnosis. Telson posterior portion not divided into 2 lobes. Ocular peduncles considerably more than half shield length. Cornea hemispherical; large, distinctly pigmented. Chelipeds not forming operculum; carpus dorsomesial face with cluster of distal tubercles. Maximum sl. 5.6 mm (Pl. 35b). Shelf, slope (100–963 m). Temperate Northern W Pacific, Tropical W Atlantic, Central Indo-Pacific, Temperate Australasia. 2 species (Forest 1987b: diagnosis, descriptions of species).

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

Xylocheles Forest, 1987 Diagnosis. Telson posterior portion divided into 2 lobes. Ocular peduncles less than half shield length. Cornea hemispherical; large, distinctly pigmented. Chelipeds together forming circular or subcircular operculum; carpus dorsomesial face without cluster of distal tubercles. Maximum sl. 11 mm. Shelf, slope (148–888 m). Western and Central Indo-Pacific. 2 species (Forest 1987b: diagnosis, descriptions of species).

Subfamily Trizochelinae Forest, 1987 The chelipeds join with other pereopods to form the operculum in five genera. Diagnosis. Carapace linea transversalis continuous; shield completely separated from posterior carapace (Fig. 13.29f, g). Telson not divided into anterior and posterior articulating plates (Fig. 13.29m–q). Maxilliped 2 with epipod. Chelipeds not forming operculum alone. Implicit generic attributes. Rostrum without subrostral spine. Pleon symmetrical. Ocular peduncles not spinose. Maxilliped 1 exopodal without flagellum. Parapylocheles Alcock, 1901 Diagnosis. Posterior carapace distinctly longer than anterior carapace. Telson not divided into anterior and posterior articulating plates; posterior margin entire, symmetrical. Ocular peduncles spinose. Maxilliped 3 crista dentata with accessory tooth or teeth. Maximum sl. 15 mm. Slope (200–925 m). Western and Central Indo-Pacific. 1 species (Forest 1987b: diagnosis, descriptions of species).

Cancellocheles Forest, 1987 Diagnosis. Posterior carapace equal to or shorter than anterior carapace; rostrum with subrostral spine. Telson not divided into anterior and posterior articulating plates; posterior margin entire, symmetrical. Maxilliped 3 crista dentata with accessory tooth or teeth. Maximum sl. 12 mm. Slope (200–360 m). Temperate Northern W Pacific. 1 species (Forest 1987b: diagnosis, descriptions of species).

Forestocheles McLaughlin & Lemaitre, 2009 Diagnosis. Posterior carapace equal to or shorter than anterior carapace. Telson not divided into anterior and posterior articulating plates; posterior margin concave or cleft. Maxilliped 3 crista dentata with accessory tooth or teeth. Maximum sl. 5 mm.

Slope (398–540 m). Temperate Australasia (New Zealand). 1 species (McLaughlin and Lemaitre 2009). Mixtopagurus A. Milne-Edwards, 1880 Diagnosis. Posterior carapace equal to or shorter than anterior carapace. Pleon asymmetrical; telson not divided into anterior

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and posterior articulating plates; telson posterior margin entire, asymmetrical. Maxilliped 3 crista dentata without accessory tooth. Maximum sl. 11 mm. Shelf, slope (196–567 m). Tropical W Atlantic. 1 species (Forest 1987b: diagnosis, descriptions of species).

Trizocheles Forest, 1987 Diagnosis. Posterior carapace equal to or shorter than anterior carapace. Telson divided into anterior and posterior portions by lateral indentations; posterior margin concave or cleft. Maxilliped 1 exopod with flagellum. Maxilliped 3 crista dentata with or without accessory tooth. Maximum sl. 11 mm (Pl. 35a).

Shelf, slope (100–1090 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 23 species (Komai 2013: Japanese species); Komai and Chan (2016: key to species). References Bracken-Grissom HD, Cannon ME, Cabezas P, Feldmann RM, Schweitzer CE, et  al. (2013) A comprehensive and integrative reconstruction of evolutionary history for Anomura (Crustacea: Decapoda). BMC Evolutionary Biology 13, 128. doi:10.1186/1471-2148-13-128 Forest J (1987a) Ethology and distribution of Pylochelidae (Crustacea Decapoda Coenobitoidea). Bulletin of Marine Science 41, 309–321. Forest J (1987b) Les Pylochelidae ou “Pagures symétriques” (Crustacea Coenobitoidea). In: Crosnier, A., Résultats des Campagnes MUSORSTOM, vol. 3. Mémoires du Muséum National d’Histoire Naturelle, Paris (sér. A, Zoologie) 137, 1–254, 9 pls. Komai T (2013) Records of four species of the pylochelid hermit crab genus Trizocheles Forest, 1987 (Crustacea: Decapoda: Anomura) from the Sagami Sea and Izu Islands, central Japan, with descriptions of three new species. Natural History Research 12, 91–112. Komai T, Chan T-Y (2016) “Symmetrical” hermit crabs of the family Pylochelidae (Crustacea: Decapoda: Anomura) collected by the “BIOPAPUA” and “PAPUA NIUGINI” expeditions in the Papua New Guinea, with descriptions of two new species. Zootaxa 4088, 301–328. doi:10.11646/zootaxa.4088.3.1 Lemaitre R, McLaughlin PA, Sorhannus U (2009) Phylogenetic relationships within the Pylochelidae (Decapoda: Anomura: Paguroidea): A cladistic analysis based on morphological characters. Zootaxa 2022, 1–14. doi:10.11646/zootaxa.2022.1.1 McLaughlin PA, Lemaitre R (2009) A new classification for the Pylochelidae (Decapoda: Anomura: Paguroidea) and descriptions of new taxa. Raffles Bulletin of Zoology Supplement 20, 159–231. Tsang LM, Ma KY, Ahyong ST, Chan T-Y, Chu KH (2008) Phylogeny of Decapoda using two nuclear protein-coding genes: Origin and evolution of the Repantia. Molecular Phylogenetics and Evolution 48, 359–368. doi:10.1016/j.ympev.2008.04.009 Tsang LM, Chan T-Y, Ahyong ST, Chu KH (2011) Hermit to king, or hermit to all: multiple transitions to crab-like forms from hermit crab ancestors. Systematic Biology 60, 616–629. doi:10.1093/sysbio/syr063

Pylojacquesidae McLaughlin & Lemaitre, 2001 Figure 13.30 McLaughlin and Lemaitre (2001) placed a new Australian species in a family distinct from Diogenidae and Paguridae

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Fig. 13.30.  Pylojacquesidae. a, Pylojacquesia colemani McLaughlin & Lemaitre, 2001, thoracic sternum, ventral view (c1–c5, coxae of pereopods 1–5). Mandible: b, Lemaitreopsis; c, Pylojacquesia. Right cheliped: d, Pylojacquesia colemani. Right pereopod 4 (carpus), propodus, dactylus: e, Lemaitreopsis; f, Pylojacquesia.

with which it shared some features. It differs in the notable separation of thoracic sternites 12–14, between the coxae of pereopods 3–5, the presence of small, calcified tergal plates on the first pleonite, and a prominently toothed mandible. A second genus and species was described later (­McLaughlin 2007). Both pylojacquesids occur in deep water. Dimensions are of the length of the shield (sl.), the anterior harder part of the carapace in front of the cervical groove. Diagnosis. Thoracic sternites 12–14 distinctly separated by membranous area; pleonite 1 fused to last thoracic somite (Fig. 13.30). Right cheliped larger than left. Mandible, incisor process mostly corneous, armed with prominent, acute teeth (Fig. 13.30b, c). Maxillipeds 3 widely separate at base. Lemaitreopsis McLaughlin, 2007 Diagnosis. 11 quadriserial gills. Mandibular incisor with 4 small teeth (Fig. 13.30b). Pereopod 5 chelate (Fig. 13.30e). Female pleopods 1 absent; left pleopods 2–4 unpaired. Maximum sl. 2.9 mm (single female known). Slope (500 m). Central Indo-Pacific (New Caledonia). 1 species (McLaughlin 2007).

Proceedings of the Biological Society of Washington 120, 56–62. doi:10.2988/​ 0006-324X(2007)120[56:ANGASI]2.0.CO;2 McLaughlin PA, Lemaitre R (2001) A new family for a new genus and new species of hermit crab of the superfamily Paguroidea (Decapoda: Anomura) and its phylogenetic implications. Journal of Crustacean Biology 21, 1062–1076. doi:10.1163/20021975-99990198

Xylopaguridae Gasparic, Fraaije, Robin & de Angeli, 2016 Figure 13.28, Plate 35d Xylopagurids live in the hollow cavities in bamboo, wood or in empty polychaete tubes, usually at substantial depth (Forest 1997; Lemaitre 1995). Long considered to be members of one genus of Paguridae (Lemaitre 1995) they were moved to a new family, along with similar fossil genera by Gašparič et al. (2016). Xylopagurids have an elongated subcylindrical pleon and carapace, a shield longer than the

Pylojacquesia McLaughlin & Lemaitre, 2001 Diagnosis. 13 biserial gills. Mandibular incisor with row of 7 prominent teeth (Fig. 13.30c). Pereopod 5 subchelate (Fig. 13.30f). Female pleopods 1 paired, modified; left pleopods 2–5 unpaired. Maximum sl. 4 mm. Shelf (139 m). Temperate Australia (SE Qld). 1 species (McLaughlin and Lemaitre 2001). References McLaughlin PA (2007) A new genus and species in the hermit crab family Pylojacquesidae (Crustacea: Anomura: Paguroidea) from New Caledonia.

Fig. 13.31.  Xylopaguridae. a, pleonite 6, Xylopagurus antonii Lemaitre, 1995. b, right cheliped: Xylopagurus. Shield: c, Prepylopagurus; d, Xylopagurus.

13 – Anomura – hermit crabs, porcelain crabs, king crabs, mole crabs, squat lobsters

posterior part of the carapace and operculate pleonite 6. The presence or absence of longitudinal grooves on the shield distinguishes the two genera (Fig. 13.28c,d). Dimensions are of the length of the shield (sl.), the anterior harder part of the carapace. Diagnosis. Pleonite 1 distinct from last thoracic somite. Pleonite 6 tergite strongly calcified, operculate (Fig. 13.28a). Right cheliped larger than left. Maxillipeds 3 widely separate at base. Pleopods 1, 2 paired in male, 2–4 unpaired in female. Prexylopagurus Gasparic, Fraaije, Robin & de Angeli, 2016 Diagnosis. Shield with pair of longitudinal sutures. Maximum sl. 12 mm (Pl. 35d). Slope (219–660 m; in cavities in wood or bamboo). Tropical Atlantic, Central Indo-Pacific. 3 species (Forest 1997: description of 2 species; Gašparič et al. 2016).

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Xylopagurus A. Milne Edwards, 1880 Diagnosis. Shield without pair of longitudinal sutures. Maximum sl. 12.4 mm. Intertidal–slope (0–389 m; in cavities in wood or bamboo). Tropical W Atlantic, Tropical Eastern Pacific. 4 species (Gašparič et  al. 2016: rediagnosis). Lemaitre’s (1995) key to five species includes one now in Prexylopagurus. References Forest J (1997) Présence du genre Xylopagurus A. Milne-Edwards, 1880 (Crustacea, Decapoda, Paguridae) dans l’Indo-Ouest Pacifique, avec la description de deux espèces nouvelles. Zoosystema 19, 421–435. Gašparič R, Fraaije RHB, Robin N, de Angeli A (2016) The first record of paguroids from the Eocene of Istria (Croatia) and further phylogenetic refinement of the Paguroidea (Crustacea, Anomura). Bulletin of Geosciences 91, 467–480. doi:10.3140/bull.geosci.1616 Lemaitre R (1995) A review of the hermit crabs of the genus Xylopagurus A. Milne Edwards, 1880 (Crustacea: Decapoda: Paguridae), including descriptions of two new species. Smithsonian Contributions to Zoology 570, 1–27. doi:10.5479/si.00810282.570.i

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Colour plates

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Plate 1.  Decapods at home. a, Jasus edwardsii (Hutton, 1875), Taputeranga Marine Reserve, Wellington, New Zealand; b, Sternostylus defensus (Benedict, 1902), deep-sea volcano, Galapagos Marine Reserve, Eastern Pacific; c, Alvinocaris komaii Zelnio & Hourdez, 2009 on hydrothermal vent, Lau Basin, Southwest Pacific; d, Lewindromia unidentata (Rüppell, 1830) on coral; e, Mictyris longicarpus Latreille, 1806, mudflat, Queensland, Australia; f, Leptomithrax gaimardii (H. Milne Edwards, 1834) mass swarming, Port Phillip Bay, Victoria, Australia.

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Marine Decapod Crustacea

Plate 2.  Dendrobranchiata. Aristeidae. a, Aristaeomorpha foliacea (Risso, 1827); b, Aristaeopsis edwardsiana (Johnson, 1868); c, Aristeus semidentatus Bate, 1881; d, Austropenaeus nitidus (Barnard, 1947); e, Cerataspis monstrosus Gray, 1828; f, Hemipenaeus sp.; g, Parahepomadus vaubani Crosnier, 1978. Benthesicymidae. h, Maorrancaris investigatoris (Alcock & Anderson, 1899).

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Plate 3.  Dendrobranchiata Penaeidae. a, Atypopenaeus dearmatus De Man, 1907; b, Funchalia woodwardi Johnson, 1868; c, Macropetasma africana (Balss, 1913); d, Metapenaeopsis sp.; e, Penaeopsis rectacuta (Bate, 1881); f, Parapenaeopsis hardwickii (Miers, 1878); g, Penaeus japonicus Bate, 1888; h, Trachysalambria curvirostris (Stimpson, 1860).

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Marine Decapod Crustacea

Plate 4.  Dendrobranchiata Sergestidae. a, Sicyonella liui Chan, 2020. Sicyoniidae. b–g, Sicyonia spp. unidentified. Solenoceridae. h, Haliporus taprobanensis Alcock & Anderson, 1899; i, Maximiliaeus odoceros Chan, 2012; j, Solenocera pectinulata Kubo, 1949.

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Plate 5.  Stenopodidea. Spongicolidae. a, Microprosthema lubricum Saito & Okuno, 2011; b, Microprosthema plumicorne (Richters, 1880); c, Microprosthema semileve (von Martens, 1872). Stenopodidae. d, Juxtastenopus spinulatus (Holthuis, 1946); e, Richardina spinicincta A. Milne-Edwards, 1881; f, Stenopus hispidus Olivier, 1811; g, Stenopus scutellatus Rankin, 1898.

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Marine Decapod Crustacea

Plate 6.  Caridea. Alpheidae. a, Alpheopsis chilensis Coutière, 1897; b, Alpheopsis yaldwyni Banner & Banner, 1973; c, Alpheus astrinx Banner & Banner, 1982; d, Alpheus villosus (Olivi, 1811); e, Athanopsis australis Banner & Banner, 1982; f, Betaeus truncatus Dana, 1852; g, Coronalpheus natator (Dana, 1852); h, Leptalpheus forceps Williams, 1965; i, Salmoneus ortmanni (Rankin, 1898); j, Leslibetaeus coibita Anker, Poddoubtchenko & Wehrtmann, 2006; k, Yagerocaris cozumel Kensley, 1988; l, Synalpheus hemphilli Coutière, 1909; m, Alpheus polystictus Knowlton & Keller, 1985; n, Alpheus cylindricus Kingsley, 1898, male and female in their host sponge.

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Plate 7.  Caridea. Hippolytidae. a, Alope orientalis (De Man, 1890); b, Hippolyte australiensis (Stimpson, 1860); c, Hippolyte caradina Holthuis, 1947; d, Latreutes antiborealis Holthuis, 1952; e, Leontocaris bulga Taylor & Poore, 1998; f, Nauticaris magellanica (A.MilneEdwards, 1891); g, Tozeuma carolinense Kingsley, 1878; h, Trachycaris rugosa (Bate, 1888); n, Saron marmoratus (Olivier, 1811). Merguiidae. i, Merguia rhizophorae (Rathbun, 1900) walking on mangrove prop root. Ogyrididae. j, Ogyrides hayi Williams, 1981. Thoridae. k, Eualus cranchii (Leach, 1817); l, Thor amboinensis (De Man, 1888). Lysmatidae. m, Lysmata hochi Baeza & Anker, 2008.

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Plate 8.  Caridea. Alvinocarididae. a, Alvinocaris solitaire Yahagi, Watanabe, Kojima & Beedesse, 2014. Bresillidae. b, Bresilia rufioculus Komai & Yamada, 2011. Bathypalaemonellidae. c, Bathypalaemonella sp. Campylonotidae. d, Campylonotus rathbunae Schmitt, 1926. Crangonidae. e, Aegaeon cataphractus (Olivi, 1972); f, Crangon crangon (Linnaeus, 1758); g, Lissosabinea indica (De Man, 1918); h, Parapontocaris aspera Chace, 1984; i, Metacrangon sp. Glyphocrangonidae. j, Glyphocrangon elephas Komai, 2005; k, Glyphocrangon pulchra Komai & Chan, 2013; l, Glyphocrangon confusa Komai, 2004.

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Plate 9.  Caridea. Eugonatonotidae. a, Eugonatonotus chacei Chan & Yu, 1991. Lipkiidae. b, Lipkius holthuisi Yaldwyn, 1960. Nematocarcinidae. c, Nematocarcinus sp. Rhynchocinetidae. d, Cinetorhynchus cf. manningi Okuno, 1996; e, Rhynchocinetes australis Hale, 1941; f, Rhynchocinetes kuiteri Tiefenbacher, 1983. Acanthephyridae. g, Acanthephyra armata A. Milne-Edwards, 1881; h, Acanthephyra sica Bate, 1888; i, Ephyrina sp.; j, Heterogenys microphthalma (Smith, 1885); k, Notostomus auriculatus Barnard, 1950. Oplophoridae. l, Oplophorus novaezeelandiae (De Man, 1931); m, Oplophorus typus H. Milne Edwards, 1837; n, Systellaspis debilis (A. Milne-Edwards, 1881).

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Plate 10.  Caridea. Palaemonidae ‘gnathophyllid’ genus. a, Gnathophyllum panamense Faxon, 1893. ‘hymenocerid’ genus. b, Hymenocera picta Dana, 1852. ‘palaemonine’ genera. c, Brachycarpus cf. biunguiculatus (Lucas, 1846); d, Leander cf. tenuicornis (Say, 1818); e, Palaemon serenus (Heller, 1862); f, Leander plumosus Bruce, 1994. ‘pontinine’ genera. g, Anchiopontonia hurii (Holthuis, 1981); h, Anchistus sp.; i, Ancylocaris brevicarpalis Schenkel, 1902; j, Ancylomenes sp.; k, Conchodytes tridacnae Peters, 1852; l, Coralliocaris superba (Dana, 1852); m, Jocaste sp.; n, Laomenes pardus Marin, 2009; o, Periclimenaeus sp.

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359

Plate 11.  Caridea. Palaemonidae ‘pontinine’ genera (continued). a, Pliopontonia sp.; b, Pontonia margarita Smith, 1869; c, Pontonia sp.; d, Thaumastocaris streptopus Kemp, 1922; e, Typton cf. carneus Holthuis, 1951; f, Zenopontonia rex (Kemp, 1922); g, Periclimenes perryae Chace, 1942 on its host basket star, Astrophyton sp. Chlorotocellidae. h, Chlorocurtis jactans (Nobili, 1904); i, Chlorotocella spinicaudus (H. Milne Edwards, 1837). Psalidopodidae. j, Psalidopus huxleyi Wood-Mason & Alcock, 1892. Stylodactylidae. k, Stylodactylus sp.

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Marine Decapod Crustacea

Plate 12.  Caridea. Pandalidae. a, Chlorotocoides spinicauda (De Man, 1902); b, Chlorotocus crassicornis (Costa, 1871); c, Dorodotes reflexus Bate, 1888; d, Procletes levicarina (Bate, 1888); e, Pandalus chani Komai, 1999; f, Heterocarpus sibogae De Man, 1917; g, Plesionika indica De Man, 1917; h, Thalassocaris crinita (Dana, 1852). Pasiphaeidae. i, Leptochela serratorbita Bate, 1888; j, Pasiphaea barnardi Yaldwyn, 1971; k, Alainopasiphaea australis (Hanamura, 1989); l, Eupasiphae latirostris (Wood-Mason, 1891); m, Glyphus marsupialis Filhol, 1884.

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Plate 13.  Astacidea. Enoplometopidae. a, b, Enoplometopus crosnieri Chan & Yu, 1998; c, Enoplometopus occidentalis (Randall, 184). Nephropidae. d, Acanthacaris tenuimana Bate, 1888; e, Dinochelus ausubeli Ahyong, Chan & Bouchet, 2010; f, Metanephrops velutinus Chan & Yu, 1991; g, Nephropsis sulcata Macpherson, 1990; h, Thaumastocheles massonktenos Chang, Chan & Ahyong, 2014.

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Plate 14.  Axiidea. Axiidae. a, Acanthaxius sapulo Poore, 2020; b, Alienaxiopsis clypeata (De Man, 1888); c, Allaxiopsis picteti (Zehntner, 1894); d, Bouvieraxius keiensis Sakai, 1992; e, Calaxius inhambane Poore, 2020; f, Calaxius manningi Kensley, Lin & Yu, 2000; g, Eiconaxius demani Sakai, 1992; h, Eiconaxius dongshaensis Poore & Dworschak, 2018.

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363

Plate 15.  Axiidea. Axiidae. a, Manaxius iro Poore, 2020; b, Montanaxius vadensis Poore, 2020; c, Parascytoleptus papua Poore & Collins, 2010; d, Paraxiopsis brocki (De Man, 1888); e, Paraxius altus Bate, 1888 (left cheliped missing); f, Ralumcaris bisquamosa (De Man, 1905); g, Spongiaxius brucei (Sakai, 1986).

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Marine Decapod Crustacea

Plate 16.  Axiidea. Callianassidae. a, Biffarius biformis (Biffar, 1971); b, Fragillianassa fragilis (Biffar, 1970); c, Neotrypaea caesari (Heard & Manning, 2000); d, Paratrypaea bouvieri (Nobili, 1904); e, Paratrypaea maldivensis (Borradaile, 1904); f, Pugnatrypaea emanata Felder & Robles, 2020; g, Scallasis amboinae Bate, 1888. Callianideidae. h, i, Callianidea typa H. Milne Edwards, 1837. Callichiridae. j, Callichirus seilacheri (Bott, 1955); k, Corallianassa sp.; l, Glypturoides trilobata (Biffar, 1970); m, Glypturus acanthochirus Stimpson, 1866; n, Glypturus rabalaisae Sakai, 2011; o, Lepidophthalmus eiseni Holmes, 1904; p, Mucrollichirus mucronatus (Strahl, 1862); q, Neocallichirus indicus (De Man, 1905).

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365

Plate 17.  Axiidea. Ctenochelidae. a, Ctenocheloides boucheti Poore, 2015 (left cheliped missing); b, Dawsonius latispina (Dawson, 1967); c, Paragourretia biffari (Blanco Rambla & Liñero Arana, 1994). Eucalliacidae. d, Calliaxina kensleyi (Dworschak, 2005); e, Eucalliaxiopsis dworschaki Poore, 2021; f, Eucalliaxiopsis jonesi (Heard, 1989). Micheleidae. g, Tethisea indica Poore, 1994. Strahlaxiidae. h, Neaxius acanthus (A. Milne-Edwards, 1879); i, j, Strahlaxius plectrorhynchus (Strahl, 1862).

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Marine Decapod Crustacea

Plate 18.  Gebiidea. Laomediidae. a, Axianassa sp.; b, Laomedia healyi Yaldwyn & Wear, 1970; c, Naushonia draconis Anker, 2014; d, Naushonia sp. Upogebiidae. e, Acutigebia simsoni (Thomson, 1893); f, Gebiacantha sp.; g, h, Upogebia spp.; i, Upogebia pugettensis, with commensal clam, Pseudopythina (Orobitella) rugifera, attached below pleon.

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Plate 19.  Achelata. Palinuridae. a, Jasus edwardsi novaehollandiae Holthuis, 1963; b, Sagmariasus verreauxi (H. Milne Edwards, 1851); c, Justitia longimana (H. Milne Edwards, 1837); d, Linuparus meridionalis Tsoi, Chan & Chu, 2011; e, Panulirus argus (Latreille, 1804); f, Puerulus gibbosus Chan, Ma & Chu, 2013.

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Plate 20.  Achelata. Scyllaridae Scyllarinae. a, Antarctus mawsoni (Bage, 1938); b, Chelarctus aureus (Holthuis, 1963; c, Eduarctus martensii (Pfeffer, 1881); d, Scyllarus chacei Holthuis, 1960; e, Scammarctus batei (Holthuis, 1946). Arctidinae. f, Scyllarides nodifer (Stimpson, 1866).

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Plate 21.  Achelata. Scyllaridae Ibacinae. a, Ibacus novemdentatus Gibbes, 1850; b, Parribacus antarcticus (Lund, 1793). Theninae. c, Thenus australiensis Burton & Davie, 2007. Polychelida. Polychelidae. d, eryonecius larva; e, Polycheles typhlops Heller, 1862; f, Stereomastis galil (Ahyong & Brown, 2002).

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Plate 22.  Anomura. Chirostylidae. a, Uroptychus scambus Benedict, 1902 on host coral, Indonesia. b, Gastroptychus. sp. on host coral, Gulf of Mexico; c, Uroptychodes grandirostris (Yokoya, 1933); d, Uroptychus babai Ahyong & Poore, 2004. Eumunididae. e, Eumunida funambulus Gordon, 1930.

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371

Plate 23.  Anomura. Kiwaidae. a, assemblage of Kiwa tyleri at the Black & White chimney at the E9 vent field, East Scotia Ridge; b, Kiwa hirsuta Macpherson, Jones & Segonzac, 2005; c, Kiwa tyleri Thatje in Thatje, Marsh, Roterman, Mavrogortdato & Linse, 2015. Sternostylidae. d, Sternostylus investigatoris (Alcock & Anderson, 1899).

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Marine Decapod Crustacea

Plate 24.  Anomura. Galatheidae. a, Allogalathea elegans (Adams & White, 1848); b, Lauriea punctata Macpherson & Robainas-Barcia, 2013; c, Macrothea bouchardi Macpherson & Cleva, 2010; d, Galathea australiensis Stimpson, 1858; e, Galathea rubromaculata Miyake & Baba, 1967; f, Galathea squamifera Leach, 1814; g, Galathea genkai Miyake & Baba, 1964; h, Galathea pilosa De Man, 1888; i, Phylladiorhynchus pusillus (Henderson, 1885).

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373

Plate 25.  Anomura. Munididae. a, Cervimunida johni Porter, 1903; b, Raymunida vittata Macperson, 2009; c, Munida andamanica Alcock, 1894; d, Agononida rubrizonata Macpherson & Baba, 2009; e, Sadayoshia tenuirostris Macpherson & Baba, 2010; f, Bathymunida balssi Van Dam, 1938.

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Marine Decapod Crustacea

Plate 26.  Anomura. Munidopsidae. a, Galacantha spinosa A. Milne Edwards, 1880; b, Leiogalathea laevirostris (Balss, 1913); c, Munidopsis similior Baba, 1988; d, Munidopsis pallida Alcock, 1894; e, Munidopsis sarissa Lin, Osawa & Chan, 2007; f, Shinkaia crosnieri Baba & Williams, 1998.

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Plate 27.  Anomura. Porcellanidae. a, Aliaporcellana kikuchii Nakasone & Miyake, 1969; b, Allopetrolisthes angulosus (Guérin, 1835); c, Euceramus panatelus Glassell, 1938; d, Liopetrolisthes mitra Dana, 1852; e, Lissoporcellana nakasonei (Miyake, 1978); f, Megalobrachium mortenseni Haig, 1962; g, Megalobrachium poeyi (Guérin-Méneville, 1855); h, Neopetrolisthes maculatus (H. Milne Edwards, 1837); i, Neopisosoma angustifrons (Benedict, 1901); j, Novorostrum indicum (De Man, 1893); k, Pachycheles biocellatus (Lockington, 1878); l, Pachycheles sculptus (H. Milne Edwards, 1837).

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Plate 28.  Anomura. Porcellanidae (continued). a, Petrolisthes elongatus (H. Milne Edwards, 1837); b, Petrolisthes galathinus (Bosc, 1802); c, Petrolisthes lamarckii (Leach, 1820); d, Petrolisthes novaezelandiae Filhol, 1885; e, Petrolisthes tomentosus (Dana, 1852); f, Polyonyx biunguiculatus (Dana, 1852); g, Polyonyx obesulus Miers, 1884; h, Porcellana sayana (Leach, 1820); i, Porcellanella haigae Sankarankutty, 1963.

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Plate 29.  Anomura. Albuneidae. a, Albunea elioti Benedict, 1904; b, Albunea holthuisi Boyko & Harvey, 1999; c, Austrolepidopa caledonia Boyko & Harvey, 1999; d, Lepidopa websteri Benedict, 1903; e, Paralbunea chani Boyko, 2020. Blepharipodidae. f, Blepharipoda occidentalis Randall, 1840.

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Marine Decapod Crustacea

Plate 30.  Anomura. Hippidae. a, Emerita taiwanensis Hsueh, 2105; b, Mastigochirus gracilis (Stimpson, 1858); c, Hippa pacifica (Dana, 1852). Lomisidae. d, Lomis hirta (Lamarck, 1818). Coenobitidae. e, Coenobita brevimanus Dana, 1852; f, Coenobita compressus H. Milne Edwards, 1836, feeding on coconut; g, Birgus latro (Linnaeus, 1767).

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Plate 31.  Anomura. Lithodidae. a, Cryptolithodes expansus Miers, 1879; b, Lithodes robertsoni Ahyong, 2010; c, Neolithodes bronwynae Ahyong, 2010; d, Paralithodes camtschaticus (Tilesius, 1815); e, Paralomis histrix (De Haan, 1849); f, Paralomis papua Ahyong, 2020; g, Rhinolithodes wosnessenskii Brandt, 1848.

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Plate 32.  Anomura. Diogenidae. a, Areopaguristes sp.; b, Aniculus maximus Edmondson, 1952; c, Calcinus gaimardii (H. Milne Edwards, 1848); d, Calcinus minutus Buitendijk, 1937; e, Calcinus lineapropodus Morgan & Forest, 1991; f, Cancellus typus H. Milne Edwards, 1836; g, Ciliopagurus strigatus (Herbst, 1804); h, Clibanarius cruentatus (H. Milne Edwards, 1848); i, Clibanarius eurysternus (Hilgendorf, 1878); j, Dardanus deformis (H. Milne Edwards, 1836); k, Dardanus gemmatus (H. Milne Edwards, 1848); l, Dardanus lagopodes (Forskål, 1775).

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381

Plate 33.  Anomura. Diogenidae (continued). a, Dardanus arrosor (Herbst, 1796); b, Paguristes cadenati Forest, 1954; c, Loxopagurus loxochelis (Moreira, 1901); d, Paguristes erythrops Holthuis, 1959; e, Paguristes runyanae Haig & Ball, 1988; f, Petrochirus diogenes (Linnaeus, 1758). Paguridae. g, Bathypaguropsis sp.; h, Paguritta harmsi Gordon, 1935; i, Goreopagurus poorei Lemaitre & McLaughlin, 2003; j, Nematopagurus crosnieri McLaughlin, 1998; k, Pagurodes inarmatus Henderson, 1888; l, Patagurus rex Anker & Paulay, 2013.

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Marine Decapod Crustacea

Plate 34.  Anomura. Paguridae (continued). a, Pagurus edwardsi (Dana, 1852); b, Phimochirus operculatus (Stimpson, 1859); c, Propagurus deprofundis (Stebbing, 1924); d, Propagurus haigae (McLaughlin, 1997); e, Pylopaguropsis bellula Osawa & Okuno, 2006; f, Solitariopagurus trullirostris McLaughlin, 2000; g, Spiropagurus spiriger (De Haan, 1849). Paraguridae h, Oncopagurus sp.; i, Parapagurus furici Lemaitre, 1999; j, Sympagurus affinis (Henderson, 1888); k, Sympagurus dimorphus (Studer, 1883); l, Sympagurus pictus Smith, 1883.

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Plate 35.  Anomura. Pylochelidae. a, Trizocheles sp.; b, Pylocheles sp.; c, Cheiroplatea sp. Xylopaguridae. d, Prexylopagurus sp.

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Marine Decapod Crustacea

Plate 36.  Brachyura. Cyclodorippidae. a, Corycodus minax Ahyong & Ng, 2011; b, c, Ketamia depressa Ihle, 1916 (dorsal, ventral with bivalve shell); d, Xeinostoma sakaii Tavares, 1993; e, Krangalangia orstomi Tavares, 1993; f, Tymolus brucei Tavares, 1991. Cymonomidae. g, Cymonomus chani Ahyong & Ng, 2017. Dynomenidae. h, Acanthodromia margarita (Alcock, 1899); i, Dynomene hispida (Latreille, 1812); j, Hirsutodynomene spinosa (Rathbun, 1911); k, Paradynomene quasimodo McLay & Ng, 2004.

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Plate 37.  Brachyura. Dromidiidae. a, Austrodromidia octodentata (Haswell, 1882); b, Conchoecetes intermedius Lewinsohn, 1984; c, Cryptodromia fukuii (Sakai, 1936); d, Cryptodromia sp.; e, f, Dromia erythropus (Edwards, 1771); g, Dromia personata (Linnaeus, 1758); h, Dromidiopsis australiensis (Haswell, 1882; i, j, Lauridromia intermedia (Laurie, 1906); k, Petalomera granulata Stimpson, 1858; l, Sphaerodromia ducoussoi McLay, 1991; m, Metadromia wilsoni (Fulton & Grant, 1902); n, Tumidodromia dormia (Linnaeus, 1763).

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Marine Decapod Crustacea

Plate 38.  Brachyura. Homolodromiidae. a, Dicranodromia doederleini Ortmann, 1892; b, Homolodromia kai Guinot, 1993. Homolidae. c, Gordonopsis velutina Ng & Richer de Forges, 2020; d, Homola mieensis Sakai, 1979; e, Homologenus malayensis Ihle, 1912; f, Homolomannia sibogae Ihle, 1912; g, Ihlopsis multispinosa (Ihle, 1912); h, Lamoha murotoensis (Sakai, 1979); i, Latreillopsis bispinosa Henderson, 1888; j, Moloha major (Kubo, 1936); k, Paromola japonica Parisi, 1915; l, Paromolopsis boasi Wood-Mason & Alcock, 1891; m, Yaldwynopsis saguili Richer de Forges & Ng, 2007. Latreilliidae. n, Latreillia metanesa Williams, 1982; o, Eplumula phalangium (De Haan, 1839).

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Plate 39.  Brachyura. Lyreididae. a, Lyreidus tridentatus De Haan, 1841; b, Lysirude channeri (Wood-Mason, 1885). Raninidae. c, Cosmonotus grayii Adams, 1848; d, Flaberhina balabacensis (Serène, 1971); e, Notopus dorsipes (Linnaeus, 1758); f, Ranilia muricata H. Milne Edwards, 1837; g, Ranina ranina (Linnaeus, 1758); h, Raninoides benedicti Rathbun, 1935; i, Raninoides lamarcki A. MilneEdwards & Bouvier, 1923; j, Symethis variolosa (Fabricius, 1787); k, Umalia misakiensis (Sakai, 1937).

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Marine Decapod Crustacea

Plate 40.  Brachyura. Aethridae. a, Aethra scruposa (Linnaeus, 1764); b, Drachiella morum (Alcock, 1896); c, Hepatella amica Smith, 1869; d, Hepatus pudibundus (Herbst, 1785) (juvenile); e, Osachila antillensis Rathbun, 1916; f, Sakaila japonica (Sakai, 1963). Belliidae. g, Acanthocyclus gayi Lucas, 1844. Bythograeidae. h, Gandalfus puia McLay, 2007; i, Bythograea thermydron Williams, 1980 on vent mussels, photo taken on board RV Akademik Mstislav Keldysh.

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389

Plate 41.  Brachyura. Calappidae. a, Acanthocarpus bispinosus A. Milne-Edwards, 1880; b, Calappa calappa (Linnaeus, 1758); c, Calappa granulata (Linnaeus, 1758); d, Calappa philargius (Linnaeus, 1758); e, Cryptosoma balguerii (Desbonne, 1867); f, Cycloes granulosa De Haan, 1837; g, Cyclozodion tuberatum Williams & Child, 1989; h, Mursia africana Galil, 1993; i, Paracyclois milneedwardsii Miers, 1886. Matutidae. j, Ashtoret lunaris (Forskål, 1775); k, Matuta banksii Leach, 1817; l, Matuta victor (Fabricius, 1781).

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Marine Decapod Crustacea

Plate 42.  Brachyura. Atelecyclidae. a, Atelecyclus rotundatus (Olivi, 1792). Cancridae. b, Anatolikos japonicus (Ortmann, 1893); c, Cancer pagurus Linnaeus, 1758; d, Glebocarcinus oregonensis (Dana, 1852); e, Metacarcinus novaezelandiae (Hombron & Jacquinot, 1846); f, Platepistoma anaglyptum Balss, 1922; g, Romaleon nadaense (Sakai, 1969). Carpiliidae. h, Carpilius convexus (Forskål, 1775). Cheiragonidae. i, Erimacrus isenbeckii (Brandt, 1848). Corystidae. j, Corystes cassivelaunus (Pennant, 1777); k, Gomeza bicornis Gray, 1831; l, Jonas distinctus (De Haan, 1835).

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391

Plate 43.  Brachyura. Dairidae. a, Daira perlata (Herbst, 1790). Dorippidae. b, Dorippe quadridens (Fabricius, 1793); c, Dorippoides facchino (Herbst, 1785); d, Heikeopsis japonica (von Siebold, 1824). Ethusidae. e, Ethusa orientalis Miers, 1886; f, Ethusina castro Ahyong, 2008; g, Parethusa hylophora Castro, 2005. Eriphiidae. h, Eriphia gonagra (Fabricius, 1781). Menippidae. i, Menippe mercenaria (Say, 1818); j, Myomenippe hardwickii (Gray, 1831). Oziidae. k, Epixanthus frontalis (H. Milne Edwards, 1834); l, Ozius deplanatus (White, 1847). Acidopsidae. m, Crinitocinus alcocki (Borradaile, 1902); n, Parapilumnus cristimanus (A. Milne-Edwards, 1873); o, Raoulia sp.

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Marine Decapod Crustacea

Plate 44.  Brachyura. Chasmocarcinidae. a, Angustopelta cribrorum (Rathbun, 1932); b, Camatopsis rubida Alcock & Anderson, 1899; c, Chasmocarcinops gelasimoides Alcock, 1900. Euryplacidae. d, Platyozius laevis (Borradaile, 1902); e, Trissoplax dentata (Stimpson, 1858); f, Xenocrate peculiaris Ng & Castro, 2007. Goneplacidae. g, Carcinoplax ischurodous (Stebbing, 1923); h Psopheticus crosnieri Guinot, 1990; i, Pycnoplax victoriensis (Rathbun, 1923). Mathildellidae. j, Mathildella sp. Progeryonidae. k, Rhadinoplax microphthalmus (Guinot & Richer de Forges, 1981). Scalopidiidae. l, Scalopidia spectabilis Ng & Castro, 2013. Vultocinidae. m, Vultocinus anfractus Ng & Manuel-Santos, 2007. Hexapodidae. n, Hexaplax aurantium Rahayu & Ng, 2014; o, Mariaplax granulifera (Campbell & Stephenson, 1970).

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Plate 45.  Brachyura. Hymenosomatidae. a, Amarinus laevis (Targioni Tozzetti, 1877); b, Elamena producta Kirk, 1878; c, Halicarcinus cookii (Filhol, 1835); d, Halicarcinus ovatus Stimpson, 1858; e, Halicarcinus whitei (Miers, 1876); f, Trigonplax longirostris McCulloch, 1908; Iphiculidae. g, Iphiculus spongiosus Adams & White, 1848; h, Pariphiculus cf. agariciferus Ihle, 1918; i, Pariphiculus coronatus (Alcock & Anderson, 1894).

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Marine Decapod Crustacea

Plate 46.  Brachyura. Leucosiidae Cryptocneminae. a, Cryptocnemus stimpsoni Ihle, 1915; b, Paranursia abbreviata (Bell, 1855). Ebaliinae. c, Acanthilia intermedia (Miers, 1886); d, Alox naispela Galil & Ng, 2015; e, Ancylodactyla elata (Zarenkov, 1994); f, Arcania heptacantha (De Haan, 1861); g, Arcania tuberculata Bell, 1855; h, Bellidilia undecimspinosa (Kinahan, 1856); i, Callidactylus asper Stimpson, 1871; j, Ebalia tumefacta (Montagu, 1808); k, Galilia petricola Komai & Tsuchida, 2014; l, Heterolithadia fallax (Henderson, 1893); m, Heteronucia laminata (Doflein, 1904; n, Hiplyra longimana (A. Milne Edwards, 1874).

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Plate 47.  Brachyura. Leucosiidae Ebaliinae (continued). a, Iliacantha subglobosa Stimpson, 1871; b, Ixa cylindrus (Fabricius, 1777); c, Lithadia cadaverosa Stimpson, 1871; d, Lyphira heterograna (Ortmann, 1892); e, Merocryptus boletisculpta Zarenkov, 1994; f, Myra brevimana Alcock, 1896; g, Myropsis quinquespinosa Stimpson, 1871; h, Neparilia tuberculata (Sakai, 1961); i, Nucia speciosa Dana, 1852; j, Nursilia dentata Bell, 1855; k, Parilia major Sakai, 1961; l, Phlyxia crassipes Bell, 1855; m, Persephona mediterranea (Herbst, 1794).

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Marine Decapod Crustacea

Plate 48.  Brachyura. Leucosiidae Ebaliinae (continued). a, Praebebalia fungifera Galil, 2015; b, Praosia punctata Tan & Ng, 1993; c, Pyrhila pisum (De Haan, 1841); d, Raylilia uenoi (Takeda, 1995); e, Ryphila bertrandi Galil & Ng, 2015; f, Speloeophorus pontifer (Stimpson, 1871); g, Tanaoa retpela Galil & Ng, 2015; h, Tokoyo eburnea (Alcock, 1896); i, Toru pilus (Tan, 1996); j, Urashima pustuloides (Sakai, 1961). Leucosiinae. k, Coleusia huilianae Promdam, Nabhitabhata & Galil, 2014; l, Euclosiana exquisita (Galil, 2003); m, Leucosia rubripalma Galil, 2003; n, Seulocia gagulae Galil & Ng, 2015; o, Urnalana chevertii (Haswell, 1879).

Colour plates

397

Plate 49.  Brachyura. Epialtidae. a, Chorinus heros (Herbst, 1790); b, Criocarcinus superciliosus (Linnaeus, 1767); c, Crocydocinus ewok Lee, Richer de Forges & Ng, 2019; d, e, Cyclocoeloma tuberculatum Miers, 1880; f, Doclea armata De Haan, 1839; g, Epialtus longirostris Stimpson, 1860; h, Herbstia condyliata (Fabricius, 1787); i, Hoplophrys oatesi Henderson, 1893; j, k, Huenia proteus De Haan, 1839; l, Hyastenus ambonensis Griffin & Tranter, 1986.

398

Marine Decapod Crustacea

Plate 50.  Brachyura. Epialtidae (continued). a, Leptopisa setirostris (Stimpson, 1871); b, Libinia spinosa Guérin, 1832; c, Macrocoeloma trispinosum (Latreille, 1825); d, Menaethius monoceros (Latreille, 1825); e, Micippa philyra (Herbst, 1803); f, Microlissa bicarinata (Aurivillius, 1889); g, Naxioides robillardi (Miers, 1882); h, Minyorhyncha crassa (A. Milne-Edwards, 1878); i, Neophrys inopinata Lee, Richer de Forges & Ng, 2019; j, Oxypleurodon christiani Richer de Forges & Corbari, 2012; k, Pelia rotunda A. Milne-Edwards, 1875; l, Picrocerus armatus A. Milne-Edwards, 1865.

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399

Plate 51.  Brachyura. Epialtidae (continued). a, Pteromaja maklayi Ng & Anker, 2014; b, Samadinia mosaica (Whitelegge, 1900); c, Scyra acutifrons Dana, 1851; d, Simocarcinus simplex (Dana, 1852); e, Stegopleurodon planirostrum Richer de Forges & Ng, 2009; f, Stratiolibinia bellicosa (de Oliveira, 1944); g, Tiarinia takedai Griffin & Tranter, 1986; h, Tunepugettia corbariae Lee, Richer de Forges & Ng, 2019; i, Tylocarcinus dumerilii (H. Milne Edwards, 1834); j, Xenocarcinus conicus (A. Milne-Edwards, 1865); k, Xenocarcinus depressus Miers, 1874.

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Marine Decapod Crustacea

Plate 52.  Brachyura. Inachidae. a, Camposcia retusa (Latreille, 18290; b, Oncinopus sp.; c, Dumea latipes (Haswell, 1880); d, Paratymolus cygnus Lo & Ng, 1999; e, Rhinospinosa sp. Inachoididae. f, Pyromaia tuberculata (Lockington, 1877); g, Stenorhynchus debilis (Smith, 1871).

Colour plates

401

Plate 53.  Brachyura. Majidae. a, Alcomaja miriky Ng & Richer de Forges, 2015; b, Cyclax suborbicularis (Stimpson, 1858); c, Eurynolambrus australis H. Milne Edwards & Lucas, 1841; d, Holthuija pauli Ng & Richer de Forges, 2015; e, Leptomithrax globifer Rathbun, 1918; f, Naxia aries (Guérin, 1832); g, Notomithrax ursus (Fabricius, 1787); h, Ovimaja compressipes (Miers, 1879); i, Paramaja kominatoensis Kubo, 1936; j, Paramaya ouch Ng & Richer de Forges, 2015; k, Prismatopus spatulifer (Haswell, 1881); l, Planotergum mirabile Balss, 1935.

402

Marine Decapod Crustacea

Plate 54.  Brachyura. Majidae (continued). a, Leptomithrax longipes (Thomson, 1902); b, Sakaija serenei Ng & Richer de Forges, 2015; c, Schizophrys aspera (H. Milne Edwards, 1831). Mithracidae. d, Mithraculus coryphe (Herbst, 1801); e, Mithraculus forceps A. Milne-Edwards, 1875; f, g, Mithrax hispidus (Herbst, 1790); h, Nemausa acuticornis (Stimpson, 1871); i, Pitho lherminieri (Desbonne, 1867); j, Teleophrys ruber (Stimpson, 1871); k, Thoe puella Stimpson, 1860.

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403

Plate 55.  Brachyura. Oregoniidae. a, Cyrtomaia murrayi Miers, 1885; b, Cyrtomaia sp.; c, Grypachaeus sp.; d, Macrocheira kaempferi (Temminck, 1836); e, Platymaia sp.; f, Hyas coarctatus Leach, 1815; g, Vitjazmaia crosnieri Ahyong & Richer de Forges, 2022; h, Pleistacantha ori Ahyong & Ng, 2007.

404

Marine Decapod Crustacea

Plate 56.  Brachyura. Orithyiidae. a, Orithyia sinica (Linnaeus, 1771). Crossonotidae. b, Crossonotus sp. Palicidae. c, Neopalicus contractus (Rathbun, 1902); d, Paliculus foliatus Castro, 2000; e, Pseudopalicus macromeles Castro, 2000; f, Pseudopalicus sexlobatus (Kensley, 1969).

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405

Plate 57.  Brachyura. Dairoididae. a, Dairoides kusei (Sakai, 1938). Parthenopidae. b, Aulacolambrus hoplonotus (Adams & White, 1849); c, Cryptopodia dorsalis White, 1847; d, Daldorfia rathbunae (De Man, 1902); e, Enoplolambrus pransor (Herbst, 1796); f, Furtipodia petrosa (Klunzinger, 1906); g, Garthambrus pteromerus (Ortmann, 1893); h, Heterocrypta colombiana Garth, 1940; i, Lambrachaeus ramifer Alcock, 1895; j, Neikolambrus polemistes Tan & Ng, 2003; k, Olenorfia cariei Bouvier, 1914; l, Parthenope longimanus (Linnaeus, 1758).

406

Marine Decapod Crustacea

Plate 58.  Brachyura. Parthenopidae (continued). a, Platylambrus granulatus (Kingsley, 1879); b, Rhinolambrus contrarius (Herbst, 1804); c, Solenolambrus typicus Stimpson, 1871; d, Spinolambrus pourtalesii (Stimpson, 1871); e, Thyrolambrus astroides Rathbun, 1894.

Colour plates

407

Plate 59.  Brachyura. Galenidae. a, Galene bispinosa (Herbst, 1783); b, Halimede ochtodes (Herbst, 1783). Pilumnidae Eumedoninae. c, Ceratocarcinus longimanus White, 1847; d, Eumedonus niger H. Milne Edwards, 1834; e, Harrovia longipes Lanchester, 1900; f, Permanotus purpureus (Gordon, 1934); g, Tiaramedon spinosum (Miers, 1879); h, Zebrida adamsii White, 1847. Pilumnidae Rhizopinae. i, Typhlocarcinops hamus Ng & Rahayu, 2020; j, Ceratoplax ciliata Stimpson, 1858.

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Marine Decapod Crustacea

Plate 60.  Brachyura. Pilumnidae Pilumninae. a, Glabropilumnus sp.; b, Heteropilumnus fimbriatus (H. Milne Edwards, 1834); c, Heteropilumnus stormi De Man, 1895; d, Pilumnopeus serratifrons (Kinahan,1856); e, Pilumnus acer Rathbun, 1923; f, Pilumnus dofleini Balss, 1933; g, Pilumnus fissifrons Stimpson, 1858; h, Pilumnus gemmatus Stimpson, 1860; i, Pilumnus lumpinus Bennett, 1964; j, Pilumnus monilifer Haswell, 1881; k, Pilumnus spinifer H. Milne Edwards, 1834.

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409

Plate 61.  Brachyura. Brusiniidae. a, Brusinia elongata (Sakai, 1969). Carcinidae. b, c, Carcinus maenas (Linnaeus, 1758). Geryonidae. d, Chaceon yaldwyni Manning, Dawson & Webber, 1990; e, Nectocarcinus tuberculosus A. Milne-Edwards, 1860. Ovalipidae. f, Ovalipes catharus (White, 1843). Pirimelidae. g, Pirimela denticulata (Montagu, 1808). Polybiidae. h, Liocarcinus corrugatus (Pennant, 1777); i, Necora puber (Linnaeus, 1767). j, Parathranites orientalis (Miers, 1886); Portunidae Achelouinae. k, Achelous sebae (H. Milne Edwards, 1834); l, Lupella forceps (Fabricius, 1793). Carupinae. m, Carupa sp.; n, Laleonectes nipponensis (Sakai, 1938). Necronectinae. o, Scylla serrata (Forskål, 1775).

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Marine Decapod Crustacea

Plate 62.  Brachyura. Portunidae Podophthalminae. a, Podophthalmus vigil (Fabricius, 1798). Portuninae. b, Arenaeus cribrarius (Lamarck, 1818); c, Callinectes ornatus Ordway, 1863; d, Portunus pelagicus (Linnaeus, 1758); e, Sanquerus validus (Herklots, 1851). Thalamitinae. f, Charybdis (Charybdis) miles (De Haan, 1835); g, Goniosupradens obtusifrons (Leene, 1936); h, Lissocarcinus orbicularis Dana, 1852; i, Thalamita pelsarti Montgomery, 1931; j, Thalamitoides quadridens A. Milne-Edwards, 1869; k, Zygita spinifera (Borradaile, 1902). Thiidae. l, Thia scutellata (Fabricius, 1793).

Colour plates

411

Plate 63.  Brachyura. Pseudocarcinidae. a, Pseudocarcinus gigas (Larmarck, 1818). Retroplumidae. b, Bathypluma spinifer de Saint Laurent, 1989. Domeciidae. c, Domecia glabra Alcock, 1899. Tetraliidae. d, Tetralia cinctipes Paul’son, 1875. Trapeziidae. e, Calocarcinus africanus Calman, 1909; f, Quadrella maculosa Alcock, 1898; g, Quadrella serenei Galil, 1986; h, Trapezia rufopunctata (Herbst, 1799); i, Trapezia tigrina Eydoux & Souleyet, 1842. Trichopeltariidae. j, Podocatactes hamifer Ortmann, 1893; k, Trichopeltarion cf. elegans (Guinot & Sakai, 1970).

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Marine Decapod Crustacea

Plate 64.  Brachyura. Panopeidae. a, Eurytium limosum (Say, 1818); b, Panopeus lacustris Desbonne, 1867; c, Rhithropanopeus harrisii (Gould, 1841). Pseudorhombilidae. d, Panoplax depressa Stimpson, 1871; e, Speocarcinus carolinensis Stimpson, 1859. Xanthidae Actaeinae. f, Actaea cf. spinosissima Borradaile, 1902; g, Epiactaea cf. margaritifera (Odhner, 1925); h, Gaillardiellus cf. orientalis (Odhner, 1925); i, Heteractaea ceratopus (Stimpson, 1860); j, Odhnea echinus (Alcock, 1898); k, Paractaea rufopunctata (H. Milne Edwards, 1834); l, Platyactaea setigera (H. Milne Edwards, 1834); m, Pseudoliomera helleri (A. Milne-Edwards, 1865). Antrocarcininae. n, Cyrtocarcinus truncatus (Rathbun, 1906). Banareiinae. o, Banareia cf. nobilii (Odhner, 1925); p, Calvactaea tumida Ward, 1933; q, Trichia horii (Miyake, 1940); r, Trichia indica (Sankarankutty, 1968).

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413

Plate 65.  Brachyura. Xanthidae Banareiinae (continued). a, Trichia horii (Miyake, 1940). Chlorodiellinae. b, Tweedieia laysani (Rathbun, 1906); c, Chlorodiella cytherea (Dana, 1852); d, Cyclodius obscurus (Hombron & Jacquinot, 1846); e, Liocarpilodes integerrimus (Dana, 1852); f, Pilodius areolatus (H. Milne Edwards, 1834). g, Ratha longimanus (H. Milne Edwards, 1834). Cymoinae. h, Cymo melanodactylus Dana, 1852; i, Cymo quadrilobatus (Dana, 1852).

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Marine Decapod Crustacea

Plate 66.  Brachyura. Xanthidae Etisinae. a, Etisus bargibanti Crosnier, 1987. Euxanthinae. b, Alainodaeus rimatara Davie, 1993; c, Danielea noelensis (Ward, 1935); d, Edwardsium spinimanus (H. Milne Edwards, 1834); e, Hypocolpus pararugosus Crosnier, 1997; f, Guinotellus melvillensis (H. Milne Edwards, 1834); g, Ladomedaeus serratus (Sakai, 1964); h, Medaeops granulosus (Haswell, 1882); i, Medaeus ornatus Dana, 1852; j, Psaumis cavipes (Dana, 1852). Polydectinae. k, Lybia tessellata (Latreille, 1812); Kraussiinae. l, Palapedia sp. Liomerinae. m, Bruciana pediger (Alcock, 1898); n, Liomera monticulosa (A. Milne-Edwards, 1873); o, Neoliomera insularis (Adams & White, 1849); Garthiellinae. p, Mabui calculus Naruse, Maeonsono & Ng, 2021.

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415

Plate 67.  Brachyura. Xanthidae Xanthinae. a, Demania cf. cultripes (Alcock, 1898); b, Euryxanthops cepros Davie, 19; c, Lachnopodus ponapensis (Rathbun, 1907); d, Leptodius sanguineus (H. Milne Edwards, 1834); e, Macromedaeus crassimanus (A. Milne-Edwards, 1867); f, Nanocassiope alcocki (Rathbun, 1902); g, Neoxanthias impressus (Latreille, 1812); h, Neoxanthops lineatus (A. Milne-Edwards, 1867); i, Paraxanthias cf. notatus (Dana, 1852); j, Xanthias maculatus Sakai, 1961. Zosiminae. k, Atergatis sp.; l, Atergatopsis amoyensis De Man, 1879; m, Pulcratis reticulatus Ng & Huang, 1997; n, Platypodiella spectabilis (Herbst, 1794); o, Zosimus aeneus (Linnaeus, 1758); p, Zozymodes cavipes (Dana, 1852); q, Paratergatis longimanus Sakai, 1964; r, Lophozozymus guezei Guinot, 1977.

416

Marine Decapod Crustacea

Plate 68.  Brachyura. Aphanodactylidae. a, Selwynia sp. Cryptochiridae. b, Dacryomaia sp.; c, Fizesereneia sp.; d, Fungicola utinomi (Fize & Serène, 1956); e, Fungicola fagei (Fize & Serène, 1956); f, Opecarcinus lobifrons Kropp, 1956; g, Pseudocryptochirus viridis Kropp, 1989; h, Lithoscaptus tri (Fize & Serène, 1956); i, Hapalocarcinus cf. marsupialis Stimpson, 1859; j, Utinomiella dimorpha (Henderson, 1906), female and smaller male.

Colour plates

417

Plate 69.  Brachyura. Grapsidae. a, Goniopsis cruentata (Latreille, 1803); b, Grapsus grapsus (Linnaeus, 1758), Galápagos Is; c, Grapsus albolineatus Latreille, 1812; d, Planes major (MacLeay, 1838); e, Pachygrapsus transversus (Gibbes, 1850). Leptograpsodidae. f, Leptograpsodes octodentatus (H. Milne Edwards, 1837). Percnidae. g, Percnon planissimum (Herbst, 1804). Plagusiidae. h, Euchirograpsus sp.; i, Guinusia chabrus (Linnaeus, 1758); j, Plagusia immaculata Lamarck, 1818.

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Marine Decapod Crustacea

Plate 70.  Brachyura. Sesarmidae. a, Aratus pisonii (H. Milne Edwards, 1837); b, Armases ricordi (H. Milne Edwards, 1853); c, Clistocoeloma merguiense De Man, 1888; d, Episesarma versicolor (Tweedie, 1940); e, Fasciarma fasciatum (Lanchester, 1900); f, Geosesarma dennerle Ng, Schubart & Lukhaup, 2015; g, Labuanium politum (De Man, 1888); h, Metasesarma obesum (Dana, 1851); i, Nanosesarma andersoni (De Man, 1888); j, Neosarmatium rotundifrons (A. Milne-Edwards, 1869); k, Parasesarma erythodactylum (Hess, 1865); l, Parasesarma eumolpe (De Man, 1895); m, Sarmatium crassum Dana, 1851; n, Manarma moeschii (De Man, 1892); o, Sesarmoides kraussi (De Man, 1888).

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419

Plate 71.  Brachyura. Varunidae. a, Brachynotus spinosus (H. Milne Edwards, 1853); b, Cyclograpsus granulosus H. Milne Edwards, 1853; c, Helograpsus haswellianus (Whitelegge, 1890); d, Hemigrapsus sexdentatus (H. Milne Edwards, 1837); e, Metaplax elegans De Man, 1888; f, Paragrapsus laevis (Dana, 1851); g, Varuna yui Hwang & Takeda, 1986. Camptandriidae. h, Baruna socialis Stebbing, 1904; i, Mortensenella forceps Rathbun, 1909; j, IIyogynnis microcheirum (Tweedie, 1937); k, Camptandrium sexdentatum Stimpson, 1858; l, Nasima dotilliformis (Alcock, 1900). Dotillidae. m, Ilyoplax orientalis (De Man, 1888); n, Dotilla wichmanni De Man, 1892. Heloeciidae. o, Heloecius cordiformis (H. Milne Edwards, 1837).

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Marine Decapod Crustacea

Plate 72.  Brachyura. Macrophthalmidae. a, Ilyograpsus paludicola (Rathbun, 1909); b, Macrophthalmus laevimanus H. Milne Edwards, 1852; c, Tasmanoplax latifrons (Haswell, 1881) (not natural colour); d, Venitus latreillei (Desmarest, 1822). Ocypodidae. e, Austruca perplexa (H. Milne Edwards, 1852); f, Gelasimus hesperiae (Crane, 1975); g, Minuca vocator (Herbst, 1804); h, Ocypode quadrata (Fabricius, 1787); i, Paraleptuca splendida (Stimpson, 1858); j, Tubuca urvillei (H. Milne Edwards, 1852); k, Uca maracoani (Latreille, 1802); l, Ucides occidentalis (Ortmann, 1897). Mictyridae. m, Mictyris platycheles H. Milne Edwards, 1852.

14. Brachyura – crabs

No decapod group is more diverse than the Brachyura, or true crabs. With some 7300 described species (Davie et al. 2015a, 2015b; Ng et al. 2008), they far exceed the next most speciose group, the caridean shrimps, with over 2500 species. Brachyurans occur in most tropical freshwater bioregions and marine waters worldwide except in the Antarctic, although diversity increases towards low latitudes to peak in the south-western Pacific. Brachyura Latreille, 1802, whose name means ‘short tail’, are usually recognisable by the familiar crab form, namely a short carapace, a highly reduced pleon (or abdomen) that is held folded beneath the carapace, a pair of chelipeds (pereopods 1) held folded across the front of the body, and walking legs (pereopods 2–5) that are more-or-less directed laterally. There are many variations on the theme, however, so not all brachyurans look particularly crab-like. Davie’s (2021) book, Crabs, is a richly illustrated natural history that examines the diversity, ecology, anatomy, behaviour and much more about crabs. The book includes profiles of some of the more iconic species. Given the high diversity of brachyurans, it should be no surprise that the classification has been highly fluid (Ng et al. 2008). For most of the last century, the higher classification of crabs focussed on the shape of the carapace and mouthparts, prompting higher level names such as ­Oxyrhyncha, Oxystomata, Brachyrhyncha and Brachygnatha. However, these features underpinning the ‘old’ system were significantly subject to convergent evolution making them misleading about how major groups of crabs are actually related. Over the past three to four decades, brachyuran classification focused instead on more fundamental aspects of the crab reproductive system and the internal structural framework of the body, in particular, the placement of the gonopores (Guinot 1977). Guinot (1977) recognised three major groups based on gonopore position: Podotremata, those with gonopores on the coxa of the walking legs in both sexes as in other decapods (on pereopod 3 in females, pereopod 5 in males); Heterotremata, with gonopores on the coxa of pereopod 5 in males, and on thoracic sternite 6 (corresponding to pereopod 3) in females; and Thoracotremata, those with gonopores of both sexes on the thoracic sternum. Further, the heterotremes and thoracotremes have a more complex level of internal structural organisation, exemplified in a unique skeletal feature, the sella turcica (Secretan 1998); they are collectively grouped in the section, Eubrachyura. As a result, the podotremes are often

referred to as ‘primitive’ crabs, and the eubrachyurans, ‘higher’ crabs. Eubrachyura, inclusive of Thoracotremata, are both monophyletic but the status of Podotremata, and to a lesser extent, Heterotremata, has been contentious. The presence of paired spermathecae in females has been argued to be a podotreme synapomorphy (e.g. Guinot et al. 2013; Jagt et al. 2015), but phylogenetic analyses consistently indicate that the podotremes consist of a paraphyletic series of clades leading to Eubrachyura (e.g. Ahyong et  al. 2007; Bracken et  al. 2009; Brösing et  al. 2007; Karasawa et  al. 2011; Ma et  al. 2019; Tsang et  al. 2014). Within Eubrachyura itself, phylogenetic relationships are unclear. It is evident that the thoracotremes must be derived from heterotreme-type forms, but these are yet to be formally identified – they may now be extinct or related to one or more surviving heterotreme groups, such as the potamoid freshwater crabs (Ma et  al. 2019). Thus, it is not yet known whether the extant heterotremes are paraphyletic or form a monophyletic group owing to extinction of the heterotreme lineages nearest to the thoracotremes (Ma et al. 2019; Tsang et al. 2014). The phylogenetics of Brachyura is an actively moving field of study and the classification employed here reflects these uncertainties. Given the well corroborated paraphyly of Podotremata, we recognise four sections for the podotreme crabs (Cyclodorippoida, Dromiacea, Homoloida and Raninoida) (Ahyong et al. 2007; De Grave et al. 2009; Karasawa et al. 2011; Schweitzer et al. 2018). Within Eubrachyura, we recognise the subsections Heterotremata and Thoracotremata primarily for taxonomic convenience, acknowledging that the former subsection is probably paraphyletic. The heterotreme-thoracotreme distinction, employed in the key, is usually readily observed by examining the thoracic sternum and pereopod 5 coxa of males, but some caveats obtain. In most male heterotremes, the genital papilla (penis) emerges from a gonopore on the pereopod 5 coxa to directly enter the sterno-pleonal cavity (Fig. 14.1i, j). Some heterotremes, however, have what is known as a ‘coxo-sternal condition’ (Guinot et  al. 2013: Fig.  8), which may superficially resemble the thoracotreme form. In thoracotremes, the male genital papilla is entirely independent of the pereopod 5 coxa and first emerges from the thoracic sternum (Fig. 14.1m–o). In the coxo-sternal condition, the male genital papilla emerges first from the pereopod 5 coxa, to then passes into a partially or completely enclosed groove formed by thoracic sternites 7 and 8 (or

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only sternite 8) before emerging into the sterno-pleonal cavity (Fig. 14.1k). As a result, crabs with the coxo-sternal condition could be mistaken for thoracotremes if the pereopod 5 coxa is not examined carefully for the initial emergence of the genital papilla. In thoracotremes the genital papilla emerges from or between thoracic sternites 7 and 8 (see Ng et  al. 2007: fig.  4 for examples in Grapsoidea). Therefore, in making identifications, be sure to examine both the coxa and the sternum. A further ‘complication’ is the anomaly presented by Hymenosomatoidea. Hymenosomatoids phylogenetically belong amongst the heterotremes, despite the male gonopores actually emerging from the thoracic sternum, as in thoracotremes (Fig. 14.1l). Molecular phylogenetic and wider anatomical studies show that the unusual thoracotreme-like condition in hymenosomatoids is independently derived (Ahyong et al. 2007; Guinot 2011; Guinot et al. 2013). The key below includes all recognised extant brachyuran families (marine and freshwater), but only marine groups are considered further in subsequent sections. The family and superfamily system largely follows Davie et al. (2015b) and Ng and Davie (2020), but with several modifications. Aphanodactylidae, previously considered a pinnotheroid, is closely related to various grapsoid and ocypodoid families rather than Pinnotheridae (Tsang et  al. 2018), and is here recognised in its own superfamily, Aphanodactyloidea. The two bellioid families, Belliidae and Heteroziidae, are recognised following Števčić (2005). The portunoid subfamily Nautilocorystinae, usually placed within Thiidae or Portunidae, is treated as a separate family, Nautilocorystidae. In the key, where practicable, families corresponding to their respective superfamilies are brought out together. In several cases, however, such families are brought out in separate parts of the key, reflecting use of more easily observed characters as in Calappoidea and Portunoidea, or superfamily polyphyly, as in G ­ rapsoidea and Goneplacoidea (Castro et al. 2010; Schubart et  al. 2006; Tsang et  al. 2018; Tsang et  al.

2014), for instance. Similarly, morphological heterogeneity within some families, such as Goneplacidae and Platyxanthidae, has required that they appear more than once in the key. Keys to families are also included under superfamily headings in the body of this chapter. Most brachyurans are marine, although some members of otherwise marine superfamilies, particularly grapsoids, may be found in fresh water. Adults of two grapsoid families, Gecarcinucidae and Glyptograpsidae are semi-­terrestrial and freshwater, respectively, but have marine larvae. However, four superfamilies (Gecarcinucoidea, Potamoidea, Pseudothelphusoidea, Trichodactyloidea) are wholly restricted to fresh water and all members complete their life cycles entirely in fresh water. These groups, having no marine representatives and with direct development, are known as primary freshwater crabs (Klaus et  al. 2011). Gecarcinucoidea and Potamoidea, with one and two families, respectively, are centred in Eurasia and extend to ­Australia and Africa – the socalled ‘Old World’ freshwater crabs. Pseudothelphusoidea and Trichodactyloidea, with two families each, are restricted to Central and South America – the ‘New World’ freshwater crabs (Álvarez et al. 2021). Each of the aforementioned terrestrial and freshwater families is included in the key below, but are not further treated in this work. Diagnosis. Carapace usually flat, usually with lateral edge, usually wider or as long as wide, or more or less globular; epistome short, fused to anterior and lateral margins of carapace and basal antennal articles. Exoskeleton usually calcified. Thoracic sternites 1–8 fused. Pleon short, usually flat, usually folded under carapace, tightly so in male; pleonal pleura not overlapping. Antennular peduncle articles folding in Z-shape, main flagellum short, thick, compact, aesthetascs concentrated on one side forming dense brush. Pereopod 1 chelate, pincer-like; pereopods 2–5 usually simple. Pereopod 5 of similar length to more anterior pereopods, reduced, or absent. Uropods absent or vestigial (Dromiacea, some Hymenosomatidae).

Key to sections, subsections, superfamilies and families of Brachyura Eight families of terrestrial and freshwater crabs not dealt with in this volume are marked with an asterisk*. Figures 14.1 and 14.2 illustrate key features used to distinguish the main brachyuran sections and some superfamilies. Most figure references are to those dealing with individual families. The same or similar keys to families, with more specific figure references, are placed within the larger superfamilies.

1. – 2. – 3.

Male and female genital openings coxal (Fig. 14.1a, b)���������������������������������������������������������������������������������������������������������������2 Male genital openings coxal, coxo-sternal or sternal (Fig. 14.1d, l–o); female genital openings sternal (Fig. 14.1c)��������������������������������������������������������������������������������������������������������������������������������������� Eubrachyura … 12 Maxilliped 3 operculiform (Figs 14.1p, 14.4d, h, 14.5e, f, 14.6d, 14.10u, 14.11k)��������������������������������������������������������������������3 Maxilliped 3 pediform (Figs 14.12h, 14.14o–r)����������������������������������������������������������������������������������������������������������������������������9 Carapace longer than wide. Pereopods 2–5 propodus and dactylus flattened, often forming paddle, none subchelate (Figs 14.17–14.19)���������������������������������������������������������������������������������������������������� Raninoida, Raninoidea … 4

14 – Brachyura – crabs

423



Carapace about as long as wide (slightly longer than wide to slightly wider than long). Pereopods 2–5 propodus and dactylus subcylindrical or weakly compressed, not flattened, never forming paddle pereopods 4, 5 slender, subcylindrical, pereopod 5 or 4 and 5 subchelate�������������������������������������������������������������������������5 4. Carapace widest behind anterior one-third; front at most half carapace width; surface smooth. Pleon with median spine or tubercle on either or both pleonites 3, 4 (Fig. 14.17a, b). Thoracic sternum with pleonal locking mechanism (Fig. 14.17d, e)��������������������������������������������������������������������������������������������������������������������������Lyreididae – Carapace widest in anterior one-quarter, usually at or near anterior margin; front usually more than half carapace width; surface smooth, granulate, minutely spinulate or uniformly scabrous (Figs 14.18, 14.19). Pleon without median spines or tubercles. Thoracic sternum without pleonal locking mechanism��������������Raninidae 5. Uropods visible externally as small flattened plates (rarely absent) (Fig. 14.2q, r)������������������������������������������������������������������������������������������������������������������ Dromiacea (part), Dromioidea … 6 – Uropods absent����������������������������������������������������������������������������������������������������������Cyclodorippoida, Cyclodorippoidea … 7 6. Pereopods 2, 3 similar, functioning as walking legs; pereopods 4, 5 smaller than pereopods 2, 3, subchelate, subdorsal, modified for carrying objects (Figs 14.7–14.10)������������������������������������������������������������ Dromiidae – Pereopods 2–4 functioning as walking legs; pereopod 5 markedly smaller than pereopods 2–4. No known carrying behaviour (Fig. 14.11)����������������������������������������������������������������������������������������������������� Dynomenidae 7. Carapace endostome deep, projecting anteriorly with rounded anterior margin giving buccal cavity semi-elliptical appearance. Maxilliped 3 endopod palp inserted on inner surface of merus, insertion not visible externally, palp largely concealed; exopod without flagellum (Fig. 14.4h)������������������������� Cyclodorippidae – Carapace endostome shallow, at most only slightly projecting anteriorly, giving buccal cavity rectangular appearance. Maxilliped 3 endopod palp inserted near inner angle of merus, insertion visible externally, palp fully visible. Maxilliped 3 exopod with flagellum (Figs 14.2e, 14.5e–f)��������������������������������������������������������������������8 8. Carapace square to subovate. Orbits absent; eyes fixed or only slightly movable (Fig. 14.5)���������������������� Cymonomidae – Carapace subhexagonal. Orbits developed; eyes retractile (Fig. 14.6)�������������������������������������������������������Phyllotymolinidae 9. Pereopods 2, 3 similar, as walking legs; pereopods 4–5 both reduced and subdorsal, or dorsal, chelate or subchelate (Fig. 14.12)������������������������������������������������������Dromiacea (part), Homolodromioidea, Homolodromiidae – Pereopods 2, 3 or 2–4 similar, as walking legs; pereopod 5 reduced and subchelate, or not reduced but similar to pereopods 2–4 (Figs 14.13, 14.15, 14.16)�������������������������������������Homoloida, Homoloidea … 10 10. Carapace triangular, with long neck occupying half of carapace length. Rostrum sharp, small. Eyestalk extremely long, basal segment much longer than terminal article, not included in orbit (Fig. 14.15)���������������������������������������������������������������������������������������������������������������������������������������������������������������� Latreilliidae – Carapace longitudinally oblong to sub-ovate, barrel-shaped, without long neck. Rostrum simple or bifid. Eyestalk short or long, basal segment sometimes longer than terminal article, rarely exceeding hepatic spines�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������11 11. Carapace strongly setose, linea homolica absent. Pereopods 2–5 similar, as walking legs. Pereopod 5 about as long as pereopod 4, dactylus linear, half as long as propodus, not carried dorsally (Fig. 14.16)��������������Poupiniidae – Carapace sparsely setose or glabrous, linea homolica present (Fig. 14.1e). Pereopods 2–4 similar, as walking legs. Pereopod 5 much shorter than pereopod 4, chelate or subchelate, carried dorsally (Fig. 14.13)���������Homolidae 12. Male gonopores clearly coxal, with genital papilla protruding directly from coxa of pereopod 5 (Fig. 14.1i–k)�������������������������������������������������������������������������Heterotremata (except Hymenosomatidae, Fig. 14.l) … 13 – Male gonopores sternal, with genital papilla first emerging from surface of sternite of pereopod 5 (Fig. 14.1d, m, o)����������������������������������������������������������������������������������������������������������������������������������������������������������������������� 94 Heterotremata

13. – 14.

Pereopods 2–4 only visible; pereopod 5 absent in adults (Figs 14.55, 14.56)������������������������ Hexapodoidea, Hexapodidae Pereopods 2–5 visible in adults����������������������������������������������������������������������������������������������������������������������������������������������������14 Buccal cavity trapezoid-triangular, margins distinctly convergent anteriorly. Maxilliped 3 merus triangular, tapering distally, rarely subquadrate (Fig. 14.1q). IF buccal cavity quadrate and merus

424

– 15. – 16. – 17. – 18. – 19. – 20. – 21. – 22. – 23. – 24. – 25. – 26. – 27. –

Marine Decapod Crustacea

irregularly subquadrate, carapace transversely ovate with lateral margins expanded to fully overhang folded chelipeds and walking legs������������������������������������������������������������������������������������������������������������������������������������������15 Buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular, never clearly triangular (Fig. 14.1r–u). IF carapace transversely ovate and laterally expanded to overhang walking legs, chelipeds visible dorsally���������������������������������������������������������������������������������������� 20 Carapace longitudinally ovate, distinctly longer than wide. Pereopods 2–4 dactyl slender, pointed; pereopod 5 dactylus as wide, flattened paddle. Female gonopores (vulvae) exposed, not concealed by pleon (Fig. 14.94)������������������������������������������������������������������������������������������������������������������������ Orithyioidea, Orithyiidae Carapace variously shaped, about as wide as long to wider than long. Pereopods 2–5 dactyls all slender pointed, or all flattened, paddle-like. Female gonopores (vulvae) fully concealed by pleon, not exposed�����������������16 Opening for inhalent respiratory current below frontal margin or orbits, adjacent to endostome, with distinct, shallow canal (inhalent branchial channel) along sides of buccal cavity beneath maxilliped 3 exopod, visible when maxillipeds 3 pushed aside (Fig. 14.1v)�������������������������������������������������������������� Leucosioidea … 17 Opening for inhalent respiratory current at base of cheliped, without canal along sides of buccal cavity����������������������18 Female pleonites freely articulating, not forming brood-chamber with thoracic sternum, egg-mass protruding from sides of pleon when ovigerous (Fig. 14.61)�����������������������������������������������������������������������������Iphiculidae Female pleonites mostly fused (rarely all free), forming brood-chamber with thoracic sternum, fully enclosing egg-mass (Figs 14.62–14.70)������������������������������������������������������������������������������������������������������������������Leucosiidae Pereopods 2–5 forming swimming legs, propodus and dactylus flattened, paddle-like (Fig. 14.26). Maxilliped 3 exopod without flagellum����������������������������������������������������������������������������Calappoidea (part), Matutidae Pereopods 2–5 as walking legs, dactylus and propodus slender (Figs 14.20, 14.24). Maxilliped 3 exopod with flagellum����������������������������������������������������������������������������������������������������������������������������������������������������������������������������19 Chelipeds asymmetrical, dactylus of one chela (usually right side) with curved tooth on outer proximal surface of dactylus occluding with shallow cavity on fixed finger; curved tooth absent on other cheliped dactylus (Figs 14.24, 14.25)���������������������������������������������������������������������������������� Calappoidea (part), Calappidae Chelipeds symmetrical, dactylus of both chelipeds similar (Fig. 14.20)�������������������������������������������Aethroidea, Aethridae Pereopods 4–5 inserted obliquely on body, held dorsally (Figs 14.33, 14.34)������������������������������������������Dorippoidea … 21 Pereopods 4–5 inserted laterally on body���������������������������������������������������������������������������������������������������������������������������������� 22 Pereopods 4, 5 subchelate, dactylus occluding with tubercle on propodus (Fig. 14.33). Inhalent branchial openings on carapace narrow, elongate, adjacent to and separated from cheliped by carapace wall������������� Dorippidae Pereopods 4, 5 simple, dactylus not occluding with propodus (Fig. 14.34). Inhalent branchial opening on carapace semioval or semicircular, placed against base of cheliped������������������������������������������������������������������� Ethusidae Pereopod 5 strongly reduced, rudimentary or vestigial���������������������������������������������������������������������������������������������������������� 23 Pereopod 5 subequal in size to other legs, or if smaller, functional as walking or swimming leg����������������������������������� 25 Carapace quadrate, dorsum smooth, with or without transverse ridges, anterolateral margin entire, unarmed. Pereopod 5 setose, feather-like (Fig. 14.136)������������������������������������������������� Retroplumoidea, Retroplumidae Carapace quadrate to ovate, dorsal surface rugose and granulate, never with transverse ridges; anterolateral margin with spines. Pereopod 5 simple, slender, not setose����������������������������������������������������������������������Palicoidea … 24 Pleonites 1, 2 of both sexes much shorter than other pleonites, about half or less as long as pleonite 3 (Fig. 14.96)����������������������������������������������������������������������������������������������������������������������������������������������������Palicidae Pleonites 1, 2 of both sexes about as long as pleonite 3 (Fig. 14.95)�������������������������������������������������������������� Crossotonotidae Wholly freshwater. Eggs large, developing directly into juvenile crabs; females brooding juveniles for short period������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 26 Marine or estuarine. Eggs almost always developing into planktonic zoea, rarely as megalopa; females rarely brooding hatchlings����������������������������������������������������������������������������������������������������������������������������������������31 Mandibular palp terminal segment simple, not bilobed������������������������������������������������������������������������������ Potamoidea … 27 Mandibular palp terminal segment bilobed����������������������������������������������������������������������������������������������������������������������������� 28 Mandibular palp with 2-articles. Africa����������������������������������������������������������������������������������������������������������Potamonautidae* Mandibular palp with 3-articles. Eurasia, Africa����������������������������������������������������������������������������������������������������Potamidae*

14 – Brachyura – crabs

425

28. Gonopod 1 terminal segment dilated, with complex spines, lobes��������������������������������������������Pseudothelphusoidea … 29 – Gonopod 1 terminal segment simple, undilated without complex spines, lobes���������������������������������������������������������������� 30 29. Maxilliped 3 exopod distinctly longer than ischium, extending to proximal one-third of merus. Central and South America���������������������������������������������������������������������������������������������������������������������������� Epiloboceridae* – Maxilliped 3 exopod shorter than to slightly overreaching ischium, extending at most to, proximal one-tenth of merus. Central and South America������������������������������������������������������������� Pseudothelphusidae* 30. Pereopods 2–5 dactyli with longitudinal rows of corneous spines. Maxilliped 1 endopod without portunoid lobe. Asia–Australia������������������������������������������������������������������������������������� Gecarcinucoidea, Gecarcinucidae* – Pereopods 2–5 dactyli with longitudinal rows of soft setae, without corneous spines. Maxilliped 1 endopod with portunoid lobe. Central and South America������������������������������� Trichodactyloidea, Trichodactylidae* 31. Carapace usually pyriform, usually longer than wide, sometimes squarish. Carapace, chelipeds and walking legs usually with hooked setae (sometimes very dense) used for attaching debris and objects used in camouflage (Fig. 14.1h)�����������������������������������������������������������������������������������������������������������������������Majoidea … 32 – Carapace usually wider than long. Carapace, chelipeds and walking legs with simple or plumose setae, without hooked setae����������������������������������������������������������������������������������������������������������������������������������������������������������������37 32. Antennal basal article article broad, at most twice as long as wide. Orbits present, formed by supraorbital eave, adjacent spines and postorbital spine or lobe (Figs 14.87j–x, 14.90g–m)����������������������������������������������������������������33 – Antennal basal article slender, at least twice as long as broad. Orbits absent or with narrow, weakly developed supraorbital eave and small postorbital lobe (Figs 14.78, 14.82a–h, 14.84p–w, 14.92i–n)�������������������������� 34 33. Antennal basal article broad, not expanded to form floor of orbit (Fig. 14.87j–x)��������������������������������������������������� Majidae – Antennal basal article broad, expanded to form floor of orbit (Fig. 14.90g–m)�������������������������������������������������Mithracidae 34. Orbits with narrow, weakly developed supraorbital eave partially overhanging eyes; with or without small postorbital lobe (Fig. 14.78)�����������������������������������������������������������������������������������������������������������������������������Epialtidae – Orbits absent, eyes unprotected though orbital margin usually with several small spines and postorbital spine������������������������������������������������������������������������������������������������������������������������������������������������������������������������35 35. Male pleonite 6 posterior margin concave, telson recessed into concavity of pleonite 6 (if pleonite 6 and telson fused, traces of concave posterior margin of pleonite 6 visible); pleonites 3–5 free (Fig. 14.93a–k). Gonopod 1 with prominent longitudinal groove (Fig. 14.93l–t)���������������������������������������������������������������������Oregoniidae – Male pleonite 6 distal margin straight, telson not recessed into margin (or if recessed, pleonites 3–5 fused). Gonopod 1 without prominent longitudinal groove�������������������������������������������������������������������� 36 36. Lateral parts of thoracic pleura 5–8 and often pleonite 1 integrated with carapace and fully visible in dorsal view. Pleotelson formed by fused pleonite 6 and telson in males, fused pleonites 5–6 and telson in females (Fig. 14.84o)����������������������������������������������������������������������������������������������������������������������������� Inachoididae – Lateral parts of thoracic pleura 5–8 and pleonite 1 entirely separate from carapace and not visible or only partially visible in dorsal view. Pleonite 6 and telson usually freely articulating in both sexes (Fig. 14.82t–v)��������������������������������������������������������������������������������������������������������������������������������������������������������������� Inachidae 37. Antennules folding longitudinally or almost so, or without fossa and directed anteriorly (Fig. 14.2a)�������������������������� 38 – Antennules folding transversely, obliquely or vertically, not directed anteriorly (Fig. 14.2b)������������������������������������������ 44 38. Antennal peduncle (excluding basal article) and flagellum minute, shorter than cornea (Fig. 14.2a)��������������������������������������������������������������������������������������������������������������������������������������� Bellioidea (part), Belliidae – Antennal peduncle (excluding basal article) as long as or longer than entire eye��������������������������������������������������������������� 39 39. Antenna as long as or longer than carapace (Fig. 14.31)�������������������������������������������������������������������Corystoidea, Corystidae – Antenna markedly shorter than carapace��������������������������������������������������������������������������������������������������������������������������������� 40 40. Male cheliped strongly unequal. Male telson short, rounded-subtriangular (Figs 14.2o, 14.140)������������������������������������������������������������������������������������������������������Trichopeltarioidea, Trichopeltariidae – Male chelipeds subequal. Male telson elongate, pointed (Fig. 14.2l), or rarely short, rounded (Nautilocorystidae, one species of Cancridae only)������������������������������������������������������������������������������������������������������������41 41. Carapace subhexagonal to longitudinally subrectangular. Male pleonites 1–6 and telson free. Female gonopores (vulvae) exposed externally, not covered by folded pleon (Fig. 14.2p)����� Cheiragonoidea, Cheiragonidae

426

– 42. – 43. – 44. – 45.

– 46. – 47. – 48. – 49. – 50. – 51. – 52. – 53. – 54. –

Marine Decapod Crustacea

Carapace longitudinally ovate or subcircular to transversely ovate. Male pleonites 3–5 fused. Female gonopores (vulvae) not exposed externally, fully concealed by folded pleon����������������������������������������������������������������� 42 Front divided into 2 or 3 angular lobes. Cheliped dactylus occlusal margin with prominent molariform tooth proximally, significantly larger than other teeth. Pereopod 5 dactylus spatulate, ~2.5 times as long as wide (Fig. 14.118)���������������������������������������������������������������������������������������������������Portunoidea (part), Nautilocorystidae Front divided into 3 or more angular lobes or spines. Cheliped dactylus occlusal margin with teeth of similar size. Pereopod 5 dactylus styliform or lanceolate, more than 4 times as long as wide�������������������������� Cancroidea … 43 Carapace subcircular to longitudinally ovate. Antennal flagellum densely setose, appearing ‘furry’ (Fig. 14.27)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Atelecyclidae Carapace transversely ovate. Antennal flagellum sparsely, minutely setose (Fig. 14.28)������������������������������������� Cancridae Carapace ‘thumb-nail’ shaped, dorsally smooth, surface convex transversely, straight longitudinally; entire margin with continuous fringe of long golden setae (Fig. 14.130)���������������������������������������Portunoidea (part), Thiidae Carapace variously shaped and ornamented, convex both transversely and longitudinally; margins glabrous or variously setose, but without continuous fringe of long golden setae�������������������������������������������������������� 45 Male pleonite 3 usually with transverse ridge (Fig. 14.2m). Pereopod 5 dactylus and propodus flat, broad, ovate to fusiform, forming swimming paddle, or dactylus slender, lanceolate. If male pleonite 3 without transverse ridge, pereopod 5 paddle-like, or narrowly crescentic, flattened (Libystes only)������������������������������������������������������������������������������������������������������������������������������������Portunoidea (part) … 46 Male pleonite 3 without transverse ridge. Pereopod 5 dactylus slender, similar to pereopod 4 dactylus, not forming flattened paddle; cross-section rounded to ovate, not flattened����������������������������������������������������������������������� 52 Carapace longer than wide (Fig. 14.115)�������������������������������������������������������������������������������������������������������������������� Brusiniidae Carapace as wide as or wider than long������������������������������������������������������������������������������������������������������������������������������������� 47 Male pleonites 1–6 and telson clearly demarcated, even if pleonites 3–5 immovable (Fig. 14.119c)�������������������������������� 48 Male pleonites 3–5 fused, sutures absent or only partially indicated (Fig. 14.116g)����������������������������������������������������������� 49 Stridulatory ridges present on lower surface of cheliped propodus or on pterygostomial surface of carapace. Male pleon slender, linguiform, margins weakly tapering, anterior width of pleonite 6 about two-thirds width of pleonite 3 (Figs 14.2m, 14.119)����������������������������������������������������������������������������������������������������������������� Ovalipidae Stridulatory ridges absent on carapace and chelipeds. Male pleon triangular, margins strongly tapering, anterior width of pleonite 6 half or less that of pleonite 3 (Fig. 14.117)�����������������������������������������������������������Geryonidae Chelipeds longer than longest walking leg (pereopod 2 or 3) (Figs 14.122–14.129)���������������������������������������������Portunidae Chelipeds shorter than longest walking leg������������������������������������������������������������������������������������������������������������������������������ 50 Endostomial ridges long, extending to epistome (Fig. 14.121g, h)��������������������������������������������������������������������������� Polybiidae Endostomial ridges absent or short, not extending to epistome (Fig. 14.116d)��������������������������������������������������������������������51 Pereopod 5 dactylus narrowly lanceolate to spatuliform, flattened, wider than dactylus of other legs (Fig. 14.116)���������������������������������������������������������������������������������������������������������������������������������������������������������������� Carcinidae Pereopod 5 dactylus styliform, similar to that of other legs, with longitudinal midrib, not flattened (Fig. 14.120)���������������������������������������������������������������������������������������������������������������������������������������������������������������Pirimelidae Orbits absent or reduced. Eyes vestigial, reduced; cornea absent or if present, without pigment or facets. Carapace without teeth, margins rounded (Fig. 14.23). Deep-sea hydrothermal vents�����������������������������������������������������������������������������������������������������������������������������������������Bythograeoidea, Bythograeidae Eyes and orbits well developed; cornea with facets, pigmented; carapace with or without spines. If eyes and orbits reduced, unpigmented, carapace always with teeth on margin���������������������������������������������������������53 Male pleonites all freely articulating������������������������������������������������������������������������������������������������������������������������������������������ 54 Male pleonites 3–4, 3–5 or 4–5 fused, immovable, even if some or all sutures visible�������������������������������������������������������78 Gonopod 1 slender, usually sinuous, distal part never with large spines or complex folds, apex usually recurved. Gonopod 2 minute, less than one-fourth as long as gonopod 1, sigmoidal, comma-shaped (Fig. 14.2x)���������������������������������������������������������������������������������������������������������������������������������������������������� Pilumnoidea … 55 Gonopod 1 otherwise. Gonopod 2 length one-third gonopod 1 length or longer, straight or weakly curved����������������57

14 – Brachyura – crabs

55. – 56. – 57. – 58. – 59. – 60. – 61. – 62. – 63. – 64. – 65. – 66. – 67. – 68.

427

At least one cheliped long and slender, twice carapace length or more; fingers spoon-tipped (Fig. 14.114)�����������������������������������������������������������������������������������������������������������������������������������������������������������Tanaochelidae Chelipeds about as long as or little longer than carapace; finger tips pointed, not spoon-tipped������������������������������������ 56 Carapace usually densely setose; endostomial ridges usually present. Male pleon triangular, pleonites 3–6 with margins tapering (Fig. 14.105–14.113)���������������������������������������������������������������������������������������������������������Pilumnidae Carapace usually glabrous or sparely setose; endostomial ridges absent. Male pleon distinctly T-shaped, wide at pleonite 3, strongly narrowing to pleonites 4–6 with margins subparallel (Fig. 14.104)�������������������Galenidae Carapace posterolateral margins strongly converging, posterior margin strongly narrowed, distance between coxae of pereopods 5 less than one-fifth carapace width (Fig. 14.138). Obligate symbionts on zooxanthellate scleractinian corals, usually Acropora����������������������������������������������� Trapezioidea (part), Tetraliidae Carapace posterior margin with distance between coxae of pereopods 5 equal to or exceeding one-third carapace width�������������������������������������������������������������������������������������������������������������������������������������������������������� 58 Gonopod 2 two-thirds or less as long as gonopod 1 (Fig. 14.2w, y)����������������������������������������������������������������������������������������59 Gonopod 2 as long as gonopod 1 or longer (Fig. 14.2u, v)������������������������������������������������������������������������������������������������������ 69 Cheliped merus with stridulatory ridge (rasp) on inner distal margin; ischium and merus fused (Fig. 14.48e)������������������������������������������������������������������������������������������������Goneplacoidea, Goneplacidae (part), Bathyplax Cheliped merus without stridulatory ridge on inner distal margin; ischium and merus separate���������������������������������� 60 Gonopod 2 two-thirds length of gonopod 1. Female vulva with 2 or more rounded lobes on margin (Fig. 14.49)��������������������������������������������������������������������������������������������������������������������Goneplacoidea, Litocheiridae Gonopod 2 shorter than half length of gonopod 1. Female vulva with simple margin or single vulvar cover��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������61 Male telson reaching anteriorly onto thoracic sternite 4 beyond level of cheliped coxa condyle (Figs 14.45p, 14.53b)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������62 Male telson reaching anteriorly onto thoracic sternite 4 at most to level of cheliped coxa condyle (Fig. 14.52e, f)��������������� 64 Male telson lateral margins concave; pleonite 3 narrow, not reaching laterally as far as pereopod 5 coxa (Fig. 14.53b)������������������������������������������������������������������������������������������������������������������������Goneplacoidea, Sotoplacidae Male telson lateral margins straight to convex; pleonite 3 wide, reaching laterally to pereopod 5 coxa (Fig. 14.45p)���������������������������������������������������������������������������������������������������������������������������������������������������������������������� 63 Eyes no longer than frontal width (Fig. 14.45)���������������������������������������������������������������������������Goneplacoidea, Euryplacidae Eye length 1.4 times frontal width or longer (Fig. 14.47i, o)�����������������������������������������������������������Goneplacoidea, Goneplacidae (part), Microgoneplax, Paragoneplax Male pleonites 1–3 not reaching laterally to pereopod 5 coxa leaving thoracic sternite 8 exposed either side of pleonites (Fig. 14.52e, f)���������������������������������������������������������������������������������������������� Goneplacoidea, Scalopidiidae Male pleonites 1–3 reaching laterally to pereopod 5 coxa, completely covering thoracic sternite 8�������������������������������� 65 Carapace front prominently quadridentate, flanked by triangular inner orbital angle; endostomial ridges absent (Fig. 14.40a, e)�������������������������������������������������������������������������Eriphioidea, Platyxanthidae (part), Otmaroxanthus Carapace front irregularly transverse with short median emargination or with low rounded lobes; inner orbital angle obsolete; endostomial ridges present��������������������������������������������������������������������Pseudozioidea … 66 Carapace oval-subcircular. Male thoracic sternite 3 wider than anterior width of sternite 4 (Fig. 14.133d)������������������������������������������������������������������������������������������������������������������������������������������������������� Pilumnoididae Carapace transversely ovate to subhexagonal. Male thoracic sternite 3 as wide as or narrower than anterior margin of sternite 4������������������������������������������������������������������������������������������������������������������������������������������������������������������ 67 Thoracic sternite 3 width distinctly greater than combined length of thoracic sternites 3 and 4 (Fig. 14.134f–h)�����������������������������������������������������������������������������������������������������������������������������������������������Planopilumnidae Thoracic sternite 3 width subequal to combined length of thoracic sternites 3 and 4������������������������������������������������������� 68 Carapace narrowly subovate, 1.4 times as wide as long; with 2 sharp, anteriorly recurved spines near lateral extremities. Thoracic sternites 1–2 forming approximately right-angled triangle, margins straight, width almost twice length. Antennae, when folded, not completely retracting into fossa (Fig. 14.132)���������������������������������������������������������������������������������������������������������������������������������������������������� Christmaplacidae

428



Marine Decapod Crustacea

Carapace transversely elliptical, 1.6–1.7 times as wide as long; anterolateral margins unarmed or with shallow notches. Thoracic sternites 1–2 forming broad triangle, margins concave, divergence angle obtuse although apex acute, more than twice as wide as long. Antennae, when folded, completely retracting into fossa (Fig. 14.135)������������������������������������������������������������������������������������������������������������������������ Pseudoziidae 69. Endostomial ridges absent (Fig. 14.40i)��������������������������������������������������������������������������� Eriphiioidea, Platyxanthidae (part) – Endostomial ridges present (Fig. 14.38g)������������������������������������������������������������������������������������������������������������������������������������70 70. Male telson half as wide as pleonite 3 or less���������������������������������������������������������������������������������Goneplacoidea (part) … 71 – Male telson width two-thirds as wide as pleonite 3 or more�������������������������������������������������������������������������������������������������� 73 71. Maxilliped 3 merus distolateral margin produced to acute, triangular lobe (Fig. 14.2f)���������������������������������� Conleyidae – Maxilliped 3 merus distoolateral margin rounded or obtusely angular������������������������������������������������������������������������������ 72 72. Male thoracic sternum with ‘press-button’ of pleonal-locking mechanism on anterior half of sternite 5, near suture 4/5������������������������������������������������������������������������������������������������������������������������� Goneplacidae (part) – Male thoracic sternum with ‘press-button’ of pleonal-locking mechanism on posterior half of sternite 5, near suture 5/6 (Fig. 14.51g)����������������������������������������������������������������������Progeryonidae (part), Rhadinoplax, Progeryon 73. Carapace frontal width of adults one-sixth carapace width or less. Adult male major chela distinctly longer than carapace width. Pereopods 2–5 meri with row of spines on extensor margin only, other articles unarmed (Fig. 14.131)������������������������������������������������������������������������������������������������Pseudocarcinoidea, Pseudocarcinidae – Carapace frontal width of adults one-quarter carapace width or greater. Adult male major chela length not exceeding carapace width. Pereopods 2–5 margins either all unarmed, or with numerous spines on merus, carpus and propodus������������������������������������������������������������������������������������������������������������ Eriphioidea (part) … 74 74. Eye positioned dorsally, slightly inward of anterior edge of carapace; entire lower margin of orbit visible in dorsal view when eyes retracted. Carapace posterolateral margins strongly converging, posterior margin strongly narrowed, distance between coxae of pereopods 5 less than one-fifth carapace width (Fig. 14.35)���������������������������������������������������������������������������������������������������������������������������������������������������� Dacryopilumnidae – Eye positioned on anterior edge of carapace; only upper margin of orbit visible in dorsal view when eyes retracted. Carapace posterior margin with distance between coxae of pereopods 5 equal to or exceeding one-third carapace width���������������������������������������������������������������������������������������������������������������������������������������������������������75 75. Orbits widely separated, usually placed close to or at lateral edges of anterior margin, completely closed, antenna widely separated from and excluded from inner margin of orbit (Fig. 14.36)���������������������������������� Eriphiidae – Orbits placed well inward of lateral edge of carapace, not completely closed, antenna positioned at inner edge of orbit��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������76 76. Carapace subhexagonal, anterior surface of covered in sharp, curved spines. Chelipeds and walking legs covered with numerous sharp spines all over dorsal and lateral surface; larger chela with indistinct cutting/crushing tooth (Fig. 14.37)�������������������������������������������������������������������������������������������������������������Hypothalassiidae – Carapace subhexagonal to transversely ovate; surface rugose smooth, without upright spines; anterolateral teeth low, angular or with short points. Chelipeds and walking legs smooth to tuberculate, never prominently spinose; larger chela usually with distinct molariform crushing teeth���������������������������������������������������� 77 77. Thoracic sternite 3 wider than anterior width of sternite 4. Carapace anterolateral margin (excluding exorbital tooth) with 4 broad, angular teeth or lobes, apices often pointed (Fig. 14.38)����������������������������� Menippidae – Thoracic sternite 3 narrower than to as wide as anterior width of sternite 4. Carapace anterolateral margin (excluding exorbital tooth) with 5 (rarely 4) broad, angular teeth or lobes, or margins uneven to irregularly nodulose or spinose (Fig. 14.39)�����������������������������������������������������������������������������������������������������������Oziidae 78. Antennal peduncle (excluding basal article) and flagellum minute, shorter than cornea (Fig. 14.2b)�������������������������������������������������������������������������������������������������������������������������������� Bellioidea (part), Heteroziidae – Antennal peduncle (excluding basal article) and flagellum longer than entire eye������������������������������������������������������������ 79 79. Carapace transversely ovate, prominently convex, surface smooth or finely rugose, without spines, almost glabrous. Cheliped surfaces and margins without spines; fixed finger of larger chela with massive molariform tooth proximally on occlusal margin (Fig. 14.29). Gonopod 2 distally looped, when straightened, exceeding gonopod 1 length (Fig. 14.2u)�������������������������������������������������������������� Carpilioidea, Carpiliidae

14 – Brachyura – crabs



80. – 81.



82.



83. – 84. – 85. – 86. – 87. – 88.

429

Carapace squarish to triangular, subhexagonal, or variously rounded; dorsally gently convex to almost flat; anterolateral margins usually dentate or lobate and/or setose. Cheliped carpus and merus usually with spines or tubercles and setae; fixed finger of larger chela crenulated or with rows of teeth, but without massive molariform tooth proximally on occlusal margin. Gonopod 2 shorter than or subequal to gonopod 1 length, distally straight or curved, not looped������������������������������������������������������������������������������������������������ 80 Carapace broadly triangular or diamond-shaped to cordiform, anteriorly strongly angular. Cheliped palm triangular to ovate in cross-section (Figs 14.97–14.103)������������������������������������������������������������������� Parthenopoidea … 81 Carapace subtrapezoid to transversely ovate, anteriorly relatively straight or arcuate. Cheliped palm ovate to fusiform in cross-section���������������������������������������������������������������������������������������������������������������������������������������������������� 82 Carapace approximately triangular to diamond-shaped; surface densely covered in interlocking perforated tubercles forming pavement-like surface. Cheliped palm ovate in cross-section. Male pleonites 3–4 fused, immovable, sutures visible. Gonopod 2 longer than gonopod 1, distal portion strongly recurved, as long as proximal portion (Fig. 14.97)�������������������������������������������������������������������������������������������������������������� Dairoididae Carapace approximately triangular to cordiform; surface variously tuberculate and eroded but not covered in interlocking perforated tubercles to form pavement-like surface. Cheliped palm usually trangular in cross-section. Male pleonites 3–5 fused, immovable, even if sutures visible. Gonopod 2 usually shorter than, occasionally slightly longer than, gonopod 1, distal portion straight or gently curved, usually shorter than proximal portion (Figs 14.98–14.103)����������������������������������������������������������������������������������������Parthenopidae Male genital papilla emerging from gonopore on pereopod 5 coxa to be either exposed, lying in shallow groove on sternite 7 before entering base of gonopod 1, or partially concealed by ‘supplementary plate’ formed by invagination of thoracic sternite 8 before entering base of gonopod 1 (Fig. 14.1k, n)����������������������������������������������������������������������������������������������������������������������������������Goneplacoidea (part) … 83 Male genital papilla emerging from gonopore on pereopod 5 coxa to either directly enter gonopod 1, fully exposed but not lying in shallow groove, or to be partially or almost fully concealed within closed ‘gutter’ beneath appressed margins of sternites 7 and 8, to then enter base of gonopod 1; sternite 8 without ‘supplementary plate’ (Fig. 14.1j, o)������������������������������������������������������������������������������������������������������������������������� 88 Eyes with long stalks, length about half carapace width, extending to or beyond lateral edges of carapace. Carapace widest anteriorly across exorbital spines. Interantennular septum absent (Fig. 14.47l)������������������������������������������������������������������������������������������������������������� Goneplacidae (part), Neommatocarcinus Eyes short, length less than 0.2 carapace width, positioned well inward of lateral carapace margin, not reaching lateral edge of carapace. Carapace widest near or behind midlength. Interantennular septum present���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 84 Carapace and pereopods covered with thick, coarsely setose ridges and convoluted rugosites (Fig. 14.54)���������������������������������������������������������������������������������������������������������������������������������������������������������������Vultocinidae Carapace and pereopods smooth or uneven, granular or at most with few low ridges; carapace surface glabrous or evenly setose��������������������������������������������������������������������������������������������������������������������������������������������������������� 85 Male genital papilla partially concealed by ‘supplementary plate’ of thoracic sternite 8 (Fig. 14.1n)� Chasmocarcinidae Male thoracic sternite 8 without ‘supplementary plate’; when pleon deflexed, male genital papilla exposed, lying in open groove between thoracic sternites 7 and 8��������������������������������������������������������������������������������������������������� 86 Carapace anterolateral margins unarmed or at most with low, coarsely granular convex lobes, without prominent teeth (Fig. 14.41)����������������������������������������������������������������������������������������������������������������������������������Acidopsidae Carapace anterolateral margins with 1 or more prominent acute teeth������������������������������������������������������������������������������� 87 Male pleonites 3–4 or 3–5 fused but demarcation between fused somites clearly indicated by complete transverse groove (Fig. 14.50)����������������������������������������������������������������������������������������������������������������������������Mathildellidae Male pleonites 3–4 fused, demarcation indicated only medially and at lateral margins (Fig. 14.51d)������������������������������������������������������������������������������������������������������������������������� Progeryonidae (part), Paragalene Carapace and chelipeds covered in large, nodular, wart-like, perforated tubercles, often margined with tufts of short setae. Gonopod 2 longer than gonopod 1, distal portion strongly recurved, even looped (Fig. 14.32)�������������������������������������������������������������������������������������������������������������������������������������������������� Dairoidea, Dairidae

430

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Carapace and chelipeds smooth to highly ornamented and tubercular but not with nodular, wart-like, perforated tubercles. Gonopod 2 as long as, or shorter than gonopod 1, distal portion straight�������������������������������� 89 89. Carapace regions indicated, usually well marked. Endostomial ridges usually absent or rudimentary, rarely well developed. Gonopod 2 usually shorter than one-third, occasionally half, gonopod 1 length. Usually free living���������������������������������������������������������������������������������������������������������������������������������������������������������Xanthoidea … 90 – Carapace regions absent. Endostomial ridges complete, reaching anteriorly to buccal margin. Gonopod 2 half gonopod 1 length or longer, rarely one-third length. Obligate symbionts of branching hard corals�������������������������������������������������������������������������������������������������������������������������������������������������� Trapezioidea (part) … 93 90. Male thoracic sternite 3–4 less than half as long as wide (Figs 14.142b, 14.147g)����������������������������������������������������������������91 – Male thoracic sternite 3–4 half or more times as long as wide (Fig. 14.150k–q)����������������������������������������������������������������� 92 91. Pereopods 2, 3 meri with spine on lower distal margin (Fig. 14.142d)����������������������������������������������������� Linnaeoxanthidae – Pereopods 2, 3 meri without spine on lower distal margin (Figs 14.145–14.148)���������������������������������� Pseudorhombilidae 92. Males with part of lateral extremity of sternite 8 exposed, visible between lateral end of pleonite 2 and pereopod 5 coxa. Gonopod 1 tip with complex folds (Fig. 14.144)������������������������������������������������������������������� Panopeidae – Males with sternite 8 usually completely covered by pleonite 2, rarely with lateral extremity of sternite 8 exposed. Gonopod 1 tip simple (Fig. 14.2y)����������������������������������������������������������������������������������������������������������� Xanthidae 93. Carapace rounded, dorsal surface smooth or with scattered small granules and spines. Cheliped propodus with prominent round or pointed tubercles on outer surface; merus short, without row of teeth along anterior margin (Fig. 14.137)��������������������������������������������������������������������������������������������������������������������������������� Domeciidae – Carapace subtrapezoid to transversely ovate, dorsal surface smooth or at most faintly rugose. Cheliped propodus smooth, without tubercles along outer surface; merus long to very long, anterior margin usually with row of conspicuous teeth along one-third or more of length (Fig. 14.139)������������������������������Trapeziidae Thoracotremata (plus Hymenosomatidae)

94. Antennae folding longitudinally. Orbits absent (Fig. 14.2d)�������������������������������������������������������������������������������������������������� 95 – Antennae folding transversely or obliquely. Orbits complete�������������������������������������������������� Thoracotremata (part) … 96 95. Carapace slightly wider than long to about as long as wide (excluding rostrum), pyriform, triangular, to circular; dorsal surface flat, soft, pliable, poorly calcified. Walking legs long, slender, pereopod 5 about twice carapace length or longer. Maxilliped 3 ischium subequal to or larger than merus. Free living (Figs 14.57–14.60)����������������������������������������������������������� Heterotremata (part), Hymenosomatoidea, Hymenosomatidae – Carapace longer than wide, pyriform to longitudinally ovate or rectangular; dorsal surface transversely arched, weakly to well calcified. Walking legs stout, pereopod 5 shorter than carapace length. Maxilliped 3 ischium distinctly smaller than merus (Fig. 14.2g, 14.170–14.172). Forming galls in scleractinian corals������������������������������������������������������������������Thoracotremata (part), Cryptochiroidea, Cryptochiridae 96. Carapace usually poorly calcified in females, cuticle thin, often soft; carapace firm in males. Maxilliped 3 ischium and merus fused or free. Adults commensal in molluscs, various worm phyla, echinoderms or other crustaceans, not scleractinian corals�������������������������������������������������������������������������������������������������������������������������� 97 – Carapace well calcified, hard in both sexes; usually squarish or transversely ovate. Maxilliped 3 ischium and merus free (except in some Camptandriidae). Free-living���������������������������������������������������������������������������������������� 98 97. Maxilliped 3 ischium and merus usually fused. Antennal gland opening on margin of buccal cavern. Pereopods 2–5 propodi lower distal margin unarmed, without teeth opposing flexor margin of dactylus (Figs 14.2h, 14.202–14.209)�������������������������������������������������������������������������������������������������� Pinnotheroidea, Pinnotheridae – Maxilliped 3 ischium and merus free. Antennal gland opening on surface of epistome, not on margin of buccal cavern. Pereopods 2–5 propodi lower distal margin usually with small teeth opposing flexor margin of dactylus (Fig. 14.169)���������������������������������������������������������������������������� Aphanodactyloidea, Aphanodactylidae 98. Wide rhomboidal gap between closed maxillipeds 3. Mandibles usually visible when mouthparts closed (Fig. 14.1s, t)������������������������������������������ Grapsoidea (part) plus Macrophthalmidae (Lutogemma, Tritodynamia) … 99 – Narrow rhomboidal gap between closed maxillipeds 3. Mandibles not visible when mouthparts closed (Fig. 14.1u)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������103

14 – Brachyura – crabs

431

99.

Carapace rounded-ovate to trapezoid; suborbital crest smooth, without granules; pterygostomial region with very thick, soft setae. Pereopods 2–5 with strong, fixed chitinous spines on dactylus�������������������������������������� 100 – Carapace subquadrangular; suborbital crest with small granules; pterygostomial region glabrous to moderately setose. Pereopods 2–5 unarmed or with small chitinous spines on dactylus�������������������������������������������101 100. Carapace with cervical grooves not indicated or shallow, indistinct, approximately subparallel�������������� Gecarcinidae* – Carapace with well marked cervical grooves, oblique to midline, strongly converging posteriorly, terminating at cardiac region (Fig. 14.174)������������������������������������������������������������������������������������������������Leptograpsodidae 101. Maxilliped 3 merus and ischium without oblique setose ridge. Carapace pterygostomial region sparingly setose, without pattern of reticulated setae (Fig. 14.173g–l)���������������������������������������������������������������������������������Grapsidae – Maxilliped 3 merus and ischium with distinct oblique setose ridge (Fig. 14.181a, b). Carapace pterygostomial region covered in reticulated network of short, hooked, setae (Figs 14.1t, 14.179r)������������������������� 102

Fig. 14.1.  Brachyura. a, female thoracic sternum (right side, gonopore on coxa of pereopod 3, shaded), Raninoidea. b, male thoracic sternum (left side, genital papilla on coxa of pereopod 5, shaded), Raninoidea. c, female thoracic sternum (right side, gonopore on sternites 6/7, shaded), Heterotremata, Chasmocarcinidae. d, male thoracic sternum (right side, gonopore on sternite 8, shaded), Thoracotremata, Pinnotheridae. Carapace: e, Homoloidea (with linea homolica); f, Grapsidae (with wide front); g, Ocypodidae (with narrow front). h, rostral spine with hooked setae, Epialtidae. Male thoracic sternites 7, 8, (gonopore or genital papilla on coxa of pereopod 5, shaded) (1) Heterotremata: i, Corystes; j, Thia; k, Ommatocarcinus (coxa-sternal condition); l, Hymenosomatidae; n, Chasmocarcinidae. (2) Thoracotremata: m, Cyclograpsus; o, Sesarma. Pterygostomial region, maxilliped 3, orbit: p, Cyclodorippidae (endostome produced); q, Leucosiidae (triangular buccal cavity); r, Portunoidea (rectangular buccal cavity); s, Cyclograpsinae (significant gap between maxillipeds 3, mandibles shaded); t, Sesarmidae (significant gap between maxillipeds 3, reticulated pterygostomial region); u, Varuninae (small gap between maxillipeds 3). v, buccal cavity inhalent branchial channel (maxillipeds 3 removed, Leucosiidae). w, antennules, antennae (directed anteriorly), orbit, endostome (maxillipeds 3 removed, with ridges), Polybiidae. x, antennules, antennae (directed obliquely), orbit, endostome (maxillipeds 3 removed, with ridges), Portunidae. 4–8 = thoracic sternites 4–8; c1–c5 = pereopodal coxae 1–5

432

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Fig. 14.2.  Brachyura. Antennule, antenna (minute, shaded), orbit: a, Belliidae (antennule longitudinal); b, Heteroziidae (antennule transverse). Antennule, antenna (directed anteriorly), orbit: c, Cheiragonidae; d, Hymenosomatidae. Maxilliped 3: e, Cymonomidae (endostome not produced); f, Conleyidae; g, Cryptochiridae; h, Pinnotheridae. Male pleon: i, Cancridae, Cancer; j, Cancridae, Metacarcinus; k, Carcinidae; l, Cheiragonidae; m, Ovalipidae; n, Progeryonidae, Paragalene. o, Trichopeltariidae; p, female pleon, vulvae on sternum, Cheiragonidae, Telmessus. q, r, male pleonites, 5, 6 telson, uropods, Dromiidae. s, pereopod 2 propodus, dactylus: Leptograpsodidae. Gonopods 1, 2: t, Camptandriidae (gonopod 1 only); u, Carpiliidae; v, Eriphiidae; w, Euryplacidae; x, Pilumnidae; y, Xanthidae.

102. Carapace pterygostomial surface without groove subparallel to lateral margin of buccal cavity (Figs 14.1t, 14.179r)��������������������������������������������������������������������������������������������������������������������������������������� Sesarmidae – Carapace pterygostomial surface with distinct groove, subparallel to lateral margin of buccal cavity (Fig. 14.183i, j)��������������������������������������������������������������������������������������� Varunidae (part), Cyclograpsinae 103. Carapace front narrow, usually narrower than or as wide as orbit (Figs 14.191–14.201) (if wider than orbits, maxillipeds 3 without rhomboidal gap when closed, male pleon with distinct constriction near midlength and/or gonopod 1 strongly bent, U-shaped, Shenius)�������������������������������������������� Ocypodoidea … 104 – Carapace front usually wide, twice as wide as orbits or more (Figs 14.177–14.179). Male pleon without constriction near midlength. Gonopod 1 straight����������������������������������� Grapsoidea (part) … 110 104. Orbit placed dorsally, forming longitudinal slits perpendicular to carapace front (Fig. 14.201)�����������Xenophthalmidae – Orbit transverse or sloping posteriorly, placed along anterior carapace margin�������������������������������������������������������������� 105 105. Carapace high, subglobular, longer than wide. Eyestalk as long as or shorter than cornea. Pleon similar in both sexes, broad, nearly covering most of thoracic sternum (Fig. 14.199)�����������������������������������������������������Mictyridae – Carapace subglobular to subquadrate, usually wider than long. Eyestalk longer than cornea, sometimes markedly so. Pleon usually distinctly sexually dimorphic, male pleon never substantially covering thoracic sternum�������������������������������������������������������������������������������������������������������������������������������������������������������������������� 106

14 – Brachyura – crabs

106. – 107. – 108. – 109. – 110. – 111. – 112. – 113. –

433

Gonopod 1 strongly bent, U-shaped (Fig. 14.194)������������������������������������������������������������������������������������������� Camptandriidae Gonopod 1 straight or sinuous, not strongly bent����������������������������������������������������������������������������������������������������������������� 107 Male chelipeds unequal (Fig. 14.200)������������������������������������������������������������������������������������������������������������������������Ocypodidae Male chelipeds equal������������������������������������������������������������������������������������������������������������������������������������������������������������������� 108 Carapace transversely subovate, regions weakly indicated; lateral margins unarmed behind exorbital angle. Cheliped dactylus curved, twisted, without truncate tooth on occlusal margin (Fig. 14.196)��������������������Heloeciidae Carapace quadrate or polygonal, regions usually well indicated; lateral margins with teeth or lobes behind exorbital angle. Cheliped dactylus straight, not twisted, usually with large truncate tooth at or proximal to midlength of occlusal margin������������������������������������������������������������������������������������������������������������������������������������������ 109 Male pleon with constriction near pleonite 5 (pleonite 5 narrower than 6) or pleonite 4 with transverse setose brush partially covering pleonite 5. Carapace lateral margins usually without teeth or lobes behind exorbital angle. Merus of one or more walking legs with ovate typanum (Fig. 14.195)���������������������Dotillidae Male pleon without constriction near pleonite 5, without transverse setose brush on pleonite 4. Carapace lateral margins with teeth or lobes behind exorbital angle. Merus of walking legs without typana (Figs 14.197, 14.198)������������������������������������������������������������������������������������������������������������������������������������ Macrophthalmidae Carapace distinctly subcircular to square, usually about as long as wide. Male pleonites 3–5 or 3–6 fused, immobile�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������111 Carapace ovate to rectangular, usually wider than long. Male pleonites 1–6 and telson free������������������������������������������113 Carapace frontal margin without deep cleft to receive antennules. Male chelipeds markedly unequal, larger about twice size of smaller; carpus articulating distal to proximal end of propodus�������������������������Glyptograpsidae* Carapace frontal margin usually with deep cleft on each side of midline to receive antennules. Male chelipeds subequal; carpus articulating at proximal end of propodus���������������������������������������������������������������������������112 Maxilliped 3 merus as wide as or slightly wider than ischium (Fig. 14.176k, l)�������������������������������������������������� Plagusiidae Maxilliped 3 merus narrower than ischium (Fig. 14.175m)�������������������������������������������������������������������������������������� Percnidae Orbit totally closed. Maxillipeds 3 with mesial margin meeting when closed, with faint oblique setose ridge on merus and ischium (Fig. 14.190). Associated with hydrothermal vents������������������������������������ Xenograpsidae Orbit with lateral opening. Maxillipeds 3 with narrow gape when closed, usually with distinct oblique setose ridge on merus and ischium (Fig. 14.1u). Intertidal and subtidal����������������������������������Varunidae (part)

References Ahyong ST, Lai JCY, Sharkey D, Colgan DJ, Ng PKL (2007) Phylogenetics of the brachyuran crabs (Crustacea: Decapoda): the status of Podotremata based on small subunit nuclear ribosomal RNA. Molecular Phylogenetics and Evolution 45, 576–586. doi:10.1016/j.ympev.2007.03.022 Álvarez F, Ojeda JC, Souza-Carvalho E, Villalobos JL, Magalhães C, et  al. (2021) Revision of the higher taxonomy of Neotropical freshwater crabs of the family Pseudothelphusidae, based on multigene and morphological analyses. Zoological Journal of the Linnean Society 193, 973–1001. Bracken HD, Toon A, Felder DL, Martin JW, Finley M, et al. (2009) The decapod tree of life: compiling the data and moving toward a consensus of decapod evolution. Arthropod Systematics & Phylogeny 67, 99–116. Brösing A, Richter S, Scholtz G (2007) Phylogenetic analysis of the Brachyura (Crustacea, Decapoda) based on characters of the foregut with establishment of a new taxon. Journal of Zoological Systematics and Evolutionary Research 45, 20–32. doi:10.1111/j.1439-0469.2006.00367.x Castro P, Guinot D, Ng PKL (2010) A new family for Sotoplax robertsi Guinot, 1984, with a diagnosis and key to the Goneplacoidea Macleay, 1838 (Crustacea: Decapoda: Brachyura). Zootaxa 2356, 36–56. doi:10.11646/zootaxa.2356.1.2 Davie PJF (2021) Crabs. A global natural history. Princeton University Press, Princeton and Oxford.

Davie PJF, Guinot D, Ng PKL (2015a) Chapter 71–16. Phylogeny of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 921–979. Brill, Leiden. Davie PJF, Guinot D, Ng PKL (2015b) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. De Grave S, Pentcheff ND, Ahyong ST, Chan T-Y, Crandall KA, et  al. (2009) A classification of living and fossil genera of decapod crustaceans. Raffles Bulletin of Zoology Supplement 21, 1–109. Guinot D (1977) Propositions pour une novelle classification des Crustacés, Décapodes, Brachyoures. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 285, 1049–1052. Guinot D (2011) The position of the Hymenosomatidae MacLeay, 1838, within the Brachyura (Crustacea, Decapoda). Zootaxa 2890, 40–52. doi:10.11646/zootaxa.2890.1.4 Guinot D, Tavares M, Castro P (2013) Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs

434

Marine Decapod Crustacea

(Crustacea, Decapoda, Brachyura), with new nomina for higherranked podotreme taxa. Zootaxa 3665, 1–414. doi:10.11646/zootaxa.​ 3665.1.1 Jagt J, Van Bakel BWM, Guinot D, Fraaije RHB, Artal P (2015) Chapter 71–15. Fossil Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura. (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 847–920. Brill, Leiden. Karasawa H, Schweitzer CE, Feldmann RM (2011) Phylogenetic analysis and revised classification of podotrematous Brachyura (Decapoda) Including extinct and extant families. Journal of Crustacean Biology 31, 523–565. doi:10.1651/10-3423.1 Klaus S, Yeo DC, Ahyong ST (2011) Freshwater crab origins—Laying Gondwana to rest. Zoologischer Anzeiger 250, 449–456. doi:10.1016/j. jcz.2011.07.001 Ma KY, Qin J, Lin C-W, Chan T-Y, Ng PKL, et al. (2019) Phylogenomic analyses of brachyuran crabs support early divergence of primary freshwater crabs. Molecular Phylogenetics and Evolution 135, 62–66. doi:10.1016/j.ympev.2019.02.001 Ng PKL, Davie PJF (2020) A new family and superfamily for the southern giant crab of Australia, Pseudocarcinus gigas (Lamarck, 1818) (Decapoda: Brachyura). Journal of Crustacean Biology 40, 607–626. doi:10.1093/jcbiol/ruaa058 Ng NK, Davie PJF, Schubart C, Ng PKL (2007) Xenograpsidae, a new family of grapsoid crabs (Crustacea: Brachyura) associated with shallow water hydrothermal vents. Raffles Bulletin of Zoology Supplement 16, 233–256. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Schubart CD, Cannicci S, Vannini M, Fratini S (2006) Molecular phylogeny of grapsoid crabs (Decapoda, Brachyura) and allies based on two mitochondrial genes and a proposal for refraining from current superfamily classification. Journal of Zoological Systematics and Evolutionary Research 44, 193–199. doi:10.1111/j.1439-0469.2006.00354.x Schweitzer CE, Feldmann RM, Karasawa H, Luque J (2018) Part R, Revised, Volume 1, Chapter 8S: Systematic Descriptions: Section Raninoida. In Treatise Online. pp. 1–42. KU Paleontological Institute, Lawrence. Secretan S (1998) The sella turcica and the endophragmal system of decapods. Journal of Natural History 32, 1753–1767. doi:10.1080/00222939800771271 Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Tsang LM, Schubart CD, Ahyong ST, Lai JCY, Au EYC, et al. (2014) Evolutionary history of true crabs (Crustacea: Decapoda: Brachyura) and

the origin of freshwater crabs. Molecular Biology and Evolution 31, 1173–1187. doi:10.1093/molbev/msu068 Tsang LM, Ahyong ST, Shih H-T, Ng PKL (2018) Further polyphyly of pinnotheroid crabs: the molecular phylogenetic position of the polychaete-associated Aphanodactylidae. Invertebrate Systematics 32, 92–99. doi:10.1071/IS17038

Section Cyclodorippoida Ahyong, Lai, Sharkey, Colgan & Ng, 2007 carrier crabs Cyclodorippoidans are small, long-legged crabs, many from deep water, that share with some dromiaceans the last two pairs of ‘walking legs’ being subchelate and held over the back. These legs are smaller than the two pairs of legs that are used for walking. The dactylus is folded back on to the propodus enabling the crab to carry objects. These are often a bivalve shell (Pl. 36c) but fragments of algae, pebbles, sticks or sponges have been observed (Guinot and Wicksten 2015; Wicksten 1986). Individuals of Deilocerus planus (Rathbun, 1900), a 3 mm long crab, back up to a bivalve shell before placing it over its carapace with its last legs (Wicksten 1982). The objects are assumed to serve as disguise and protection. Unlike other podotrematous crabs, cyclodorippoidans have a broad sterno-pleonal cavity like that of eubrachyurans, corroborating the sister-group relationship between the two groups found by Ahyong et al. (2007) based on analysis of 18 SrRNA. The section contains a single superfamily, Cyclodorippoidea. Tavares (1991, 1993a, 1993b, 1998) reviewed the taxonomy of the superfamily, recognising three families, two subfamilies and 18 genera. Diagnosis. Carapace about as long as wide. Maxilliped 3 operculiform. Pereopods 2–5 propodus and dactylus subcylindrical or weakly compressed, not flat; pereopods 4–5 slender, subcylindrical, pereopod 5 or 4 and 5 subchelate. Male and female genital openings coxal.

Superfamily Cyclodorippoidea Ortmann, 1892 Key to families of Cyclodorippoidea 1. – 2. –

Orbits undeveloped; eyes immovable or slightly movable (Fig. 14.5)������������������������������������������������������������� Cymonomidae Orbits developed; eyes retractile (Figs 14.3, 14.6)������������������������������������������������������������������������������������������������������������������������2 Endostome deep, projecting anteriorly with rounded anterior margin, buccal cavity apearing semielliptical (Fig. 14.4a–e). Maxilliped 3 palp inserted on inner surface of merus, insertion not visible externally; exopod without flagellum (Fig. 14.4h)������������������������������������������������������������������������������������� Cyclodorippidae Endostome shallow, with ill-defined anterior margin, buccal cavity appearing rectangular. Maxilliped 3 palp inserted near inner angle of merus, insertion visible externally; exopod with flagellum (Fig. 14.6d)�����������������������������������������������������������������������������������������������������������������������������������������������������Phyllotymolinidae

14 – Brachyura – crabs

References Ahyong ST, Lai JCY, Sharkey D, Colgan DJ, Ng PKL (2007) Phylogenetics of the brachyuran crabs (Crustacea: Decapoda): the status of Podotremata based on small subunit nuclear ribosomal RNA. Molecular Phylogenetics and Evolution 45, 576–586. doi:10.1016/j.ympev.2007.03.022 Guinot D, Wicksten MK (2015) Chapter 71–11. Camouflage: carrying behaviour, decoration behaviour, and other modalities of concealment in Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 583–638. Brill, Leiden. Tavares MS (1991) Révision préliminaire du genre Tymolus Stimpson, avec la description de Tymolus brucei sp. nov. d’Australie occidentale (Crustacea, Brachyura, Cyclodorippoidea). Bulletin du Muséum National d’Histoire Naturelle , Paris, 4e série, A 13, 439–456. Tavares MS (1993a) Crustacea Decapoda: Les Cyclodorippidae et Cymonomidae de l’Indo-Ouest Pacifique à l’exclusion du genre Cymonomus. In: Crosnier A (Ed.), Résultats des campagnes MUSORSTOM Vol. 10. Mémoires du Muséum National d’Histoire Naturelle, Paris 156, 253–313. Tavares MS (1993b) Description préliminaire de quatre nouveaux genres et trois nouvelles espèces de Cyclodorippoidea Américains (Crustacea, Decapoda, Brachyura). Vie et Milieu 43, 137–144. Tavares MS (1998) Phyllotymolinidae, nouvelle famille de Brachyoures Podotremata (Crustacea, Decapoda). Zoosystema 20, 109–122. Wicksten M (1982) Behavior in Clythrocerus planus (Rathbun, 1900) (Brachyura, Dorippidae). Crustaceana 43, 306–308. doi:10.1163/156854082X00245 Wicksten MK (1986) Carrying behavior in brachyuran crabs. Journal of Crustacean Biology 6, 364–369. doi:10.2307/1548176

435

Cyclodorippidae Ortmann, 1892 carrier crabs Figures 14.3, 14.4, Plate 36a–f Cyclodorippidae is one of few families of primitive crabs known as carrier crabs. Tavares (1992) recognised two subfamilies on differences in the way that the embryos are incubated. In cyclodorippines the female pleon is calcified and forms a chamber enclosing the brood of embryos (Fig. 14.4n–p), much as in typical crabs. The female pleon in xeinostomatines comprises six or seven poorly calcified short narrow segments carrying calcified pleopods held laterally to form a basket supporting the brood (Fig. 14.4q–s) – this is unique within Brachyura. Tavares (1993a) provided a key to these two subfamilies and nine genera, two of which have since been removed to a separate family, Phyllotymolinidae (Tavares 1998); other genera are now included (Tavares 1993b). Diagnosis. Carapace ovate to trapezoid. Orbit developed; eyes retractile. Endostome deep, projecting anteriorly with rounded anterior margin giving buccal cavity semi-elliptical appearance. Maxilliped 3 endopod palp inserted on inner surface of merus, insertion not visible externally, palp largely concealed; exopod without flagellum.

Key to subfamilies and genera of Cyclodorippidae 1. – 2. – 3. – 4. – 5. – 6. –

Female pleopods 2–5 articulating on inner surface of pleonites, not at margins (Fig. 14.4o). Distance between exorbital teeth less than half to two-thirds maximum carapace width. Male pleon of 5 or 6 segments���������������������������������������������������������������������������������������������������������������������������������������������������Cyclodorippinae … 2 Female pleopods 2–5 articulating at margins of pleonites (Fig. 14.4q–s). Distance between exorbital teeth about three-quarters maximum carapace width. Male pleon of 5 or 7 segments����������������������� Xeinostomatinae … 8 Antennules shorter than half carapace width. Antenna article 2+3 valviform (with prominent lobe) (Fig. 14.4f) or slightly flat (Fig. 14.4g) or cylindrical (Fig. 14.4c)����������������������������������������������������������������������������������������3 Antennules longer than half carapace width. Antenna article 2+3 cylindrical (Figs 14.3g, h, 14.4e)���������������������������������6 Carapace subhexagonal, much wider than long (Fig. 14.3e). Cheliped fingers longer than palm, slender, with erect spines (Fig. 14.4j, m)�������������������������������������������������������������������������������������������������������������������������������������������������4 Carapace subcircular (Fig. 14.3a, d). Cheliped fingers about as long as palm, without erect spines (Fig. 14.4i, l)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������5 Distance between exorbital teeth half carapace width (Fig. 14.3e). Antenna separated from orbit by suborbital lobe (Fig. 14.4d)��������������������������������������������������������������������������������������������������������������������������������� Neocorycodus Distance between exorbital teeth much less than half carapace width (Fig. 14.3b). Antenna not separated from orbit, suborbital lobe weak (Fig. 14.4b)��������������������������������������������������������������������������������������������������������� Corycodus Front 4-lobed, with shallow median cleft wider than lateral clefts. Front-orbital width greater than half carapace width (Fig. 14.3a). Antenna article 2+3 valviform (with prominent lobe) (Fig. 14.4f)��������������� Clythrocerus Front bilobed, with wide median cleft. Front-orbital width half carapace width (Fig. 14.3d). Antenna article 2+3 slightly flat (Fig. 14.4g)�������������������������������������������������������������������������������������������������������������������������������������� Deilocerus Carapace dorsal surface strongly ornamented with granules and truncate spines (Fig. 14.3f)���������������������� Simodorippe Carapace dorsal surface weakly ornamented with tubercles����������������������������������������������������������������������������������������������������7

436

Marine Decapod Crustacea

Fig. 14.3.  Cyclodorippidae. Cyclodorippinae. Carapace: a, Clythrocerus; b, Corycodus; c, Cyclodorippe; d, Deilocerus; e, Neocorycodus; f, Simodorippe. g, h, Tymolus similis (Grant, 1905) with detail of front, antennules, antennae, orbits. Xeinostomatinae. i, Ketamia handokoi Tavares, 1993. Carapace: j, k, Krangalangia, with detail of front; l, Xeinostoma.

7. – 8. – 9. –

Front 4-lobed, with 3 narrow clefts, median similar to or smaller than lateral clefts, rarely truncate; distance between exorbital teeth no more than half carapace width. Eyes aligned longitudinally (Fig. 14.3g, h). Indo-West Pacific���������������������������������������������������������������������������������������������������������������������������������Tymolus Front semicircular; distance between exorbital teeth about two-thirds carapace width. Eyes aligned transversely (Fig. 14.3c). E Atlantic���������������������������������������������������������������������������������������������������������������������Cyclodorippe Front triangular, with V-shaped median notch, not exceeding exorbital teeth (Fig. 14.3i). Female pleonites and telson free (Fig. 14.4q)��������������������������������������������������������������������������������������������������������������������������������������������Ketamia Front rounded or triangular, exceeding exorbital teeth. Female pleon of 5 pleonites plus fused pleotelson (Fig. 14.4r, s)���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������9 Front trilobed, ending in a prominent triangular rostrum; orbit shallow, upper and lower margins ill-defined (Fig. 14.3j, k). Pleotelson tapering, twice as wide as long (Fig. 14.4r)����������������������������������������Krangalangia Front semicircular, fringed with small teeth; orbit deep, upper and lower margins well defined (Fig. 14.3l). Pleotelson almost parallel-sided, 3 times as long as wide (Fig. 14.4s)��������������������������������������������������������������Xeinostoma

Subfamily Cyclodorippinae Ortmann, 1892 Diagnosis. Distance between exorbital teeth at most twothirds maximum carapace width. Female pleopods 2–5 articulating on inner surface of pleonites, not at margins. Male pleon of 5 or 6 segments. Implicit attributes of genera of Cyclodorippinae. Orbit open to antennal cavity, suborbital lobe small; eyes aligned longitudinally or slightly obliquely, retractable. Only

maxillipedal 3 exopod without flagellum. Cheliped fingers about as long as palm, without erect spines. Female pleon of 5 pleonites, plus pleotelson. Clythrocerus A. Milne-Edwards & Bouvier, 1899 Diagnosis. Cyclodorippinae. Carapace subcircular, almost flat; distance between exorbital teeth about two-thirds carapace width; front 4-lobed, with shallow median cleft wider than lateral clefts.

14 – Brachyura – crabs

437

Fig. 14.4.  Cyclodorippidae. Right orbit, ventral, eye, antenna, distal maxilliped 3/endostome: a, Clythrocerus; b, Corycodus; c, Cyclodorippe; d, Neocorycodus; e, Tymolus. Antenna: f, Clythrocerus; g, Deilocerus. Maxilliped 3: h, Corycodus. Cheliped: i, Clythrocerus; j, Corycodus; k, Cyclodorippe; l, Deilocerus; m, Neocorycodus. Female pleon, telson: n, o, Cyclodorippe with detail of ventral pleopod attachment; p, Tymolus; q Ketamia, r, Krangalangia; s, Xeinostoma.

Antennule shorter than half carapace width. Antenna article 2+3 valviform. Endostome overlapping front, visible in anterior view. Maximum cl. 7 mm. Subtidal–slope (0.5–1036 m). Temperate Northern Atlantic, Tropical W Atlantic. 6 species (Campos and de Melo 1999a: key to 4 Brazilian species; Goeke and Heard 1984: figures, outdated taxonomy; Tavares 1993a: descriptions, key to 5 species; Tavares 1996: rediagnosis, key to 5 species).

front semicircular. Eyes aligned transversely, not retractable. Antennule longer than half carapace width. Antenna article 2+3 cylindrical. Endostome not overlapping front, invisible in anterior view. Maximum cl. 6.2 mm. Shelf, slope (90–686 m). Tropical Atlantic (Gulf of Mexico, Caribbean, Brazil). 7 species (Campos and de Melo 1999b: key to 3 Brazilian species; Tavares 1991: discussion of genus; Tavares 1996: rediagnosis, key to 6 species).

Corycodus A. Milne-Edwards, 1880

Deilocerus Tavares, 1993

Diagnosis. Cyclodorippinae. Carapace subhexagonal, much wider than long, swollen; distance between exorbital teeth much less than half carapace width; front semicircular-truncate, deflexed. Antennule shorter than half carapace width. Antenna article 2+3 valviform. Endostome not overlapping front, invisible in anterior view. Cheliped fingers longer than palm, cylindrical, thin, armed with long spines. Female pleon of 6 pleonites, plus telson. Maximum cl. 5.4 mm (Pl. 36a). Shelf, slope (113–647 m). Tropical Atlantic (Caribbean), Western and Central Indo-Pacific, Temperate Southern Africa. 7 species (Ahyong and Ng 2011; Naruse 2013: key to 4 species of W Indian Ocean; Tavares 1996: rediagnosis, key to 5 species).

Diagnosis. Cyclodorippinae. Carapace subcircular, almost flat; distance between exorbital teeth half carapace width; front bilobed, with broad median gap. Antennule shorter than half carapace width. Antenna article 2+3 slightly flat. Endostome overlapping front, visible in anterior view. Maximum cl. 6 mm. Subtidal–slope (18–270 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 8 species (Campos and de Melo 1998; Tavares 1993b; 1996: rediagnosis, key to 6 species; Tavares 1999).

Cyclodorippe A. Milne-Edwards, 1880 Diagnosis. Cyclodorippinae. Carapace subcircular, regions defined; distance between exorbital teeth about two-thirds carapace width;

Neocorycodus Tavares, 1993 Diagnosis. Cyclodorippinae. Carapace subhexagonal, much wider than long, swollen; distance between exorbital teeth half carapace width; front truncate, with broad median cleft. Orbit separate from antennal cavity, suborbital lobe well developed, closing orbital cavity. Antennule shorter than half carapace width.

438

Marine Decapod Crustacea

Antenna article 2+3 valviform. Endostome overlapping front, visible in anterior view. Maxillipedal 1–3 exopods without flagellum. Cheliped fingers longer than palm, cylindrical, thin, armed with long spines. Maximum cl. 3.5 mm. Shelf (67–180 m). Tropical Atlantic (Caribbean). 1 species (Tavares 1993b, 1996).

Simodorippe Chace, 1940 Diagnosis. Cyclodorippinae. Carapace subcircular, granular; distance between exorbital teeth half carapace width; front truncate, upturned, with 2 pairs of small teeth. Antennule longer than half carapace width. Antenna article 2+3 cylindrical. Endostome not overlapping front, invisible in anterior view. Maxillipedal 1–3 exopods without flagellum. Maximum cl. 12 mm. Slope (400 m). Tropical Atlantic. 1 species (Chace 1940; Tavares 1996: redescription).

Tymolus Stimpson, 1858 Diagnosis. Cyclodorippinae. Carapace subcircular, regions defined; distance between exorbital teeth less than half carapace width; front 4-lobed, with 3 narrow clefts, median similar to or smaller than lateral clefts, rarely truncate. Antennule longer than half carapace width. Antenna article 2+3 cylindrical. Endostome not overlapping front, invisible in anterior view. Maximum cl. 10 mm (Pl. 36f). Shelf, slope (44–950 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia (S Australia). 10 species (Ahyong et al. 2009: key to 4 Taiwanese species; Spiridonov and Turkay 2007; Tan and Huang 2000; Tavares 1991: key to 5 species; Tavares 1993a: rediagnosis, key to 6 species).

Subfamily Xeinostomatinae Tavares, 1992 Diagnosis. Distance between exorbital teeth about threequarters maximum carapace width. Female pleopods 2–5 articulating at margins of pleonites. Male pleon of 5 or 7 segments. Ketamia Tavares, 1992 Diagnosis. Xeinostomatinae. Front triangular, with V-shaped median notch, not exceeding exorbital teeth. Orbit deep, upper and lower margins well defined. Female pleon of 6 segments plus telson. Maximum cl. 10 mm (Pl. 36b). Subtidal–slope (15–440 m). Western and Central Indo-Pacific. 6 species (Ahyong and Ng 2011: figures; Ahyong and Mendoza 2021: key to species).

Krangalangia Tavares, 1992 Diagnosis. Xeinostomatinae. Front trilobed, ending in a prominent triangular rostrum. Orbit shallow, upper and lower margins ill-defined. Female pleon of 5 segments plus pleotelson tapering, twice as wide as long. Maximum cl. 6.2 mm (Pl. 36e).

Slope (411–1223 m). Central Indo-Pacific. 3 species (Tavares 1992; 1993a: rediagnosis, key to species).

Xeinostoma Stebbing, 1920 Diagnosis. Xeinostomatinae. Front semicircular, fringed with small teeth. Orbit deep, upper and lower margins well defined. Female pleon of 5 segments plus pleotelson almost parallel-sided, 3 times as long as wide. Maximum cl. 10 mm (Pl. 36c). Shelf, slope (144–750 m). Temperate Northern Pacific (Japan), Western and Central Indo-Pacific, Temperate Southern Africa. 4 species (Tavares 1993a: rediagnosis, key to 3 species; Tavares 1994). References Ahyong ST, Mendoza JCE (2021) Cyclodorippidae of the SJADES 2018 biodiversity cruise in Indonesia (Crustacea: Decapoda: Brachyura). Raffles Bulletin of Zoology Supplement 36, 258–264. Ahyong ST, Ng PKL (2011) Cyclodorippoid crabs from the Philippines collected by the PANGLAO 2004–2005 and AURORA 2007 expeditions. Zoologischer Anzeiger 250, 479–487. doi:10.1016/j.jcz.2011.06.001 Ahyong ST, Naruse T, Tan SH, Ng PKL (2009) Part II. Infraorder Brachyura: Sections Dromiacea, Raninoida, Cyclodorippoida. In Crustacean fauna of Taiwan: Brachyuran crabs, volume 1 – Carcinology in Taiwan and Dromiacea, Raninoida, Cyclodorippoida. (Eds Chan T-Y, Ng PKL, Ahyong ST, Tan SH) pp. 27–198. National Taiwan Ocean University, Keelung. Campos O, Junior, de Melo GAS (1998) Nova especie brasileira do genero Deilocerus (Brachyura, Podotremata, Cyclodorippidae). Iheringia. Série Zoologia 85, 129–132. Campos O, Junior, de Melo GAS (1999a) Nova especie brasileira do genero Clythrocerus A. Milne-Edwards & Bouvier, 1899 (Crustacea, Brachyura, Podotremata). Atlântica 21, 5–10. Campos O, Junior, de Melo GAS (1999b) Nova espécie brasileira do gênero Cyclodorippe A. Milne Edwards, 1880 (Brachyura, Podotremata, Cyclodorippidae). Atlântica 21, 37–41. Chace FA (1940) Reports on the scientific results of the Atlantis expeditions to the West Indies, under the joint auspices of the University of Havana and Harvard University, The brachyuran crabs. Torreia 1–67. Goeke GD, Heard RW (1984) A review of the genus Clythrocerus (Brachyura: Dorippidae) in the eastern Gulf of Mexico with notes on Cythrocerus stimpsoni. Gulf Research Reports 7, 351–355. doi:10.18785/ grr.0704.06 Naruse T (2013) Species of Corycodus A. Milne-Edwards, 1880 (Crustacea, Brachyura, Cyclodorippidae) collected from the Mozambique MAINBAZA and Madagascar MIRIKY expeditions, with description of a new species. In: Ahyong ST, Chan T-Y, Corbari L & Ng PKL (Eds) Tropical Deep-Sea Benthos vol. 27. Mémoires du Muséum National d’Histoire Naturelle 204, 485–494. Spiridonov VA, Turkay M (2007) Podotreme crabs from the western Indian Ocean: description of a new species of Tymolus Stimpson 1858 with a review of regional distribution of the Cyclodorippidae (Crustacea: Decapoda: Brachyura: Podotremata). Arthropoda Selecta 16, 1–9. Tan SH, Huang JF (2000) Description of a new species of Tymolus (Crustacea: Decapoda: Brachyura: Cyclodorippidae) from Taiwan. In Proceedings of the International Symposium on Marine Biology in Taiwan  – Crustacea and zooplankton taxonomy, ecology and living resources, 26–27 May, 1998, Taiwan. (Eds Hwang J-S, Wang C-H, Chan T-Y) pp. 135–140. National Taiwan Museum, Taipei.

14 – Brachyura – crabs

Tavares MS (1991) Révision préliminaire du genre Tymolus Stimpson, avec la description de Tymolus brucei sp. nov. d’Australie occidentale (Crustacea, Brachyura, Cyclodorippoidea). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 13, 439–456. Tavares MS (1992) Tendances évolutives chez les crabes primitifs, avec la description d’un nouvaeau type de chambre incubatrice (Crustacea, Decapoda: Cyclodorippinae Ortmann, 1892, et Xeinostominae subfam. nov.). Comptes Rendus Hebdomadaires des Séances de l’Académie des Sciences, Série III 314, 509–514. Tavares MS (1993a) Crustacea Decapoda: Les Cyclodorippidae et Cymonomidae de l’Indo-Ouest Pacifique à l’exclusion du genre Cymonomus. In: Crosnier A (Ed.), Résultats des campagnes MUSORSTOM Vol. 10. Mémoires du Muséum National d’Histoire Naturelle, Paris 156, 253–313. Tavares MS (1993b) Description préliminaire de quatre nouveaux genres et trois nouvelles espèces de Cyclodorippoidea Américains (Crustacea, Decapoda, Brachyura). Vie et Milieu 43, 137–144. Tavares MS (1994) Xeinostoma inopinatum sp. nov., a new crab from Réunion Island, South Indian Ocean (Crustacea: Brachyura: Cyclodorippidae: Xeinostomatinae). Memoirs of the Museum of Victoria 54, 121–123. doi:10.24199/j.mmv.1994.54.04 Tavares MS (1996) Révision systématique des Cyclodorippidae américains (Crustacea, Decapoda, Brachyura). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 18, 233–295. Tavares MS (1998) Phyllotymolinidae, nouvelle famille de Brachyoures Podotremata (Crustacea, Decapoda). Zoosystema 20, 109–122. Tavares MS (1999) Deilocerus captabilis, a new species of cyclodorippid crab from southeastern Brazil (Crustacea: Decapoda: Brachyura: Cyclodorippidae). Proceedings of the Biological Society of Washington 112, 141–144.

439

Cymonomidae carrier crabs Figure 14.5, Plate 36g Cymonomids are small, rare, cryptic primitive crabs with elongate, usually immovable eyestalks, a carapace without functional orbits, and reduced pereopods 4 and 5 held over the carapace. Most cymonomids occur in deep outer shelf or slope waters below 200 m depth and are known worldwide. Three of almost 50 species have been allocated to their own genera. The remainder belong to Cymonomus of which new species are regularly being discovered, especially from the Indo-West Pacific. It is supposed that cymonomids carry shells or other camouflage or protection as in other cyclodorippoids but this has not been directly observed. In one genus, Elassopodus, pereopods 4 and 5 are vestigial suggesting this behaviour has been lost (Guinot and Wicksten 2015). Diagnosis. Carapace square to subovate. Orbit absent; eyes fixed or only slightly movable. Endostome shallow, at most only slightly projecting anteriorly, giving buccal cavity rectangular appearance. Maxilliped 3 endopod palp inserted near inner angle of merus, insertion visible inserted externally, palp fully visible; exopod flagellum well developed or rudimentary.

Fig. 14.5.  Cymonomidae. a, Curupironomus agassizii (A. Milne-Edwards & Bouvier, 1899); b, Cymonomus soela Ahyong & Brown, 2003; c, Cymopolus asper A. Milne-Edwards, 1880; d, Elassopodus stellatus Tavares, 1993, posterolateral right carapace, pereopod 3 coxa, pereopods 4, 5, pleonites 1, 2. Maxilliped 3: e, Cymonomus; f, Cymopolus.

440

Marine Decapod Crustacea

Key to genera of Cymonomidae 1. – 2. – 3. –

Maxilliped 3 palp articulating near midlength of merus mesial margin (Fig. 14.5e). Pereopods 2, 3 thin, much longer than carapace length, pereopod 3 dactylus longer than propodus (Fig. 14.5b)���������������������Cymonomus Maxilliped 3 palp articulating with merus mesiodistally (Fig. 14.5f). Pereopods 2, 3 robust, as long or up to 1.5 times carapace length, pereopod dactylus 3 shorter than propodus (Fig. 14.5a)��������������������������������������������������������2 Rostrum short, exorbital projections longer. Eyestalk fixed to rostrum. Pereopods 4, 5 vestigial, of 1 article (Fig. 14.5d). Central Indo-Pacific������������������������������������������������������������������������������������������������������������������������� Elassopodus Rostrum prominent, triangular, exorbital projections shorter. Eyestalk mobile (Fig. 14.5a, c). Pereopods 4, 5 short, of 6 articles (Fig. 14.5a). Caribbean����������������������������������������������������������������������������������������������������������������������������3 Carapace parallel-sided; front produced, laterally concave, widest part of carapace, with broad spinose triangular rostrum and pair of spinose projections. Eyestalks and antennae almost completely hidden by rostrum (Fig. 14.5c)��������������������������������������������������������������������������������������������������������������������������������������������� Cymopolus Carapace lateral margins convex, widest at midpoint; front produced, slightly narrower than widest part of carapace, with broad dentate triangular rostrum and pair of smaller spinose exorbital teeth. Eyestalks and antennae not hidden by rostrum (Fig. 14.5a)����������������������������������������������������������������������������������������Curupironomus Curupironomus Tavares, 1993

Diagnosis. Carapace lateral margins convex, widest at midpoint; front produced, slightly narrower than widest part of carapace, with broad dentate triangular rostrum and pair of smaller spinose exorbital teeth. Eyestalk almost completely visible, antennae hidden; eyestalk mobile. Maxilliped 3 merus broad; palp articulating mesiodistally. Pereopods 2, 3 robust, ~1.5 times carapace length, dactylus of pereopod 3 shorter than propodus. Pereopods 4, 5 short, of 6 articles. Female pleon of 6 pleonites, plus telson. Maximum cl. 9 mm. Shelf, slope (120–546 m). Tropical Atlantic (Caribbean). 1 species (Milne-Edwards and Bouvier 1902: figures; Tavares 1993b).

Cymonomus A. Milne-Edwards, 1880 Diagnosis. Carapace parallel-sided; front transverse, about half as wide as carapace or slightly wider, with short triangular, spiniform or rod-like rostrum and pair of exorbital projections. Eyestalk and antennae not hidden by rostrum; eyestalk fixed to rostrum base or slightly movable. Maxilliped 3 merus elongated, distally produced; palp articulating on mesial margin close to merus midlength. Pereopods 2, 3 thin, much longer than carapace length, dactylus of pereopod 3 longer than propodus. Pereopods 4, 5 short, of 6 articles. Female pleon of 5 segments plus pleotelson, or of 6 segments plus telson. Maximum cl. 11 mm (Pl. 36g). Shelf–bathyal (176–2525 m). Cosmopolitan except polar seas. 47 species (Ahyong 2019: 6 species groups recognised, key to 14 species from New Zealand and Australia; Ahyong et al. 2020; Ahyong and Ng 2009: 4 species described, sexual dimorphism; Ahyong and Ng 2017: key to 7 species from East Asia; Guzmán 2003: record from Chile; Tavares 1991: Brazilian species; Tavares 1994: key to 7 American species). We treat Cymonomoides Tavares, 1993 as a junior synonym following the discussion of Ahyong and Ng (2009).

Cymopolus A. Milne-Edwards, 1880 Diagnosis. Carapace parallel-sided; front produced, laterally concave, widest part of carapace, with broad spinose triangular rostrum

and pair of spinose projections. Eyestalk and antennae almost completely hidden by rostrum, cornea visible; eyestalk mobile. Maxilliped 3 merus broad; palp articulating mesiodistally. Pereopods 2, 3 robust, ~1.5 times carapace length, dactylus of pereopod 3 shorter than propodus. Pereopods 4, 5 short, of 6 articles. Female pleon of 5 segments plus pleotelson. Maximum cl. 8.5 mm. Slope (270 m). Tropical Atlantic (Caribbean). 1 species (MilneEdwards and Bouvier 1902: figures; Tavares 1993b: rediagnosis).

Elassopodus Tavares, 1993 Diagnosis. Carapace lateral margins convex, widest at midpoint; front quarter carapace width, with rostrum and pair of longer lateral projections. Eyestalk and antennae hidden by rostrum; eyestalk fixed to rostrum. Maxilliped 3 merus broad; palp articulating mesiodistally. Pereopods 2, 3 robust, about equal to carapace length, dactylus of pereopod 3 shorter than propodus. Pereopods 4, 5 vestigial, of 1 article. Female pleon of 5 segments plus pleotelson (male), or of 6 segments plus telson (female). Maximum cl. 6.5 mm. Slope (700 m). Central Indo-Pacific. 1 species (Tavares 1993a). References Ahyong ST (2019) The cymonomid crabs of New Zealand and Australia (Crustacea: Brachyura: Cyclodorripoida). Records of the Australian Museum 71, 33–69. doi:10.3853/j.2201-4349.71.2019.1682 Ahyong ST, Ng PKL (2009) The Cymonomidae of the Philippines (Crustacea: Decapoda: Brachyura), with descriptions of four new species. Raffles Bulletin of Zoology Supplement 20, 233–246. Ahyong ST, Ng PKL (2017) East Asian cymonomid crabs (Crustacea: Brachyura). Zoological Studies (Taipei, Taiwan) 56, 1–20. Ahyong ST, Mitra S, Ng PKL (2020) Cymonomid crabs from southwestern Indonesia and redescription of Cymonomus andamanicus Alcock, 1905. Raffles Bulletin of Zoology 68, 62–69. Guinot D, Wicksten MK (2015) Chapter 71–11. Camouflage: carrying behaviour, decoration behaviour, and other modalities of concealment in Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1).

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(Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 583–638. Brill, Leiden. Guzmán GL (2003) Primer registro para Chile y descripción de la hembra de Cymonomus menziesi Garth, 1971 (Decapoda, Archaeobrachyura, Cymonomidae). Investigaciones Marinas 31, 91–94. doi:10.4067/ S0717-71782003000200008 Milne-Edwards A, Bouvier E-L (1902) Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877–78), in the Caribbean Sea (1878–79), and along the Atlantic coast of the United States (1880), by the U. S. Coast Survey steamer “Blake,” Lieut.-Com. C.D. Sigsbee, U.S.N., and Commander J.R. Bartlett, U.S.N., commanding. XXXIX: Les Dromiacés et Oxystomes. Memoirs of the Museum of Comparative Zoology at Harvard College 27, 1–127, pls 1–25. Tavares MS (1991) Espèces nouvelles de Cyclodorippoidea Ortmann et remarques sur les genres Tymolus Stimpson et Cyclodorippe A. Milne Edwards (Crustacea, Decapoda, Brachyura). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 12, 623–648. Tavares MS (1993a) Crustacea Decapoda: Les Cyclodorippidae et Cymonomidae de l’Indo-Ouest Pacifique à l’exclusion du genre Cymonomus. In: Crosnier A (Ed.), Résultats des campagnes MUSORSTOM Vol. 10. Mémoires du Muséum National d’Histoire Naturelle, Paris 156, 253–313. Tavares MS (1993b) Description préliminaire de quatre nouveaux genres et trois nouvelles espèces de Cyclodorippoidea Américains (Crustacea, Decapoda, Brachyura). Vie et Milieu 43, 137–144. Tavares MS (1994) Description de Cymonomus leblondi sp. nov. de Guadeloupe, avec une clef des Cymonomus américains. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série 16, 203–208.

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Phyllotymolinidae Tavares, 1998 Figure 14.6 The last two pairs of ‘walking legs’ of phyllotymolinids are carried over the carapace as in other cyclodorippoids. Phyllotymolinids differ from cymonomids in having movable eyestalks in well developed orbits. Whereas the buccal cavity of cyclodorippids is closed closely by the operculiform maxillipeds 3 and has a curved, bluntly pointed anterior margin, that of phyllotymolinids is more rectangular and lacks anterior definition (Fig. 14.6d) (Tavares 1998). Only four species are known, all from lower shelf depths in the Western Atlantic, eastern and central Indo-Pacific. Tavares’ (1998) key to genera relies on differences between the position of spermathecal openings and the relationship between thoracic sternites 6 and 7. Our key uses more easily seen characters. Diagnosis. Carapace subhexagonal. Orbit developed; eyes retractile. Endostome shallow, at most only slightly projecting anteriorly, giving buccal cavity rectangular appearance. Maxilliped 3 palp fully visible, inserted externally near inner angle of merus, insertion visible; exopod with flagellum.

Key to genera of Phyllotymolinidae 1. – 2. –

Carapace subpentagonal, with 2 tubercles on anterolateral margin (Fig. 14.6b)������������������������������������������������������Genkaia Carapace subrectangular or subpentagonal, lateral margin smooth���������������������������������������������������������������������������������������2 Carapace subpentagonal (Fig. 14.6e). Pereopods 2, 3 propodus and dactylus cylindrical����������������������� Phyllotymolinum Carapace subrectangular, lateral margin smooth, scarcely converging posteriorly. Pereopods 2, 3 propodus and dactylus flattened, with setose margin (Fig. 14.6a)������������������������������������������������������������������������������ Lonchodactylus Genkaia Miyake & Takeda, 1970

Phyllotymolinum Tavares, 1993

Diagnosis. Carapace subpentagonal, with 2 tubercles on anterolateral margin. Pereopods 2, 3 propodus and dactylus cylindrical. Thoracic sternite 7 overlapping sternite 6; spermathecal openings well separated, indicated by distinct boss. Female pleopod 1 vestigial; pleopods 2–5 exopods linear. Maximum cl. 4 mm. Shelf (68–150 m). Temperate Northern W Pacific, Central Indo-Pacific. 2 species (Miyake and Takeda 1970: diagnosis; Tavares 1993: rediagnosis, key to species; Tavares 1998: rediagnosis, key to species).

Diagnosis. Carapace subpentagonal, anterolateral margins smooth. Pereopods 2, 3 propodus and dactylus cylindrical. Thoracic sternites 6, 7 contiguous; spermathecal openings well separated, indicated by distinct boss. Female pleopod 1 absent; pleopods 2–5 exopods oval. Maximum cl. 5.7 mm. Slope (205–307 m). Central Indo-Pacific. 1 species (Tavares 1993).

Lonchodactylus Tavares & Lemaitre, 1996 Diagnosis. Carapace subrectangular, lateral margin smooth, scarcely converging posteriorly. Pereopods 2, 3 propodus and dactylus flattened, with setose margin. Thoracic sternites 6, 7 contiguous; spermathecal openings almost contiguous, not indicated by distinct boss. Female pleopod 1 vestigial; pleopods 2–5 exopods linear. Maximum cl. 6.8 mm. Slope (265 m). Tropical W Atlantic. 1 species (Tavares and Lemaitre 1996).

References Miyake S, Takeda M (1970) A remarkable species of the Dromiacea (Crustacea Decapoda) from the Tsushina Islands, Japan. Ohmu 3, 19–28. Tavares MS (1993) Crustacea Decapoda: Les Cyclodorippidae et Cymonomidae de l’Indo-Ouest Pacifique à l’exclusion du genre Cymonomus. In: Crosnier A (Ed.), Résultats des campagnes MUSORSTOM Vol. 10. Mémoires du Muséum National d’Histoire Naturelle, Paris 156, 253–313. Tavares MS (1998) Phyllotymolinidae, nouvelle famille de Brachyoures Podotremata (Crustacea, Decapoda). Zoosystema 20, 109–122. Tavares MS, Lemaitre R (1996) Lonchodactylus messingi, a new genus and species of Cyclodorippidae (Crustacea: Decapoda: Brachyura) from the Bahamas. Proceedings of the Biological Society of Washington 109, 464–469.

442

Marine Decapod Crustacea

Fig. 14.6.  Phyllotymolinidae. a, Lonchodactylus messingi Tavares & Lemaitre, 1996; b–d, Genkaia keijii Tavares, 1993, with detail of dorsal and ventral front; e, f, Phyllotymolinum crosnieri Tavares, 1993, with detail of dorsal front.

Section Dromiacea De Haan, 1833 Dromiacea include two superfamilies, one with one family and the other with two families. All have one or two posterior pairs of reduced, chelate or subchelate legs. The superfamilies were for many years included as part of Podotremata, the primitive crabs (Ng et  al. 2008). Podotremata have been shown to be paraphyletic (Ahyong et al. 2007) and divided into four sections. Affinities between the

three families of Dromiacea have long been realised (Guinot 2019; Karasawa et al. 2011; Schweitzer et al. 2012). Diagnosis. Carapace wider than to longer than wide, usually about as long as wide. Maxilliped 3 operculiform, or pediform. Pereopods 2–5 propodus and dactylus subcylindrical or weakly compressed, not flat; pereopods 4–5 slender, subcylindrical, pereopod 5 or 4 and 5 subchelate, reduced. Male and female genital openings coxal.

Key to families of Dromiacea 1. – 2. –

Uropod minute, ventral (not visible externally; Fig. 14.12i). Carapace branchiostegite decalcified. Maxilliped 3 pediform (Fig. 14.12h)�������������������������������������������������������������������� Homolodromioidea, Homolodromiidae Uropod absent or visible externally (Fig. 14.10d–t). Carapace branchiostegite calcified. Maxilliped 3 operculiform (Fig. 14.10u, 14.11k) or pediform������������������������������������������������������������������������������������������Dromioidea … 2 Pereopods 2, 3 similar, functioning as walking legs; pereopods 4, 5 smaller than pereopods 2, 3, subchelate, subdorsal, modified for carrying objects (Figs 14.7–14.9)���������������������������������������������������������������������������������� Dromiidae Pereopods 2–4 functioning as walking legs; pereopod 5 markedly smaller than pereopods 2–4. No known carrying behaviour (Fig. 14.11)������������������������������������������������������������������������������������������������������������������������� Dynomenidae

References Ahyong ST, Lai JCY, Sharkey D, Colgan DJ, Ng PKL (2007) Phylogenetics of the brachyuran crabs (Crustacea: Decapoda): the status of Podotremata based on small subunit nuclear ribosomal RNA. Molecular Phylogenetics and Evolution 45, 576–586. doi:10.1016/j.ympev.​2007.​ 03.022

Guinot D (2019) New hypotheses concerning the earliest brachyurans (Crustacea, Decapoda, Brachyura). Geodiversitas 41, 747–796. doi:10.5252/geodiversitas2019v41a22 Karasawa H, Schweitzer CE, Feldmann RM (2011) Phylogenetic analysis and revised classification of podotrematous Brachyura (Decapoda) Including extinct and extant families. Journal of Crustacean Biology 31, 523–565. doi:10.1651/10-3423.1

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Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Schweitzer CE, Feldmann RM, Karasawa H (2012) Part R, Revised, Volume 1, Chapter 8M: Systematic descriptions: Infraorder Brachyura, Section Dromiacea. Treatise Online 51, 1–43. doi:10.17161/to.v0i0.4336

Superfamily Dromioidea De Haan, 1833 Dromioidea include two families with one or two pairs of reduced, chelate or subchelate legs. Both families have uropods, albeit vestigial, almost unique within Brachyura. Diagnosis. Maxilliped 3 operculiform or pediform. Pereopods 2, 3 similar, functioning as walking legs; pereopods 4, 5 smaller than pereopods 2, 3, subchelate, subdorsal, prehensile, or 2–4 functioning as walking legs; pereopod 5 markedly smaller than pereopods 2–4. Uropods absent (rare), or visible externally as small flattened plates.

Dromiidae De Haan, 1833 sponge crabs Figures 14.7–14.10, Plate 37 The dromiids are the sponge crabs, so called because of their habit of disguise using a piece of sponge or other encrusting animal carried over the back. In many species the sponge appears to have been carefully trimmed to fit the crab and is sometimes much larger than its owner. Solitary or colonial ascidians are also used by many species but others use algae (McLay 2020). Sponge crabs do this using the hooked or subchelate dactyli of the last two pairs of legs which are held over the carapace (Guinot and Wicksten 2015). Bivalve mollusc shells are also used by at least three genera, this strategy being apparently convergent as the grasping mechanisms are very different (Guinot and Tavares 2000; McLay 2001c). Most species are behaviourally cryptic and are rarely seen intertidally.

443

McLay’s (1993) systematic review of the family synthesised earlier studies such as those of Borradaile (1903) and Lewinsohn (1977, 1984). McLay addressed considerable nomenclatural confusion while providing several new generic names. While he concentrated on descriptions of New Caledonian and Philippine species, his revision is world-wide at the generic level, recognising 29 genera for ~100 species and providing keys to many. McLay et  al.’s (2001) examination of evidence of interrelationships based on the larvae of 11 species of Dromiidae partially supported a later revision of Dromia (McLay 2009). The systematics of the family was revisited by Guinot and Tavares (2003) who proposed three subfamilies (Dromiinae, Hypoconchinae, Sphaerodromiinae) based largely on the male and female reproductive structures. They also rediagnosed some genera and erected eight new genera. In the only phylogenetic study of dromiids based on molecular and spermatozoal morphology, Garcia Bento et al. (2018) recovered the distinction between Dromiinae and Hypoconchinae, although Sphaerodromiinae were not included. The presence of an epipod attached to the coxa of the cheliped is a character of major taxonomic importance in distinguishing genera but it can be difficult to see. It is concealed inside the gill chamber by the edge of the carapace but is visible on the posterior edge of the coxa if the cheliped is moved forward or detached (Fig. 14.9a). The front of the carapace of dromiids, comprising the rostrum may be undivided, but is more often divided into three lobes, referred to in most taxonomic literature as ‘tridentate’ referring to the median and pair of lateral rostral lobes. The median rostral lobe is often on a lower plane than the lateral rostral lobes and may not always be visible from dorsal view. Sometimes the lateral rostral lobe merges with the orbital eave, including the inner orbital tooth. In most genera pleonites 5 and 6 are freely articulating. Diagnosis. Pereopods 2, 3 similar, functioning as walking legs; pereopods 4, 5 smaller than pereopods 2, 3, subchelate, subdorsal, prehensile.

Key to subfamilies and genera of Dromiidae 1. – 2. – 3. –

Carapace flat, partly membranous, frontal and anterolateral margins expanded, covering eyes (Fig. 14.8b, c). Pereopods 4, 5 dactyli short, set in deep notch in propodus, ending in hooked unguis (Fig. 14.9m). Americas only�����������������������������������������������������������������������������������������������������������������������Hypoconchinae … Hypoconcha Carapace frontal and lateral margins not as above. Pereopods 4, 5 dactyli exposed, not set in notch in propodus (Fig. 14.9h–l)���������������������������������������������������������������������������������������������������������������������������������������������������������������2 Pereopods 2, 3 propodi with distal spine; dactyli almost straight (Fig. 14.9g). Male pereopod 5 coxa prolonged into hard process enclosing genital papilla (Fig. 14.9r)���������������������������������Sphaerodromiinae … 3 Pereopods 2, 3 propodi without distal spine; dactyli usually strongly curved (Fig. 14.9f). Male pereopod 5 coxa bearing articulated tube enclosing genital papilla (Fig. 14.9q)�����������������������������������Dromiinae … 5 Carapace longer than wide. Median and lateral rostral lobes prominent, narrow, pointed (Fig. 14.7s)������������Frodromia Carapace as long as wide. Median rostral lobe absent, lateral lobes broadly angular�����������������������������������������������������������4

444

4. – 5.

Marine Decapod Crustacea

Carapace dorsal surface and lateral margins smooth (Fig. 14.8q)�������������������������������������������������������������������Sphaerodromia Carapace dorsal surface granulate, lateral margins with row of tubercles (Fig. 14.7m)���������������������������������������� Eodromia Carapace flat, subpentagonal; median rostral lobe short, lateral rostral lobes longer, triangular (Fig. 14.7f). Pereopod 4 dactylus large, talon-like (Fig. 14.9j)��������������������������������������������������������������������������������������������� Conchoecetes – Carapace variously shaped, usually convex; rostral lobes of various forms. Pereopod 4 dactylus not talon-like, usually opposed or surrounded by 1 or more propodal spines������������������������������������������������������������������������6 6. Carapace 1.6 times as wide as long; anterolateral margin with deep reniform cavity lateral to each eye (Fig. 14.7u)�����������������������������������������������������������������������������������������������������������������������������������������������������������������Haledromia – Carapace at most 1.3 times as wide as long; anterolateral margin without pair of cavities�������������������������������������������������7 7. Cheliped with epipod (Fig. 14.9a)��������������������������������������������������������������������������������������������������������������������������������������������������8 – Cheliped without epipod (exceptional species rare)���������������������������������������������������������������������������������������������������������������� 22 8. Uropod visible only as ventral plate or absent, not visible dorsally (Fig. 14.10n, o)���������������������������������������������������������������9 – Uropod visible as dorsal plate (Fig. 14.10h, i, l, m, p, t)�������������������������������������������������������������������������������������������������������������10 9. Carapace as long as wide; lateral rostral lobes separated by deep U-shaped concavity, median lobe scarcely or not visible dorsally; with shallowly concave margin between lateral rostral lobe and inner orbital tooth (Fig. 14.8w). Uropod absent������������������������������������������������������������������������������������������������������������������������ Tunedromia – Carapace wider than long; lateral rostral lobes and anterolateral margin forming continuous, broadly convex rim with shallow median notch and 1 blunt anterolateral tooth on each side (hidden by setal fringe) (Fig. 14.8d, e). Uropod visible as ventral plate (Fig. 14.10n, o)����������������������������������������������������� Lamarckdromia 10. Carapace strongly convex, lateral rostral lobes low, bluntly rounded, anterolateral margin rounded, without teeth or teeth small to well developed���������������������������������������������������������������������������������������������������������������������11 – Carapace flat or weakly convex, lateral rostral lobes and anterolateral teeth always well developed, prominent�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������13 11. Pleonites 5, 6 fused, sometimes suture partially visible. Uropod oriented transversely in female, longitudinally in male (Fig. 14.10h, i)���������������������������������������������������������������������������������������������������������������� Dromidiopsis – Pleonites 5, 6 free. Uropod oriented transversely in female, obliquely in male (Fig. 14.10l, m)���������������������������������������� 12 12. Carapace anterolateral margins smooth, without teeth. Cheliped fixed finger and dactylus regularly toothed, without strong proximally directed molariform tooth at midpoint���������������������������������Hemisphaerodromia – Carapace anterolateral margins with 2 or 3 blunt, prominent teeth. Cheliped fixed finger and dactylus with strong proximally directed molariform tooth at midpoint���������������������������������������������������������������������������� Mclaydromia 13. Carapace clearly wider than long�������������������������������������������������������������������������������������������������������������������������������������������������14 – Carapace longer than or as long as wide�������������������������������������������������������������������������������������������������������������������������������������18 14. Carapace anterolateral margin with 1 rectangular and 1 triangular teeth (Fig. 14.8r)������������������������������� Stebbingdromia – Carapace anterolateral margin with 2–4 triangular teeth (Figs 14.7j, 14.8i, s, v)�����������������������������������������������������������������15 15. Thoracic sternite 4 anterior section parallel-sided with convex tip, longer than wide, episternites truncate (Fig. 14.9u)��������������������������������������������������������������������������������������������������������������������������������������������������������������Sternodromia – Thoracic sternite 4 anterior section tapering or dilated, episternites with rounded tip�����������������������������������������������������16 16. Thoracic sternite 4 anterior section dilated anteriorly, with broad truncated tip (Fig. 14.9t)��������������������������Metadromia – Thoracic sternite 4 anterior section tapering anteriorly, episternites with rounded tip�����������������������������������������������������17 17. Thoracic sternite 4 triangular, anterior and lateral sections poorly differentiated (Fig. 14.9v)������������������ Tumidodromia – Thoracic sternite 4 with anterior and lateral sections separated by deep concavity (Fig. 14.9y)����������������������������� Dromia 18. Carapace subcircular (Fig. 14.8f). Cheliped carpus, propodus upper margins with 2–4 large tubercles (Fig. 14.9b). Sternal grooves 7/8 of mature female ending apart in prominent tubes behind chelipeds (Fig. 14.9z)��������������������������������������������������������������������������������������������������������������������������������������������������������������� Lauridromia – Carapace oval or pentagonal. Cheliped carpus, propodus smooth or nodular. Sternal grooves 7/8 of mature female ending together on medial prominence (Fig. 14.9x, A)������������������������������������������������������������������������������������������19 19. Carapace subpentagonal, widest at anterior third, anterolateral margin transverse-oblique (Fig. 14.8k, t)������������������� 20 – Carapace ovoid, widest at midpoint (Figs 14.7t, 14.8l)�������������������������������������������������������������������������������������������������������������21

14 – Brachyura – crabs

445

20. Carapace anterolateral margin with 3 short broad lobes and sharper lateral tooth (Fig. 14.8k). Chelipeds, pereopods 2, 3 meri–propodus smooth����������������������������������������������������������������������������������������������������������������Paradromia – Carapace anterolateral margin with 3 sharp teeth (Fig. 14.8t). Chelipeds, pereopods 2, 3 meri–propodus strongly tuberculate�������������������������������������������������������������������������������������������������������������������������������������������������Stimdromia 21. Carapace anterolateral margin with 2 or 3 triangular teeth (Fig. 14.8l). Chelipeds, pereopods 2, 3 meri– propodus carinate (Fig. 14.9d)��������������������������������������������������������������������������������������������������������������������������������Petalomera – Carapace anterolateral margin with 4 irregular strong teeth (Fig. 14.7t). Chelipeds, pereopods 2, 3 meri– propodus tuberculate����������������������������������������������������������������������������������������������������������������������������������������������Fultodromia 22. Uropod absent (Fig. 14.10d) or visible as ventral plate, not or scarcely visible dorsally (Fig. 14.10b, e, f, g, j, k, n, o, r, s)��������������������������������������������������������������������������������������������������������������������������������������������� 23 – Uropod clearly visible only as dorsal plate (Fig. 14.10a, c)�������������������������������������������������������������������������������������������������������33 23. Carapace 1.1–1.5 times as long as wide�������������������������������������������������������������������������������������������������������������������������������������� 24

Fig. 14.7.  Dromiidae. a, Alainodromia timorensis McLay, 1998; b, Alcockdromia fallax (Latreille in Milbert, 1812); c, Ascidiophilus; d, Austrodromidia australis (Rathbun, 1923); e, Barnardromia hirsutimana (Kensley & Buxton, 1984); f, Conchoecetes chantyi McLay & Naruse, 2019; g, Cryptodromia longipes McLay, 1993; h, Cryptodromiopsis; i, Desmodromia griffini McLay, 2001; j, Dromia; k, Dromidia; l, Dromidiopsis tridentata Borradaile, 1903; m, Eodromia denticulata McLay, 1993; n, Epigodromia rugosa McLay, 1993; o, Epipedodromia thomsoni (Fulton & Grant, 1902); p, Eudromidia; q, Exodromidia; r, Foredromia; s, Frodromia atypica (Sakai, 1936); t, Fultodromia nodipes (Guérin, 1832); u, Haledromia bicavernosa (Zietz, 1888); v, w, Hemisphaerodromia, dorsal, anterior.

446

Marine Decapod Crustacea

Fig. 14.8.  Dromiidae. a, Homalodromia coppingeri Miers, 1884; b, c, Hypoconcha, dorsal, ventral front; d, e, Lamarckdromia globosa (Lamarck, 1818), dorsal, front; f, Lauridromia; g, Lewindromia unidentata (Rüppell, 1830); h, Mclaydromia; i, Metadromia wilsoni (Fulton & Grant, 1902); j, Moreiradromia; k, Paradromia; l, Petalomera; m, Platydromia; n, Pseudodromia; o, p, Speodromia, dorsal, ventral; q, Sphaerodromia kendalli (Alcock & Anderson, 1894); r, Stebbingdromia; s, Sternodromia; t, Stimdromia; u, Takedromia cristatipes (Sakai, 1969); v, Tumidodromia dormia (Linnaeus, 1763); w, Tunedromia.

– Carapace as wide as or wider than long to little longer than wide���������������������������������������������������������������������������������������� 25 24. Carapace branchial groove indistinct, margin with or without notch but not strongly constricted; rostrum produced strongly forwards, well in advance of eyes (Fig. 14.7p). Pereopods 4, 5 propodi with movable spine opposing dactylus������������������������������������������������������������������������������������������������������������������������������������������Eudromidia – Carapace branchial groove deep giving carapace waisted appearance (Figs 14.7c, 14.8n); rostrum produced forwards little in advance of eyes (Fig. 14.7c, 14.8n). Pereopods 4, 5 propodi without spiniform setae opposing dactylus (Fig. 14.9h)������������������������������������������������������������������������������������������������������������������������������������������������ 25 25. Rostrum broadly triangular, undivided lateral lobes absent (Fig. 14.7c)����������������������������������������������������������� Ascidiophilus – Rostrum tridentate, median lobe usually visible in dorsal view (Fig. 14.8n)�������������������������������������������������� Pseudodromia 26. Carapace widest anteriorly, flat, with prominent anterolateral tooth. Lateral rostral lobes broad with rounded corners, shelf-like, depressed (Fig. 14.7o)���������������������������������������������������������������������������������������Epipedodromia – Carapace widest near midpoint, convex. Lateral rostral lobes triangular or rounded������������������������������������������������������ 27 27. Carapace 1.3–4 times as wide as long, lateral margin bulging, anterior portion protruding, gastric and branchial regions swollen, with deep sub-branchial cavity (Fig. 14.8o, p)�����������������������������������������������������Speodromia – Carapace as long or longer than wide as, gastric and branchial regions not swollen, without deep subbranchial cavity������������������������������������������������������������������������������������������������������������������������������������������������������������������ 28 28. Carapace dorsally sculptured; anterolateral margin with prominent uneven teeth (Fig. 14.7a, e)���������������������������������� 29 – Carapace dorsally smooth; anterolateral margin smooth or with regular teeth����������������������������������������������������������������� 30

14 – Brachyura – crabs

447

Fig. 14.9.  Dromiidae. a, cheliped coxa with podobranch and epipod (shaded). Cheliped: b, Lauridromia; c, Mclaydromia; d, Petalomera (male, major); d, Stebbingdromia. Pereopod 2, propodus, dactylus: f, Exodromidia; g, Sphaerodromia. Pereopod 4 or 5: h, Ascidiophilus (P5); i, Barnardromia (P4); j, Conchoecetes (P4); k, Dromidiopsis (P5); l, Hemisphaerodromia (P5); m, Hypoconcha (P5); n, Pseudodromia (P4); o, Speodromia (P4). Pereopod 5 coxa, genital papilla: p, Hypoconcha; q, Metadromia; r, Sphaerodromia. Male thoracic sternum, coxa 1–5, genital papilla: s, Dromidia. Thoracic sternite 4, (coxae 1, 2): t, Metadromia (male); u, Sternodromia (female); v, Tumidodromia (female). Female thoracic sternum: w, Cryptodromia; x, Cryptodromiopsis; y, Dromia; z, Lauridromia; A, Moreiradromia. Thoracic somites 4–8, coxa 1–4 (c1–c4), episternites (e) labelled.

29.

Carapace lateral margin with well developed teeth, some bifid, on higher level than orbit. Lateral rostral lobes narrow (Fig. 14.7a)������������������������������������������������������������������������������������������������������������������������������������� Alainodromia – Carapace lateral margin dominated by branchial projection, with 1 or 2 marginal teeth. Lateral rostral lobes rounded, triangular (Fig. 14.7e)�������������������������������������������������������������������������������������������������������������� Barnardromia 30. Southern Australia. Carapace anterolateral margin evenly convex, with or without teeth (Fig. 14.7d). Telson with rounded apex (Fig. 14.10e)���������������������������������������������������������������������������������������������������������Austrodromidia – Southern Africa. Carapace anterolateral margin straight, convex or concave, with or without teeth (Figs 14.7p, q, 14.8m). Telson with terminal spine (Fig. 14.10b, k, r, s)�����������������������������������������������������������������������������31 31. Carapace about as wide as long. Lateral rostral lobes triangular or spike-like (Fig. 14.7q)�����������������������������Exodromidia – Carapace wider than long. Lateral rostral lobes acute or obsolete�����������������������������������������������������������������������������������������32 32. Carapace front, orbit and anterolateral margins convex, without teeth (Fig. 14.8m); dorsal tomentum spongiose. Pereopod 2 coxa without spine����������������������������������������������������������������������������������������������������������Platydromia – Carapace with short median rostral lobe, lateral rostral lobes triangular or spike-like; anterolateral margins almost transverse, with or without teeth; dorsal tomentum short, stiff (Fig. 14.7k). Pereopod 2 coxa with posteriorly directed spine (Fig. 14.9s)�����������������������������������������������������������������������������������������������������Dromidia 33. Lateral rostral lobes merged with inner orbital angle forming broad, quadrate, prominently protruding lobe��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 34 – Lateral rostral lobes triangular, pointed or bluntly angular, separate from inner orbital angle, not usually prominently protruding���������������������������������������������������������������������������������������������������������������������������������������������������������� 36

448

Marine Decapod Crustacea

Fig. 14.10.  Dromiidae. Male posterior pleonites, telson, coxae 1, 2: a, Cryptodromiopsis; b, Dromidia; c, Moreiradromia. Male posterior somites, telson (dorsal unless noted): d, Alainodromia; e, Austrodromidia (ventral); f, g, Dromidia (dorsal, ventral); h, i, Dromidiopsis (female, male); j, Eudromidia (female); k, Exodromidia; l, m, Hemisphaerodromia (female, male); n, o, Lamarckdromia (dorsal, ventral); p, Mclaydromia; q, Moreiradromia (ventral); r, s, Platydromia (dorsal, ventral); t, Stebbingdromia. u, maxilliped 3, Cryptodromia. Pleonites 5, 6, coxa 2 labelled.

34. Carapace lateral rostral lobes with anterior margin oblique or rounded, with mesial basal lobe; anterolateral margin with 2 blunt teeth, separate lateral tooth, with subhepatic tubercles (Fig. 14.7r)������������������������������������������������������������������������������������������������������������������������������������������������������������������ Foredromia – Carapace front with 2 prominent lobes separated by deep U-shaped concavity, each lobe composed of acute lateral rostral lobe and acute inner orbital tooth separated by concave margin, altogether giving front 4-pointed appearance; anterolateral margin smooth, without subhepatic tubercles�����������������������������������������������������35 35. Carapace postorbital tooth present (Fig. 14.8a)��������������������������������������������������������������������������������������������������Homalodromia – Carapace postorbital tooth absent (Fig. 14.8g)������������������������������������������������������������������������������������������������������ Lewindromia 36. Carapace subpentagonal, anterolateral margin with 1 broad, eave-like tooth separated from orbit by deep notch, second tooth less prominent (Fig. 14.7i)�������������������������������������������������������������������������������������Desmodromia – Carapace ovoid, subrectangular or subpentagonal; anterolateral margin granulate or with at least 2 teeth������������������37 37. Carapace lateral margin convex, with 4 or more irregular granulate teeth (Figs 14.7n, 14.8u), without distinct separation between anterolateral and posterolateral margins��������������������������������������������������������������������������� 38 – Carapace with distinct anterolateral and posterolateral margins; anterolateral margin with simple teeth (Figs 14.7b, g, h, 14.8j)�������������������������������������������������������������������������������������������������������������������������������������������������������������� 39 38. Lateral rostral lobes truncate, median rostral lobe low, blunt (Fig. 14.8u)���������������������������������������������������������� Takedromia – Lateral rostral lobes triangular, median rostral lobe pointed (Fig. 14.7n)��������������������������������������������������������� Epigodromia 39. Female sternal sutures 7/8 ending on median tubercle (Fig. 14.9A)�������������������������������������������������������������������������������������� 40 – Female sternal sutures 7/8 ending apart on prominent tubercles (Fig. 14.9w)���������������������������������������������������������������������41 40. Carapace about as wide as long, tomentose except for pair of well defined posterolateral naked areas (Fig. 14.7h). Male pleon held by serrate process on pereopod 2 coxa (Fig. 14.10a)������������������������������Cryptodromiopsis – Carapace wider than long, uniformly tomentose (Fig. 14.8j). Male pleon held by spine on pereopod 2 coxa (Fig. 14.10c)������������������������������������������������������������������������������������������������������������������������������������������������Moreiradromia 41. Carapace sculptured with deep grooves, unevenly pubescent, shaggy median rostral lobe longer than lateral lobes, lateral lobes expanded upturned, forming erect parapet-like rim (Fig. 14.7b)������������������� Alcockdromia

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Carapace evenly convex, evenly tomentose; median rostral lobe similar to or smaller than lateral lobes, pointed or bluntly subconical (Fig. 14.7g)����������������������������������������������������������������������������������������������������������������� 42 42. Maxilliped 3 merus with 1 and carapace subhepatic area with 2 large, smooth, hemispherical tubercles. Female thoracic sternal grooves ending between bases of chelipeds�������������������������������������������������������������Baccadromia – Maxilliped 3 merus with carapace subhepatic area without large, smooth, hemispherical tubercles. Female thoracic sternal grooves ending between bases of pereopod 2 (Fig. 14.9w)������������������������������������������������Cryptodromia

Subfamily Dromiinae De Haan, 1833 Dromiinae contains most genera of the family. All have an articulated tube derived from the coxa of pereopod 5 housing the male genital papilla (Guinot and Tavares 2003). Diagnosis. Pereopods 2, 3 propodi without distal movable spines. Pereopods 4, 5 dactyli usually strongly curved. Male pereopod 5 coxa bearing articulated tube enclosing genital papilla. Alainodromia McLay, 1998 Diagnosis. Carapace as wide as long, subpentagonal, strongly convex, with row of prominent gastric tubercles; rostrum median lobe shorter than triangular lateral lobes; lateral margins with prominent teeth, some bifid, on higher level than orbit. Uropod absent. Cheliped without epipod. Maximum cl. 12 mm. Shelf (27–44 m). Central Indo-Pacific. 1 species (McLay 1998).

Alcockdromia McLay, 2019 Diagnosis. Carapace wider than long, sculptured with deep grooves, unevenly pubescent, shaggy; rostrum median lobe longer than lateral lobes, lateral lobes expanded upturned, forming erect parapet-like rim; anterolateral margin and orbit elevated, with shallow groove behind, with 1 or 2 blunt teeth. Thoracic sternite groove 7/8 of mature female ending apart on prominent tubercle between pereopods 2 or 3. Uropod visible as dorsal plate. Cheliped without epipod. Maximum cl. 13.5 mm. Intertidal, shallow subtidal (camouflaged by sheets of algae). Western Indo-Pacific. 1 species (McLay 2020).

Ascidiophilus Richters, 1880 Diagnosis. Carapace ~1.5 times as long as wide, smooth, with distinct branchial grooves; rostrum broadly triangular, undivided, lateral lobes absent, little advanced beyond eyes; anterolateral margin smooth. Uropod absent. Cheliped without epipod. Pereopods 4, 5 propodi without movable spine opposing dactylus. Maximum cl. 11.5 mm. Subtidal, shelf (1–60 m). Western Indo-Pacific. 1 species (Lewinsohn 1977: description as Pseudodromia caphyraeformis (Richters, 1880); McLay 1993: rediagnosis).

Austrodromidia McLay, 1993 Diagnosis. Carapace about as wide as long or wider than long, tomentose; rostrum median lobe shorter than triangular lateral

lobes; anterolateral margin smooth, or with 4 or 5 teeth. Telson rounded, or tapering to rounded apex. Uropod vestigial on ventral face, or visible as ventral plate. Cheliped without epipod. Maximum cl. 35 mm (Pl. 37a). Intertidal–shelf (0–218 m). Temperate Australasia. 4 species (Guinot and Tavares 2003 rediagnosis; McLay 1993: diagnosis, key to species; Poore 2004: figures, key to species).

Baccadromia McLay & Hosie, 2022 Diagnosis. Carapace about as wide as long, subpentagonal, evenly convex, coarsely tomentose; rostrum median lobe as long as but narrower than lateral lobes, pointed, subconical; anterolateral margin with 2 prominent teeth and few smaller tubercles. Thoracic sternite groove 7/8 of mature female ending slightly apart on elevated platform between base of chelipeds. Male telson wider than long. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 8 mm. Shelf (30–60 m). Western and Central Indo Pacific. 1 species (McLay and Hosie 2022: diagnosis and figures).

Barnardromia McLay, 1993 Diagnosis. Carapace wider than long, gastric and branchial regions inflated; rostrum median lobe short, triangular, lateral lobes prominent, rounded, well exceeding median lobe; lateral margins dominated by prominent branchial tooth, with 1 or 2 marginal teeth. Uropod visible as ventral plate. Cheliped without epipod. Maximum cl. 12 mm. Shelf (27–33 m). Temperate Southern Africa. 2 species (Barnard 1950: figures as Eudromia bituberculata Stebbing, 1920; Kensley and Buxton 1984: description, figures as Cryptodromia hirsutimana Kensley & Buxton, 1984; McLay 1993: key to species).

Conchoecetes Stimpson, 1858 Diagnosis. Carapace subpentagonal, flat, granular to almost smooth; Rostrum median lobe shorter than triangular lateral lobes; anterolateral margin with teeth. Pereopod 4 dactylus large, talon-like, opposing propodal extension. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 42 mm (Pl. 37b). Subtidal, shelf (2–150 m; camouflage: bivalve shells). Temperate Northern W Pacific, Western and Central Indo-Pacific. 10 species (Guinot and Tavares 2000: illustrations and discussion of pereopods 4, 5; McLay and Naruse 2019: rediagnosis, key to species).

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Cryptodromia Stimpson, 1858 Diagnosis. Carapace about as wide as long or wider than long, evenly convex, evenly tomentose; rostrum median lobe similar to or smaller than lateral lobes, bluntly subconical; anterolateral margin with 1–3 blunt teeth. Thoracic sternite groove 7/8 of mature female ending apart on prominent tubercle between pereopods 2 or 3. Telson wider than long. Uropod visible as dorsal plate. Cheliped with epipod (C. fukuii (Sakai, 1936) only) or without epipod. Maximum cl. 15 mm (Pl. 37c, d). Intertidal–shelf (0–108 m). Indo-West Pacific. 22 species (McLay 2001a: rediagnosis, key to 21 species; McLay 2001b: key to species, some since removed to other genera).

Cryptodromiopsis Borradaile, 1903 Diagnosis. Carapace wider than long, tomentose except for pair of posterolateral defined naked areas; rostrum median lobe small, much shorter than sharp, triangular, lateral lobes; anterolateral margin obliquely transverse, with 2 well spaced teeth, 1 smaller lateral tooth. Thoracic sternite groove 7/8 of mature female ending on median tubercle. Male pleon held by serrate process on pereopod 2 coxa. Uropod visible as dorsal plate. Cheliped without epipod. Maximum cl. 8 mm. Intertidal–shelf (0–122 m). Indo-West Pacific. 1 species (Borradaile 1903: figures; Guinot and Tavares 2003: rediagnosis; McLay 1993: description of only species).

Desmodromia McLay, 2001 Diagnosis. Carapace as wide as long, subpentagonal, flat, smooth or minutely granulate; rostrum median lobe small, slightly shorter than triangular lateral lobes; anterolateral margin with 1 broad, eave-like tooth separated from orbit by deep notch, second tooth less prominent. Uropod visible as dorsal plate. Cheliped without epipod. Maximum cl. 9 mm. Subtidal, shelf (9–81 m; camouflaged with bivalve shell). Central Indo-Pacific (N Australia). 2 species (McLay 2001c).

Dromia Weber, 1795 Diagnosis. Carapace wider than long, transversely ovoid, sculptured; rostrum median and lateral lobes short, triangular; anterolateral margin with teeth. Thoracic sternite 4 anterior section tapering anteriorly, episternites with rounded tip, anterior and lateral sections separated by deep concavity. Pereopod 5 dactylus without spine on outer margin. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 80 mm (Pl. 37e–g). Intertidal–slope (0–360 m). Temperate Northern and Tropical Atlantic, Temperate Southern Africa. 5 species (Forest 1974: key to species; McLay 2009: rediagnosis).

Dromidia Stimpson, 1858 Diagnosis. Carapace about as wide as long or wider than long, tomentose; rostrum median lobe short, lateral lobes spike-like or triangular; anterolateral margin smooth, or with teeth. Telson

with terminal spine. Uropod visible as ventral plate, scarcely visible dorsally. Cheliped without epipod. Maximum cl. 70 mm. Shelf (31–59 m). Temperate Southern Africa. 5 species (Guinot and Tavares 2003: rediagnosis; McLay 1993: key to 6 species includes Platydromia).

Dromidiopsis Borradaile, 1900 Diagnosis. Carapace as long as or longer than wide, strongly convex, smooth; rostrum median not visible in dorsal view, lateral lobes low, narrow, bluntly rounded; anterolateral margin smooth, or with teeth. Pereopod 5 dactylus with spine on outer margin. Pleonites 5, 6 fused, suture sometimes visible. Uropod visible as dorsal plate (oriented transversely in female, longitudinally in male). Cheliped with epipod. Maximum cl. 33 mm (Pl. 37h). Intertidal–shelf (0–62 m). Central Indo-Pacific, Temperate Australasia. 4 species (Guinot and Tavares 2003: rediagnosis; McLay 2001a: key to species, some now excluded).

Epigodromia McLay, 1993 Diagnosis. Carapace longer than wide or wider than long, granulate; rostrum median lobe short, triangular, lateral lobes triangular or bluntly rounded, as long as or longer than median lobe; anterolateral margin granular or with 2 or 3 broad granulated teeth. Uropod visible as dorsal plate. Cheliped with epipod (E. nodosa (Sakai, 1936) only), or without epipod. Maximum cl. 10 mm. Intertidal–slope (0–350 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 10 species (McLay 1993: key to 9 species).

Epipedodromia André, 1932 Diagnosis. Carapace as wide as long, widest along anterior margin, flat, minutely granulate; rostrum without median lobe, lateral lobes broad with rounded corners, shelf-like, depressed; anterolateral margin with angular concavity. Uropod absent. Cheliped without epipod. Maximum cl. 12 mm. Subtidal, shelf (to 60 m). Temperate Australasia (SE Australia). 1 species (Guinot and Tavares 2003: rediagnosis; Poore 2004: figure).

Eudromidia Barnard, 1946 Diagnosis. Carapace longer than wide, tomentose; rostrum strongly produced forwards, well in advance of eyes, median lobe absent, lateral lobes well developed as overhanging rounded or triangular eaves; anterolateral margin with 1 tooth. Telson of males with pointed apex, females rounded, or tapering to rounded apex. Uropod visible as ventral plate. Cheliped without epipod. Pereopods 4, 5 propodi with movable spine opposing dactylus. Maximum cl. 15 mm. Shelf, slope (35–275 m). Temperate Southern Africa. 2 species (Barnard 1950: key to species; McLay 1993: rediagnosis).

Exodromidia Stebbing, 1905 Diagnosis. Carapace about as wide as long, smooth or granulate or spinose; rostrum median lobe scarcely visible in dorsal view, lateral

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lobes spike-like or triangular; anterolateral margin smooth, or with teeth, or with short spines. Telson with terminal spine. Uropod visible as ventral plate. Cheliped without epipod. Maximum cl. 34 mm. Shelf, slope (90–600 m). Temperate Southern Africa. 3 species (Barnard 1950: key to 2 species; Kensley 1977; McLay 1993: rediagnosis).

Foredromia McLay, 2002 Diagnosis. Carapace about as wide as long, granulate; rostrum median lobe not visible in dorsal view, lateral lobes broad lobes, with medial shoulder; anterolateral margin with 2 blunt teeth, separate lateral tooth, with subhepatic tubercle. Uropod visible as dorsal plate. Cheliped without epipod. Maximum cl. 6 mm. Intertidal–shelf (0–38 m). Central Indo-Pacific. 1 species (McLay 2002).

Fultodromia McLay, 1993 Diagnosis. Carapace slightly longer than wide, strongly convex, ovoid, minutely granulate; rostrum median lobe short, lateral lobes prominent, rounded, well exceeding median lobe; anterolateral margin with 4 irregular strong teeth. Cheliped, pereopods 2, 3 tuberculate. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 30 mm. Intertidal–shelf (0–123 m). Central Indo-Pacific, Temperate Australasia (S and NW Australia). 2 species (Guinot and Tavares 2003: rediagnosis; McLay 1993; Poore 2004: figure).

Haledromia McLay, 1993 Diagnosis. Carapace 1.6 times as wide as long, tomentose; rostrum median lobe short, triangular, lateral lobes broad, short; anterolateral margin lateral to eyes with pair of deep reniform cavities, otherwise smooth. Uropod vestigial on ventral face. Cheliped with epipod. Intertidal, shallow subtidal. Temperate Australia. 1 species (McLay 1993; Poore 2004: figure).

Hemisphaerodromia Barnard, 1954 Diagnosis. Carapace wider than long, strongly convex, smooth; rostrum median lobe short. indistinct, scarcely visible dorsally, lateral lobes low, broadly rounded; anterolateral margin smooth. Pereopod 5 dactylus without spine on outer margin. Uropod visible as dorsal plate (oriented transversely in female, obliquely in male). Cheliped with epipod. Pleonites 5, 6 free. Maximum cl. 20 mm. Intertidal–shelf (0–25 m; carries compound ascidians). Western Indo-Pacific (Madagascar). 1 species (Barnard 1954: description; Guinot and Tavares 2003: rediagnosis).

Homalodromia Miers, 1884 Diagnosis. Carapace longer than wide, strongly convex, tomentose; front with 2 prominent lobes separated by deep U-shaped

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concavity, each lobe composed of acute lateral rostral lobe and acute inner orbital tooth separated by concave margin, altogether giving front 4-pointed appearance; median rostral lobe absent; with postorbital tooth; anterolateral margin smooth. Uropod visible as dorsal plate. Cheliped without epipod. Maximum cl. 13 mm. Shelf (24–50 m). Indo-West Pacific. 1 species (Guinot and Tavares 2003: rediagnosis; McLay 1993: figures, rediagnosis).

Lamarckdromia Guinot & Tavares, 2003 Diagnosis. Carapace wider than long, tomentose, with fringe of long plumose setae along anterior and lateral margins; rostrum median lobe short, deflexed, lateral lobes and anterolateral margin forming continuous, broadly convex rim with shallow median notch and 1 blunt anterolateral tooth on each side (hidden by setal fringe). Uropod visible as ventral plate. Cheliped with epipod. Maximum cl. 41 mm. Intertidal–shelf (0–180 m). Temperate Australia. 1 species (Guinot and Tavares 2003; Poore 2004: figures).

Lauridromia McLay, 1993 Diagnosis. Carapace about as wide as long, smooth; rostrum median lobe scarcely visible in dorsal view, lateral lobes triangular, longer than median lobe; anterolateral margin with teeth. Thoracic sternite groove 7/8 of mature female ending apart on prominent tubercle between pereopods 2 or 3. Cheliped carpus, propodus upper margins with 2–4 large tubercles. Pereopod 5 dactylus with spine on outer margin. Pleonites 5, 6 fused, suture sometimes visible. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 57 mm (Pl. 37i, j). Subtidal–slope (7–300 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 2 species (Ihle 1913: description, figures; McLay 1993; McLay et al. 2001: larvae).

Lewindromia Guinot & Tavares, 2003 Diagnosis. Carapace longer than wide, smooth; front with 2 prominent lobes separated by deep U-shaped concavity, each lobe composed of acute lateral rostral lobe and acute inner orbital tooth separated by concave margin, altogether giving front 4-pointed appearance; median rostral lobe short; without postorbital tooth; anterolateral margin smooth. Telson much longer than wide. Uropod visible as dorsal plate. Cheliped without epipod. Maximum cl. 13 mm. Intertidal–slope (0–300 m). Indo-West Pacific. 1 species (­Guinot and Tavares 2003; McLay 1993: desciption, figures as Cryptodromiopsis unidentata Rüppell, 1830).

Mclaydromia Guinot & Tavares, 2003 Diagnosis. Carapace 1.2 times as long as wide, convex, smooth; rostrum median lobe minute but visible dorsally, lateral lobes low, rounded, bluntly obtuse; anterolateral margin with 2 or 3 blunt teeth. Uropod visible as dorsal plate (oblique). Cheliped fixed

452

Marine Decapod Crustacea

finger and dactylus with strong proximally directed molariform tooth at midpoint; with epipod. Pereopod 5 dactylus with or without spine on outer margin. Maximum cl. 13 mm. Subtidal, shelf (14–75 m). Western and Central Indo-Pacific. 2 species (Guinot and Tavares 2003; Lewinsohn 1984: description, figures as Dromidiopsis dubia Lewinsohn, 1984).

Metadromia McLay, 2009 Diagnosis. Carapace wider than long, transversely ovoid, smooth or sculptured; rostrum median lobe short, triangular, lateral lobes broad, bluntly angular; anterolateral margin with teeth. Thoracic sternite 4 anterior section dilated anteriorly, with broad truncated tip. Pereopod 5 dactylus without spine on outer margin. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 45 mm (Pl. 37m). Intertidal–slope (0–520 m; without camouflage as adults). Tropical Atlantic, Indo-West Pacific, Temperate Southern Africa, Temperate Australasia. 1 species – M. wilsoni (Fulton & Grant, 1902) is the most widespread dromiid species (Ahyong et  al. 2009: figures; McLay 2009: diagnosis, description and figures of M. wilsoni).

Moreiradromia Guinot & Tavares, 2003 Diagnosis. Carapace about as wide as long, tomentose; rostrum median lobe short, lateral lobes triangular, sharp, as long as median lobe; anterolateral margin with postorbital tooth, 4 or 5 teeth, second lower. Thoracic sternite groove 7/8 of mature female ending on median tubercle. Male pleon held by spine on pereopod 2 coxa. Uropod visible as dorsal plate. Cheliped without epipod. Maximum cl. 32 mm. Subtidal–slope (1–330 m). Tropical W Atlantic, Eastern IndoPacific. 2 species (Guinot and Tavares 2003; Rathbun 1937: figures; Williams 1984: figures as Dromidia antillensis Stimpson, 1858).

Paradromia Balss, 1921 Diagnosis. Carapace as wide as long, subpentagonal, strongly convex, granulate; rostrum median lobe short, deflexed, lateral lobes barely exceeding median lobe; anterolateral margin with 3 short, broad lobes and sharper lateral tooth. Cheliped, pereopods 2, 3 smooth. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 14 mm. Intertidal–shelf (0–37 m). Temperate Northern W, Central Indo-Pacific. 2 species (Henderson 1888: description, figures; McLay 1993: rediagnosis).

Petalomera Stimpson, 1858 Diagnosis. Carapace slightly longer than wide, strongly convex, ovoid, granulate; rostrum median lobe short,lateral lobes triangular, prominent; anterolateral margin with 2 or 3 blunt teeth. Cheliped, pereopods 2, 3 merus-propodus upper margins carinate. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 23 mm (Pl. 37k).

Subtidal, shelf (7–86 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 4 species (McLay and Ng 2007: rediagnosis, key to species).

Platydromia Brocchi, 1877 Diagnosis. Carapace wider than long, with spongy tomentum; rostrum median lobe short, lateral lobes broad, obtuse; anterolateral margin convex, undulated. Telson with terminal spine. Uropod visible as ventral plate. Cheliped without epipod. Maximum cl. 14 mm. Intertidal–slope (0–300 m). Western Indo-Pacific, Temperate Southern Africa. 1 species (Barnard 1950: figures; Guinot and Tavares 2003: rediagnosis).

Pseudodromia Stimpson, 1858 Diagnosis. Carapace ~1.1–1.5 times as long as wide, tomentose, with distinct branchial grooves; rostrum median lobe short, deflexed, lateral lobes triangular, advanced little beyond eyes; anterolateral margin smooth, unarmed or with 1 or 2 blunt teeth. Uropod visible as ventral plate, or visible as ventral plate, scarcely visible dorsally. Cheliped without epipod. Pereopods 4, 5 propodi without movable spine opposing dactylus. Maximum cl. 36 mm. Intertidal–slope (0–400 m). Western Indo-Pacific, Temperate Southern Africa. 3 species (Guinot and Tavares 2003: rediagnosis; McLay 1993: key to species).

Speodromia Barnard, 1946 Diagnosis. Carapace 1.3–42 times as wide as long, anterior portion protruding, gastric and branchial regions swollen, with deep subbranchial cavity; rostrum median lobe short, deflexed, lateral lobes reduced to low obtuse swellings; lateral margins bulging, gastric and branchial regions swollen, with deep sub-branchial cavity; anterolateral margin with irregular dentate. Uropod visible as ventral plate. Cheliped without epipod. Maximum cl. 40 mm. Shelf–slope (37–1281 m). Temperate Southern Africa. 1 species (Barnard 1950: description, figures; McLay 1993: rediagnosis; Ng 2016: description, figures).

Stebbingdromia Guinot & Tavares, 2003 Diagnosis. Carapace wider than long, tomentose, with fringe of long plumose setae along anterior and lateral margins; rostrum median lobe short, triangular, lateral lobes triangular, longer than median lobe; anterolateral margin with 1 rectangular and 1 triangular teeth. Pereopod 4 dactylus with spine on outer margin. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 7 mm. Subtidal, shelf (15–55 m). Central and Eastern Indo-Pacific, Tropical Eastern Pacific. 1 species (Guinot and Tavares 2003; Lewinsohn 1984: description as Dromidiopsis plumosa Lewinsohn, 1984).

Sternodromia Forest, 1974 Diagnosis. Carapace wider than long, transversely ovoid, tomentose; rostrum median lobe short, lateral lobes triangular;

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anterolateral margin with teeth. Thoracic sternite 4 anterior section parallel-sided with convex tip, longer than wide, episternites truncate. Pereopod 4 dactylus without spine on outer margin. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 58 mm. Subtidal, shelf (12–100 m). Tropical E Atlantic. 1 species (Forest 1974; McLay 2009: differences from Dromia).

Stimdromia McLay, 1993 Diagnosis. Carapace wider than long or as wide as long, subpentagonal, strongly convex, smooth or minutely granulate; rostrum median lobe scarcely visible in dorsal view, lateral lobes triangular; anterolateral margin with row of tubercles. Cheliped, pereopods 2, 3 strongly tuberculate, lobed. Pereopod 4 dactylus without spine on outer margin. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 24 mm. Intertidal–slope (0–220 m). Temperate Northern W Pacific, Central Indo-Pacific, Temperate Australasia. 6 species (McLay 1993: key to 5 species).

Takedromia McLay, 1993 Diagnosis. Carapace wider than long, granulate; rostrum median lobe short, triangular or truncate, lateral lobes spike-like or broad, angular; anterolateral margin with 5 or 6 granulate teeth. Uropod visible as dorsal plate. Cheliped without epipod. Maximum cl. 20 mm. Shelf, slope (48–430 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 5 species (McLay 1993: key to 4 species).

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Subfamily Hypoconchinae Guinot & Tavares, 2003 The subfamily contains one genus restricted to the coasts of North and Latin America (Guinot and Tavares 2003). All species camouflage using bivalve shells (Wicksten 1986). Diagnosis. Carapace flat, partly membranous, frontal and anterolateral margins expanded, covering eyes. Pereopods 2, 3 propodi without distal movable spines. Pereopods 4, 5 dactyli short, set in deep notch in propodus, ending in hooked unguis. Male pereopod 5 coxa bearing articulated tube enclosing genital papilla. Hypoconcha Guérin-Méneville, 1854 Subtidal, shelf (1–110 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. Maximum cl. 23 mm. 6 species (Guinot and Tavares 2000: figures of pereopods; Rathbun 1937: key to species; Williams 1984: rediagnosis, key to 3 species).

Subfamily Sphaerodromiinae Guinot & Tavares, 2003 The subfamily contains three genera from the Western Pacific, differing from other dromiids in the fixed process enclosing the genital papilla (Guinot and Tavares 2003). Diagnosis. Pereopods 2, 3 propodi with distal movable spines. Pereopods 4, 5 dactyli almost straight. Male pereopod 5 coxa prolonged into hard process enclosing genital papilla.

Tumidodromia McLay, 2009

Eodromia McLay, 1993

Diagnosis. Carapace wider than long, transversely ovoid, tomentose; rostrum median lobe short, lateral lobes triangular, longer than median lobe; anterolateral margin with teeth. Thoracic sternite 4 anterior section tapering anteriorly, episternites with rounded tip, anterior and lateral sections poorly differentiated. Pereopod 5 dactylus without spine on outer margin. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 160 mm (Pl. 37n). Subtidal, shelf (8–156 m). Indo-West Pacific. 1 species (McLay 2009).

Diagnosis. Carapace about as wide as long, minutely granulate; rostrum median lobe absent, lateral lobes broadly angular; anterolateral margin with row of tubercles. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 8 mm. Slope (335–350 m). Central Indo-Pacific. 1 species (McLay 1993).

Tunedromia McLay, 1993 Diagnosis. Carapace about as wide as long, tomentose; rostrum lateral lobes separated by deep U-shaped concavity, median lobe scarcely or not visible dorsally; with shallowly concave margin between lateral rostral lobe and inner orbital tooth. Uropod absent. Cheliped with epipod. Maximum cl. 10 mm. Shelf–slope (35–270 m). Temperate Northern W Pacific. 1 species (McLay 1993; Takeda and Miyake 1970: description).

Frodromia McLay, 1993 Diagnosis. Carapace longer than wide, minutely granulate;median and lateral rostral lobes prominent, narrow, pointed;anterolateral margin with row of tubercles. Cheliped with epipod. Maximum cl. 10.6 mm. Shelf, slope (50–437 m). Temperate Northern W Pacific, Central Indo-Pacific. 2 species (McLay 1993).

Sphaerodromia Alcock, 1899 Diagnosis. Carapace about as wide as long, smooth; Rostrum median lobe absent, lateral lobes broadly angular; anterolateral

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margin smooth. Uropod visible as dorsal plate. Cheliped with epipod. Maximum cl. 58 mm (Pl. 37l). Shelf, slope (100–450 m). Temperate Northern W Pacific, Indo-West Pacific. 5 species (McLay 2001b: rediagnosis, key to species). References Ahyong ST, Naruse T, Tan SH, Ng PKL (2009) Part II. Infraorder Brachyura: Sections Dromiacea, Raninoida, Cyclodorippoida. In Crustacean fauna of Taiwan: Brachyuran crabs, volume 1 – Carcinology in Taiwan and Dromiacea, Raninoida, Cyclodorippoida. (Eds Chan T-Y, Ng PKL, Ahyong ST, Tan SH) pp. 27–198. National Taiwan Ocean University, Keelung. Barnard KH (1950) Descriptive catalogue of South African decapod Crustacea (crabs and shrimps). Annals of the South African Museum 38, 1–837. Barnard KH (1954) Notes sur une collection de crustacés décapodes de la region Malagache. Mémoires de l’Institut Scientifique de Madagascar, Sér. A 9, 95–104. Borradaile LA (1903) Marine crustaceans. IX. The sponge crabs (Dromiacea). In The Fauna and Geography of the Maldive and Laccadive Archipelagoes; Being the Account of the Work carried on and of the Collections made by an Expedition during the years 1899 and 1900. (Ed. Gardiner JS) pp. 574–578, pl. 23. University Press, Cambridge, England. Forest J (1974) Les Dromies de l’Atlantique oriental. Description de Sternodromia gen. nov. et de deux espèces nouvelles du genre Dromia Weber (Crustacea Decapoda Dromiidae). Annales de l’Institut Océanographique 50, 71–123. Garcia Bento MA, Miranda I, Mantelatto FL, Zara FJ (2018) Comparative spermatozoal ultrastructure and molecular analysis in dromiid crabs and their phylogenetic implications for Dromiidae and Podotremata (Decapoda: Brachyura). Arthropod Structure & Development 47, 627–642. doi:10.1016/j.asd.2018.10.001 Guinot D, Tavares M (2000) Conchoeodromia alcocki Chopra, 1934: megalopa of Conchoecetes artificiosus (Fabricius, 1798) (Decapoda, Brachyura, Dromiidae). Journal of Crustacean Biology 20, 301–309. doi:10.1163/1937240X-90000032 Guinot D, Tavares M (2003) A new subfamilial arrangement for the Dromiidae De Haan, 1833, with diagnoses and descriptions of new genera and species (Crustacea, Decapoda, Brachyura). Zoosystema 25, 43–129. Guinot D, Wicksten MK (2015) Chapter 71–11. Camouflage: carrying behaviour, decoration behaviour, and other modalities of concealment in Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 583–638. Brill, Leiden. Henderson JR (1888) Zoology Part LXIX. Report on the Anomura collected by H.M.S. Challenger during the years 1873–76. Report of the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873–76 27, i–xi, 1–221, 21 pls. Ihle JEW (1913) Die Decapoda Brachyura der Siboga-Expedition. I. Dromiacea. Siboga-Expéditie 39b, 1–94, pls 1–4. Kensley B (1977) The South African Museum’s Meiring Naude Cruises. Part 2. Crustacea, Decapoda, Anomura and Brachyura. Annals of the South African Museum 72, 161–187. Kensley B, Buxton CD (1984) Inshore small mesh trawling survey of the Cape south coast. Part 5. Crustacea, Stomatopoda, Isopoda and Decapoda. South African Journal of Zoology 19, 189–193. doi:10.1080/02541858.​ 1984.11447881 Lewinsohn C (1977) Die Dromiidae des Roten Meeres (Crustacea, Decapoda, Brachyura). Zoologische Verhandelingen 151, 1–41, pl. 1.

Lewinsohn C (1984) Dromiidae (Crustacea, Decapoda, Brachyura) from Madagascar and the Seychelles. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 6, 89–129. McLay CL (1993) Crustacea Decapoda: the sponge crabs (Dromiidae) of New Caledonia and the Philippines with a review of the genera. In: Crosnier A (Ed.), Resultats des Campagnes MUSORSTOM, vol. 10. Mémoires du Muséum National d’Histoire Naturelle, Paris 156, 111–251. McLay CL (1998) A new genus and species of dromiid crab (Brachyura, Dromiidae) from the Timor Sea, north-west Australia with records of other species from the China Sea. Zoosystema 20, 339–350. McLay CL (2001a) The Dromiidae of French Polynesia and a new collection of crabs (Crustacea, Decapoda, Brachyura) from the Marquesas lslands. Zoosystema 23, 77–100. McLay CL (2001b) Dynomenidae and Dromiidae (Decapoda, Brachyura) from Guam, Philippine Islands, Tonga and Samoa. Zoosystema 23, 807–856. McLay CL (2001c) A new genus and two new species of unusual dromiid crabs (Brachyura: Dromiidae) from northern Australia. Records of the Australian Museum 53, 1–8. doi:10.3853/j.0067-1975.53.2001.1319 McLay CL (2002) Foredromia rostrata, a new genus and species of sponge crab (Decapoda, Brachyura, Dromiidae) from Southeast Asia. Crustaceana 75, 505–515. doi:10.1163/156854002760095552 McLay CL (2009) New records of crabs (Decapoda: Brachyura) from the New Zealand region, including a new species of Rochinia A. MilneEdwards, 1875 (Majidae), and a revision of the genus Dromia Weber, 1795 (Dromiidae). Zootaxa 2111, 1–66. doi:10.11646/zootaxa.2111.1.1 McLay CL (2020) Rediscovery of the sponge crab Cryptodromia fallax (Latreille in Milbert, 1812) (Decapoda: Brachyura: Dromiidae) at Mauritius, with the description of a new genus and the confirmation of an unusual seaweed-carrying camouflage mode. Journal of Crustacean Biology 40, 82–88. doi:10.1093/jcbiol/ruz085 McLay CL, Hosie AM (2022) The sponge crabs of Western Australia and the Northwest Shelf with descriptions of new genera and species (Crustacea: Brachyura: Dromiidae). Zootaxa 5129, 301–355. doi:10.11646/​ zootaxa McLay CL, Naruse T (2019) Revision of the shell-carrying crab genus Conchoecetes Stimpson, 1858 (Crustacea: Brachyura: Dromiidae). Zootaxa 4706, 1–47. doi:10.11646/zootaxa.4706.1.1 McLay CL, Ng PKL (2007) Revision of the Indo-West Pacific sponge crabs of the genus Petalomera Stimpson, 1858 (Decapoda: Brachyura: Dromiidae). Raffles Bulletin of Zoology 55, 107–120. McLay CL, Lim SSL, Ng PKL (2001) On the first zoea of Lauridromia indica (Gray, 1831), with an appraisal of the generic classification of the Dromiidae (Decapoda: Brachyura) using larval characters. Journal of Crustacean Biology 21, 733–747. doi:10.1163/20021975-99990170 Ng PKL (2016) The taxonomy of Speodromia platyarthrodes (Stebbing, 1905) (Crustacea: Brachyura), an unusual dromiid crab endemic to South Africa. Zootaxa 4111, 261–275. doi:10.11646/zootaxa.4111.3.4 Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Rathbun MJ (1937) The oxystomatous and allied crabs of America. Bulletin of the United States National Museum 166, 1–278, pls 1–86. Takeda M, Miyake S (1970) Crabs from the East China Sea. IV. Gymnopleura, Dromiacea and Oxystomata. Journal of the Faculty of Agriculture, Kyushu University 16, 193–235, pl. 1. Wicksten M (1986) Shell-carrying in Hypoconcha sabulosa (Herbst, 1799) (Decapoda, Brachyura). Crustaceana 50, 319–320. doi:10.1163/​ 156854086X00340 Williams AB (1984) Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C.

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Dynomenidae Ortmann, 1892 Figure 14.11, Plate 36h–k Dynomenids are principally reef-dwellers associated with corals in the Indo-West Pacific but some species occur down to upper slope depths. Only 20 species are known. The family is unknown in the Atlantic. They can be recognised by the broad triangular or convex frontal margin and the reduced last walking leg. This last leg, less than one third the length of the leg in front, is carried alongside the carapace above the other legs and, in males only, ends in a minute claw. The coarse, wiry setation and reef-habitat of many dynomenids make them easily mistaken in the field as pilumnids, but the reduced pereopod 5 makes them immediately recognisable. Unlike dromiids and homolodromiids, modern dynomenids do not carry camouflage, although it is likely that stem dynomenids were camouflage carriers like other dromiaceans. Dynomenids are most closely related to the

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dromiids but it is not yet clear whether both groups are reciprocally monophyletic (Guinot 2008; McLay 1999) or whether the dynomenids are derived from within the Dromiidae (e.g. Ahyong et al. 2007; Brösing et al. 2007; Guinot 1994). Dynomenids share several features with the dromiid subfamily Sphaerodromiinae, presumed to be basal in Dromiidae, in the vestigial male pleopods 3–5, short female sternal sutures 7/8, spermathecal apertures that are close to the female gonopores on the pereopod 3 coxae, an exopod on gonopod 2, and the hardened process enclosing most of the genital papilla on pereopod 5 in males (Guinot, 2008). The taxonomy of the family was reviewed thoroughly by McLay (1999) who rediagnosed all genera and redescribed several species. Few other species have been described since. Guinot (2008) divided the family into four subfamilies, one with two genera, the others with one genus each. Diagnosis. Pereopods 2–4 functioning as walking legs; pereopod 5 markedly smaller than pereopods 2–4.

Key to subfamilies and genera of Dynomenidae 1. – 2. –

3.



4. –

Carapace longer than wide or as long as wide; dorsal surface strongly ornamented with spines or tubercles (Fig. 14.11a, f). Male pleon entirely filling sternal depression, fully covering sternite 4. Uropods small (Fig. 14.11g, j)�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Carapace as wide or wider than long; dorsal surface (beneath setae) smooth or weakly ornamented (Fig. 14.11b–e). Male pleon not completely filling entire length of sternal depression, anterior portion of sternite 4 exposed. Uropod large (Fig. 14.11h, i)��������������������������������������������������������������������������������������������������������������������3 Carapace elongate, ovate, densely covered by spines, obscuring areolation, cervical and branchial grooves; lateral margins poorly defined (Fig. 14.11a). Male pleon locked by ornaments on maxillipeds, pereopods 1–3. Uropod narrow (Fig. 14.11g)��������������������������������������������������������������������������Acanthodromiinae … Acanthodromia Carapace subquadrangular, densely covered by coarse granules and tubercles; areolation, cervical and branchial grooves clearly visible; lateral margins subparallel, each with posterolateral tooth (Fig. 14.11f). Male pleon locked granules on pereopods 2, 3 coxae. Uropod moderately developed dorsal plate, occupying about half lateral margin of pleonite 6 (Fig. 14.11j)��������������������������� Paradynomeninae … Paradynomene Carapace ovoid, densely covered by short tomentum, undulating appearance, with transverse troughs; dorsal surface smooth (Fig. 14.11e); anterolateral margin unarmed or with nodular teeth. Male pleon restricted by tubercle on pereopod 2 coxa. Uropod large dorsal plate occupying all lateral margin of pleonite 6 in both sexes (Fig. 14.11i)�����������������������������������������������������������������������Metadynomeninae … Metadynomene Carapace broadly rounded; dorsal surface areolate or not, smooth or granulate, sometimes laterally spinose, sparsely covered with setae; anterolateral margin armed with teeth or granules (Fig. 14.11b–d). Male pleon restricted by tubercle on sternite 5. Uropod dorsal plate never whole length of pleonite 6, at least in males (Fig. 14.11h)������������������������������������������������������������������������������������������������������������������������ Dynomeninae … 4 Carapace areolate, granulate and spinous (especially near margins) under tomentum (Fig. 14.11d)��� Hirsutodynomene Carapace smooth or minutely granulate (Fig. 14.11b, c)�������������������������������������������������������������������������������������������Dynomene

Subfamily Acanthodromiinae Guinot, 2008 Diagnosis. Carapace elongate, ovate; densely covered by spines, obscuring cervical and branchial grooves; lateral

margins poorly defined. Male pleon entirely filling sternal depression, locked by ornaments of maxilliped 3, pereopods 1–3. Uropod narrow.

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Fig. 14.11.  Dynomenidae. a, Acanthodromia margarita (Alcock, 1899); b, c, Dynomene pilumnoides Alcock, 1900; d, Hirsutodynomene vespertilio McLay & Ng, 2005; e, Metadynomene tuamotu Ng & McLay, 2010; f, Paradynomene tuberculata Sakai, 1963. Male pleon, thoracic sternum, right limbs: g, Acanthodromia; h, Dynomene; i, Metadynomene. Male, pleonite 6, uropod, telson: j, Paradynomene. k, maxilliped 3, Dynomene. t, telson; u, uropod; mp3, p1–p4, coxae of maxilliped 3, pereopods 1–4; s4, sternite 4; tubercle on sternite 5 (h) and coxa 2 (i) shaded.

Acanthodromia A. Milne-Edwards, 1880

Dynomene Desmarest, 1823

Maximum cl. 18 mm (Pl. 36h). Shelf, slope (120–540 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 2 species (McLay 1999: key to species).

Diagnosis. Carapace smooth or minutely, sparsely granulate. Maximum cl. 22 mm (Pl. 36i). Intertidal–slope (0–400 m). Tropical Atlantic, Indo-West Pacific, Temperate Australasia. 7 species (McLay 2001: rediagnosis, key to species).

Subfamily Dynomeninae Ortmann, 1892 Diagnosis. Carapace broadly rounded; areolate or not, smooth or granulate, sometimes laterally spinose, sparsely covered with setae; lateral margins with teeth or granules. Male pleon not entirely filling sternal depression, restricted by tubercle on sternite 5, anterior part of sternite 4 exposed. Uropod dorsal plates never filling all length of pleonite 6, at least in males.

Hirsutodynomene McLay, 1999 Diagnosis. Carapace areolate, granulate and spinous (especially near margins) under tomentum. Maximum cl. 29 mm (Pl. 36j). Intertidal–shelf (0–125 m). Indo-West Pacific, Tropical Eastern Pacific, Temperate Australasia. 3 species (McLay 1999: rediagnosis, key to species; McLay and Ng 2005).

14 – Brachyura – crabs

Subfamily Metadynomeninae Guinot, 2008 Diagnosis. Carapace ovoid; densely covered by short tomentum, undulating with transverse troughs; dorsal surface smooth; lateral margins subparallel, each with posterolateral tooth. Male pleon not entirely filling sternal depression, restricted by tubercle on pereopod 2 coxa, anterior part of sternite 4 exposed. Uropod large dorsal plates occupying all of lateral margin of pleonite 6 in both sexes. Metadynomene McLay, 1999 Maximum cl. 27 mm. Slope (205–520 m). Temperate Northern W Pacific, Indo-West Pacific, Temperate Australasia. 4 species (McLay 1999; Ng and McLay 2010: rediagnosis, key to species).

Subfamily Paradynomeninae Guinot, 2008 Diagnosis. Carapace longitudinally ovate–subquadrangular; areolate, cervical and branchial grooves clearly visible; lateral margins subparallel, each with posterolateral tooth. Male pleon entirely filling sternal depression, restricted by granules on pereopods 2, 3. Uropod moderately developed dorsal plates, occupying about half length of lateral margin of pleonite 6. Paradynomene Sakai, 1963 Maximum cl. 23 mm (Pl. 36k). Subtidal–slope (1–402 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 6 species (McLay and Ng 2004: rediagnosis, key to species). References Ahyong ST, Lai JCY, Sharkey D, Colgan DJ, Ng PKL (2007) Phylogenetics of the brachyuran crabs (Crustacea: Decapoda): the status of Podotremata based on small subunit nuclear ribosomal RNA. Molecular Phylogenetics and Evolution 45, 576–586. doi:10.1016/j.ympev.2007.03.022 Brösing A, Richter S, Scholtz G (2007) Phylogenetic analysis of the Brachyura (Crustacea, Decapoda) based on characters of the foregut with establishment of a new taxon. Journal of Zoological Systematics and Evolutionary Research 45, 20–32. doi:10.1111/j.1439-0469.2006.00367.x Guinot D (1994) Decapoda Brachyura. Encyclopaedia Biospeologica 1, 165–179. Guinot D (2008) A re-evaluation of the Dynomenidae Ortmann, 1892 (Crustacea, Decapoda, Brachyura, Podotremata), with the recognition of four subfamilies. Zootaxa 1850, 1–26. doi:10.11646/zootaxa.1850.1.1 McLay CL (1999) Crustacea: Decapoda: Revision of the family Dynomenidae. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM. Vol. 20. Mémoires du Muséum National d’Histoire Naturelle, Paris 180, 427–569. McLay CL (2001) Dynomenidae and Dromiidae (Decapoda, Brachyura) from Guam, Philippine Islands, Tonga and Samoa. Zoosystema 23, 807–856. McLay CL, Ng PKL (2004) A taxonomic revision of the genus Paradynomene Sakai, 1963 (Crustacea: Decapoda: Brachyura: Dynomenidae). Zootaxa 657, 1–24. doi:10.11646/zootaxa.657.1.1

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McLay CL, Ng PKL (2005) On a collection of Dromiidae and Dynomenidae from the Philippines, with description of a new species of Hirsutodynomene McLay, 1999 (Crustacea: Decapoda: Brachyura). Zootaxa 1029, 1–30. doi:10.11646/zootaxa.1029.1.1 Ng PKL, McLay CL (2010) Metadynomene tuamotu, a new species of dynomenid crab from French Polynesia (Crustacea: Decapoda: Brachyura). Zootaxa 2405, 48–54. doi:10.11646/zootaxa.2405.1.2

Homolodromioidea Alcock, 1899 Homolodromioids are deep-water carrying crabs. The last two pair of legs are held aloft, each with a pincer formed from the dactylus and mobile spines at the end of the propodus. Homolodromioids have a rather thin carapace and have been observed carrying and completely hidden by a sponge (Ahyong et al. 2009; Ho and Ng 1999) or by a glass sponge (Guinot and Wicksten 2015; Ng and Naruse 2007). Homolodromiids are widely held to represent the ‘oldest’ lineage of living crabs. The modest size of modern homolodromoid fauna belies a much more diverse and extensive radiation that peaked in the Late Jurassic with six families and 30 genera (Karasawa et al. 2011). Today, only two genera in a single family remain. The systematics of Homolodromioidea was reviewed in detail by Guinot (1995). She described or redescribed 20 species; six have been described since. Diagnosis. Carapace longer than wide; dorsal surface variously setose; branchiostegite decalcified. Orbit illdefined. Maxilliped 3 pediform. Cheliped dactylus closing into cleft tip of fixed finger (Fig. 14.12e). Pereopods 4 and 5 reduced, lying obliquely on carapace and directed upwards, subchelate or ‘chelate’. Pleons of both sexes of 7 pleonites, each with distinct pleuron; pleopods 3–5 vestigial in males; uropods minute, ventral (not visible dorsally; Fig. 14.12i).

Homolodromiidae Alcock, 1899 Figure 14.12, Plate 38a, b Dicranodromia A. Milne-Edwards, 1880 Diagnosis. Pereopods 2, 3, dactyli shorter than propodi, slightly curved (Fig.  14.12a). Pereopods 4, 5 subchelate, propodus not extended beyond base of dactylus, with long distal movable spines (Fig. 14.12f). Eyestalk basal article mobile (Fig. 14.12c). Maximum cl. 36 mm (Pl. 38a). Shelf, slope (65–1330 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Western and Central IndoPacific, Tropical Eastern Pacific (Galapagos), Temperate Australasia. 22 species (Ahyong 2008; Guinot 1995: rediagnosis, keys to 8 Indo-West Pacific and 8 Atlantic species; Milne-Edwards and Bouvier 1902: figures; Ng and Naruse 2007; Tavares and Lemaitre 2014).

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Fig. 14.12.  Homolodromiidae. a, Dicranodromia sp.; b, Homolodromia paradoxa A. Milne-Edwards, 1880. Ventral front, eyestalks (antennae not shown): c Dicranodromia; d, Homolodromia. Chela: e, Dicranodromia. Pereopod 5, with detail of chelae/subchela: f, Dicranodromia; g, Homolodromia. h, maxilliped 3, Homolodromia. i, male pleon, ventral, Homolodromia.

Homolodromia A. Milne-Edwards, 1880 Diagnosis. Pereopods 2, 3, dactyli as long as propodi, falcate (Fig.  14.12b). Pereopods 4, 5 chelate, propodus extended beyond base of dactylus, with long distal movable spines (Fig. 14.12g). Eyestalk basal article fixed (Fig. 14.12d). Maximum cl. 43 mm (Pl. 38b). Slope (375–960 m). Tropical Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate South America (Chile). 7 species (Báez and Martin 1989: distribution; Guinot 1995: rediagnosis, key to 4 species; Martin et  al. 2001; MilneEdwards and Bouvier 1902: figures; Padate et  al. 2020: key to 6 species, distribution). References Ahyong ST (2008) Deepwater crabs from seamounts and chemosynthetic habitats off eastern New Zealand (Crustacea: Decapoda: Brachyura). Zootaxa 1708, 1–72. doi:10.11646/zootaxa.1708.1.1 Ahyong ST, Naruse T, Tan SH, Ng PKL (2009) Part II. Infraorder Brachyura: Sections Dromiacea, Raninoida, Cyclodorippoida. In Crustacean fauna of Taiwan: Brachyuran crabs, volume 1 – Carcinology in Taiwan and Dromiacea, Raninoida, Cyclodorippoida. (Eds Chan T-Y, Ng PKL, Ahyong ST, Tan SH) pp. 27–198. National Taiwan Ocean University, Keelung. Báez RP, Martin JW (1989) Crabs of the family Homolodromiidae, I. Description of the male of Homolodromia robertsi Garth, 1973, based on specimens from deep waters off the coast of Chile. Journal of Crustacean Biology 9, 492–500. Guinot D (1995) Crustacea Decapoda Brachyura: révision de la famille des Homolodromiidae Alcock, 1900. In: Crosnier A (ed.), Résultats des Campagnes MUSORSTOM, vol. 13. Mémoires du Muséum National d’Histoire Naturelle, Paris 163, 155–282. Guinot D, Wicksten MK (2015) Chapter 71–11. Camouflage: carrying behaviour, decoration behaviour, and other modalities of concealment in Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the

Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 583–638. Brill, Leiden. Ho P-H, Ng PKL (1999) On Homolodromia kai Guinot, 1993 (Decapoda, Brachyura, Homolodromiidae) from the South China Sea. Crustaceana 72, 1123. Karasawa H, Schweitzer CE, Feldmann RM (2011) Phylogenetic analysis and revised classification of podotrematous Brachyura (Decapoda) Including extinct and extant families. Journal of Crustacean Biology 31, 523–565. doi:10.1651/10-3423.1 Martin JW, Christiansen JC, Trautwein SE (2001) Crabs of the family Homolodromiidae, VI. Homolodromia monstrosa new species (Decapoda: Brachyura) from the western North Atlantic with a redescription of the holotype of Homolodromia paradoxa A. Milne Edwards, 1880 and comments on sexual dimorphism. Bulletin of Marine Science 68, 313–326. Milne-Edwards A, Bouvier E-L (1902) Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877–78), in the Caribbean Sea (1878–79), and along the Atlantic coast of the United States (1880), by the U. S. Coast Survey steamer “Blake,” Lieut.-Com. C.D. Sigsbee, U.S.N., and Commander J.R. Bartlett, U.S.N., commanding. XXXIX: Les Dromiacés et Oxystomes. Memoirs of the Museum of Comparative Zoology at Harvard College 27, 1–127, pls 1–25. Ng PKL, Naruse T (2007) On two species of deep-sea homolodromoid crabs of the genus Dicranodromia (Crustacea: Decapoda: Brachyura: Homolodromiidae) from the Philippines, including one new species. Raffles Bulletin of Zoology 16, 47–53. Padate VP, Cubelio SS, Jayachandran KV (2020) Description of a new species of deep-water crab of the genus Homolodromia A. MilneEdwards, 1880 from the northern Indian Ocean (Crustacea: Decapoda: Brachyura: Homolodromiidae). Marine Biology Research doi:10.1080/17451000.2020.1735641. Tavares M, Lemaitre R (2014) New morphological and distributional information on Homolodromiidae and Homolidae (Decapoda: Brachyura) from the Americas, with description of a new species and comments on Western Pacific species. Journal of Crustacean Biology 34, 504–524. doi:10.1163/1937240X-00002243

14 – Brachyura – crabs

Section Homoloida Karasawa, Schweitzer & Feldmann, 2011 Homoloidans combine a pediform maxilliped 3 with long slender walking legs. They are typically deep-water carrier crabs in which pereopods 5 are shorter than the others and end in a small grasping device formed from the dactylus and a variety of modifications of the propodus. Most occur at shelf and slope depths worldwide. A wide range of material derived from other animals is used, sponges, gorgonians, antipatharians, hydroids, algae and sea anemones (Guinot and Wicksten 2015). In situ observations of Paromola cuvieri (Risso, 1816) have shown that carrying behaviour is common and that a wide range of material is used (Braga-Henriques et al. 2012). These authors reviewed published observations and speculated that the behaviour is more for defence and distraction rather than camouflage. Not all homoloids do this. Species of Latreillia have a plumose pereopod 5 that may be used to hold on to soft coral (Poupin et al. 2012). The last leg of the only species of Poupiniidae is unmodified but may be distracting to predators (Guinot 1991).

459

Karasawa et  al. (2011) analysed phylogenetic relationships of homoloidans from a palaeontological perspective. When the paraphyly of Podotremata was first realised, the three families that comprise the superfamily Homoloidea were included in Section Dromiacea (Ahyong et al. 2007; De Grave et al. 2009). They were later revealed to stand apart and elevated to section level (Karasawa et al. 2011). The systematics of the superfamily was discussed by Guinot and Richer de Forges (1995) as a prelude to their review of Homolidae. Alternative keys to families can be found in Guinot and Richer de Forges (1995) and in Williams and Moffitt (1991). These rely in part on the presence or absence of a pair of lineae homolicae, longitudinal grooves running from the orbit to the posterior margin of the carapace (e.g. Fig. 14.15). Diagnosis. Carapace longer than wide. Maxilliped 3 pediform. Pereopods 2–3 or 2–4 similar, as walking legs; pereopod 5 reduced and subchelate, or not reduced but similar to pereopods 2–4. Male and female genital openings coxal.

Homoloidea De Haan, 1839 Key to families of Homoloidea 1. – 2. –

Carapace triangular, with long neck occupying half of carapace length. Rostrum sharp, small. Eyestalk extremely long, basal segment much longer than terminal article, not included in orbit (Fig. 14.15)������ Latreilliidae Carapace more or less rectangular or suboval, without long neck. Rostrum simple or bifid. Eyestalk short or long, basal segment sometimes longer than terminal article, rarely exceeding hepatic spines���������������������������������2 Carapace strongly setose, linea homolica absent. Pereopods 2–5 similar, as walking legs. Pereopod 5 about as long as pereopod 4, dactylus linear, half as long as propodus, not carried dorsally (Fig. 14.16)������������ Poupiniidae Carapace sparsely setose or glabrous, linea homolica present. Pereopods 2–4 similar, as walking legs. Pereopod 5 much shorter than pereopod 4, chelate or subchelate, carried dorsally (Fig. 14.13)������������������Homolidae

References Ahyong ST, Lai JCY, Sharkey D, Colgan DJ, Ng PKL (2007) Phylogenetics of the brachyuran crabs (Crustacea: Decapoda): the status of Podotremata based on small subunit nuclear ribosomal RNA. Molecular Phylogenetics and Evolution 45, 576–586. doi:10.1016/j.ympev.2007.03.022 Braga-Henriques A, Carreiro-Silva M, Tempera F, Porteiro FM, Jakobsen K, et al. (2012) Carrying behavior in the deep-sea crab Paromola cuvieri (Northeast Atlantic). Marine Biodiversity 42, 37–46. doi:10.1007/​ s12526-011-0090-3 De Grave S, Pentcheff ND, Ahyong ST, Chan T-Y, Crandall KA, et  al. (2009) A classification of living and fossil genera of decapod crustaceans. Raffles Bulletin of Zoology Supplement 21, 1–109. Guinot D (1991) Établissement de la famille des Poupiniidae pour Poupinia hirsuta gen. nov., sp. nov. de Polynésie (Crustacea Decapoda Brachyura Homoloidea). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 12, 577–605. Guinot D, Richer de Forges B (1995) Crustacea Decapoda Brachyura: révision de la famille des Homolidae De Haan, 1839. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 13. Mémoires du Muséum National d’Histoire Naturelle, Paris 163, 283–517.

Guinot D, Wicksten MK (2015) Chapter 71–11. Camouflage: carrying behaviour, decoration behaviour, and other modalities of concealment in Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 583–638. Brill, Leiden. Karasawa H, Schweitzer CE, Feldmann RM (2011) Phylogenetic analysis and revised classification of podotrematous Brachyura (Decapoda) Including extinct and extant families. Journal of Crustacean Biology 31, 523–565. doi:10.1651/10-3423.1 Poupin J, Corbari L, Pérez T, Chevaldonné P (2012) Deep-water decapod crustaceans studied with a remotely operated vehicle (ROV) in the Marquesas Islands, French Polynesia (Crustacea: Decapoda). Zootaxa 3550, 43–60. doi:10.11646/zootaxa.3550.1.3 Williams AB, Moffitt RB (1991) Crabs from the Mariana Archipelago: Bothromaia griffini new genus and species (Brachyura; Majidae), and remarks on Poupinia hirsuta Guinot (Homoloidea, Poupiniidae). Proceedings of the Biological Society of Washington 104, 569–582.

460

Marine Decapod Crustacea

Homolidae De Haan, 1839 Figures 14.13, 14.14, Plate 38c–m deep-water carrier crabs, deep-water porter crabs Homolids are much larger carrier crabs than latreillids, the other homoloid family displaying this behaviour. Inanimate objects as well as other invertebrates, such as sea anemones and soft corals may be carried (Ng and Richer de Forges 2017, 2020). The carapace of homolids is more-or-less longitudinally rectangular to ovate, somewhat barrel-shaped. The claw on the end of the last leg takes various forms. Generally, the dactylus folds back on to mobile or fixed spines on the propodus. The apparatus in Homolochunia has been likened to ‘sugar tongs’ (Guinot and Wicksten 2015). The taxonomy of the family was reviewed and many of its species redescribed in detail by Guinot and Richer de Forges (1995). The family has attracted considerable attention since with new records and species throughout the Indo-West Pacific (Ng and Richer de Forges 2017; Richer de Forges and Ng 2007, 2008, 2020). The key is derived from that of Ahyong et  al. (2009) and depends on the relative lengths of pereopods and

armature of the carapace. Homolids bear a pair of pseudorostral spines alongside the rostrum which range in size from minute to as long as the carapace. The lateral margins of the carapace are divided into three regions by the cervical and branchiocardiac grooves: the subhepatic, anterolateral and posterolateral regions. Spination of these regions is key to distinguishing some genera (Fig. 14.14a–n). Diagnosis. Carapace more-or-less rectangular to subovate, barrel-shaped sparsely setose or glabrous; linea homolica present. Eyestalk slender or stout, at most barely overeaching lateral carapace margin. Pereopods 2–4 similar, as walking legs. Pereopod 5 half or more as long as pereopod 4, chelate or subchelate, carried dorsally (Fig. 14.14u-w). Implicit generic attributes. Subhepatic region not swollen. Rostrum simple. Maxilliped 3 pediform, not closing buccal cavity. Male cheliped shorter than pereopods 2–4; cheliped margins without row of long erect fine spines. Pereopods 2–4 long, merus no more than 1.5 times carapace length. Pereopod 5 not thinner than pereopods 2–4; merus similar to that of pereopod 4; merus shorter than carapace plus rostrum; subchelate.

Key to genera of Homolidae 1. – 2. – 3. – 4. – 5. – 6.

Pereopod 5 merus clearly longer than carapace length (including rostrum) (Fig. 14.13a, b)�����������������������������������������������2 Pereopod 5 merus shorter than carapace (including rostrum) (Fig. 14.13c)���������������������������������������������������������������������������4 Pseudorostral spines distally bifid, often with row of dorsal spines; as long as or longer than maximum carapace width (Fig. 14.14e). Pereopod 5 dactylus and propodus forming large pincer; fingers long, slender, with wide gape (Fig. 14.14v)����������������������������������������������������������������������������������������������������������������Homolochunia Pseudorostral spines usually distally unidivided (with or without dorsal spines along margin); shorter than maximum carapace width (Fig. 14.14a, j). Pereopod 5 dactylus and propodus forming small subchela, occluding, without wide gape (Fig. 14.14a, b)�������������������������������������������������������������������������������������������������������������������������3 Carapace hepatic region not swollen; width across hepatic region (excluding spines) narrower than width across branchial regions (excluding spines); without constriction behind hepatic region. Pseudorostral spines with row of dorsal spines (Figs 14.13a, 14.14a)���������������������������������������������������������������������������������������� Dagnaudus Carapace hepatic region swollen, width across hepatic region equal to or greater than width across branchial regions; with distinct constriction behind hepatic region. Pseudorostral spines simple or with row of dorsal spines (Figs 14.13b, 14.14j)������������������������������������������������������������������������������������������������������������� Latreillopsis Rostrum with pair of dorsal spines, directed anterolaterally, forming trident. Carapace with long anterolateral spine at base of cervical groove and long upright median gastric spine (Fig. 14.14f)����������� Homologenus Rostrum simple or shallowly bifid distally (Fig. 14.14b–d). Carapace margin at base of cervical groove unarmed or with spines of similar length to other carapace spines; without long median gastric spine���������������������5 Maxilliped 3 operculiform, almost fully covering buccal cavity (Fig. 14.14q). Carapace hepatic region swollen (Fig. 14.14g)������������������������������������������������������������������������������������������������������������������������������������������Homolomannia Maxilliped 3 pediform or subpediform, not covering buccal cavity (Fig. 14.14o, r). Carapace hepatic region not swollen�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������6 Pseudorostral spines antler-like, with additional dorsal spines; very long, equal to or exceeding distance between bases of hepatic spines. Carapace dorsal spines long, slender (Fig. 14.14h)������������������������������������������� Ihlopsis

14 – Brachyura – crabs

461

Fig. 14.13.  Homolidae. a, Dagnaudus petterdi (Guinot & Richer de Forges, 1995); b, Latreillopsis tetraspinosa Dai & Chen, 1980 c, Paromolopsis boasi Wood-Mason & Alcock, 1891.



Pseudorostral spines not antler-like, simple or with short dorsal spine at midlength; length distinctly less than distance between bases of hepatic spines�����������������������������������������������������������������������������������������������������������������������7 7. Carapace dorsal and lateral surfaces with spinules or spines behind anterolateral or subhepatic spine (Fig. 14.14c, d, i, k)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������8 – Carapace dorsal and lateral surfaces without spinules or spines (except for anterolateral or subhepatic spine and short, conical postcervical tubercle) (Fig. 14.14b, m)���������������������������������������������������������������������������������������������������13 8. Pereopod 5 merus long, reaching anteriorly to level or orbit����������������������������������������������������������������������������������������������������9 – Pereopod 5 merus relatively short, not reaching anteriorly to the level of orbit������������������������������������������������������������������10 9. Carapace surface and margins covered with prominent conical spines. Pseudorostral spines as long as or longer than rostrum (Fig. 14.14k)��������������������������������������������������������������������������������������������������������������������������������� Moloha – Carapace surface and margins without prominent conical spines, at most some small spines. Pseudorostral spines low, much shorter than half rostral length (Fig. 14.14d)���������������������������������������������������������������������������� Homolax 10. Pseudorostral spines well developed, as long as or longer than rostrum (Fig. 14.14l, n)����������������������������������������������������11 – Pseudorostral spines small, conical, much shorter than rostrum (Fig. 14.14c, i)���������������������������������������������������������������� 12 11. Pereopod 5 merus very slender, more than 10 times as long as wide�����������������������������������������������������������������Yaldwynopsis – Pereopod 5 merus relatively stout, ~7 as long as wide�������������������������������������������������������������������������������������������������Paromola 12. Eyes on short stalk, not nearly reaching lateral extremity of carapace (Fig. 14.14i). Outer surface of cheliped palm with dark coloured sunken depression at base of pollex in males and some females (Fig. 14.14s)��������� Lamoha

462

Marine Decapod Crustacea

Fig. 14.14.  Homolidae. Carapace: a, Dagnaudus; b, Gordonopsis; c, Homola; d, Homolax; e, Homolochunia; f, Homologenus; g, Homolomannia; h, Ihlopsis; i, Lamoha; j, Latreillopsis; k, Moloha; l, Paromola; m, Paromolopsis; n, Yaldwynopsis. Maxilliped 3: o, Homola; p, Homolochunia; q, Homolomannia; r, Lamoha. Cheliped: s, Lamoha; t, Yaldwynopsis. Pereopod 5 chela/subchela: u, Gordonopsis; v, Homolochunia; w, Lamoha. Pereopods 4, 5 to same scale: x, y, Lamoha. a–n redrawn from Guinot and Richer de Forges (1995), dorsal spination not always shown; cervical and branchiocardiac grooves shown as light line; lineae homolicae dashed; al, anterolateral spine; mg, mesogastric spine; p, pseudorostral spine; pg, protogastric spine; pl, posterolateral spines; sh, subhepatic spine.

– 13. –

Eyes with long stalk, reaching laterally to or slightly beyond lateral margin of carapace (Fig. 14.14c). Outer surface of cheliped palm without sunken depression at base of pollex������������������������������������������������������������������Homola Carapace widest near midlength. Pseudorostral spines very small, much shorter than rostrum (Fig. 14.14m)����������������������������������������������������������������������������������������������������������������������������������������������������������� Paromolopsis Carapace widest posteriorly. Pseudorostral spines well developed, conical, about as long as rostrum (Fig. 14.14b)��������������������������������������������������������������������������������������������������������������������������������������������������������������Gordonopsis Dagnaudus Guinot & Richer de Forges, 1995

Diagnosis. Dorsal carapace ornamented; anterolateral margin with 1 horizontal spine; posterolateral margin with 3 spines. Pseudorostral spine multifid, with accessory spines in one plane. Eyestalk basal segment short, distal segment and cornea short, globose. Pereopod 5 merus reaching tip of rostrum. Maximum cl. 88 mm.

Shelf, slope (81–920 m). Central Indo-Pacific, Temperate ­ ustralasia. 1 species (Guinot and Richer de Forges 1995: rediagA nosis; Poore 2004: figure).

Gordonopsis Guinot & Richer de Forges, 1995 Diagnosis. Dorsal carapace unarmed except rostrum, pseudorostral, and subhepatic spines, sometimes with conical lateral

14 – Brachyura – crabs

branchial tubercle; anterolateral margin without spine; posterolateral margin smooth; subhepatic region with triangular spine. Pseudorostral spine simple, little shorter than rostrum. Eyestalk basal segment short, distal segment and cornea longer, cylindrical. Pereopods spinose. Pereopod 5 thinner than pereopods 2–4, not reaching end of pereopod 4 merus; almost chelate. Maximum cl. 45 mm (Pl. 38c). Slope (256–1362 m). Western and Central Indo-Pacific. 8 species (Ng and Richer de Forges 2020: rediagnosis, key to species).

463

half carapace width; posterolateral margin smooth or with few small spines. Rostrum long, with pair of dorsolateral spines forming trident. Pseudorostral spine simple, oblique. Eyestalk basal segment short, distal segment and cornea short, cylindrical. Pereopod 5 thinner than pereopods 2–4, not reaching end of pereopod 4 merus. Maximum cl. 19 mm (Pl. 38e). Slope, bathyal (623–2548 m). Temperate Northern Atlantic, Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific, Temperate Australasia. 13 species (Guinot and Richer de Forges 1995: rediagnosis, key to 9 species; Ng 2016: key to 11 species).

Homola Leach, 1816 Diagnosis. Dorsal carapace relatively smooth, with or without tubercles on anterior half; anterolateral margin with 1 oblique spine; posterolateral margin with longitudinal row of spinules. Rostrum bifid. Pseudorostral spine reduced, about as long as supraorbital spine. Eyestalk basal segment thin, distal segment and cornea shorter, cylindrical. Pereopod 5 merus not reaching progastric spine. Maximum cl. 48 mm (Pl. 38d). Shelf, slope (20–1000 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Temperate Australasia. 12 species (Guinot and Richer de Forges 1995: rediagnosis, key to 10 species; Ng and Richer de Forges 2016: new species).

Homolax Alcock, 1899 Diagnosis. Dorsal carapace smooth; anterolateral margin without spine; posterolateral margin with longitudinal row of spinules. Rostrum minutely bifid. Pseudorostral spine reduced, about as long as supraorbital spine. Eyestalk basal segment short, distal segment and cornea short, globose. Pereopod 5 merus reaching orbit. Maximum cl. 57 mm. Slope (250–1068 m). Western Indo-Pacific. 1 species (Guinot and Richer de Forges 1995: rediagnosis).

Homolochunia Doflein, 1904 Diagnosis. Dorsal carapace ornamented; anterolateral margin with 1 oblique spine; posterolateral margin with 1 long and 1 short spine. Pseudorostral spine antler-like, with or without accessory spines, sometimes in different planes. Eyestalk basal segment thin, distal segment and cornea shorter, cylindrical. Maxilliped 3 operculiform, almost completely closing buccal cavity. Pereopods 2–4 extremely long, thin, merus less than to slightly more than twice carapace length. Pereopod 5 merus reaching tip of rostrum; chelate, with subequal fingers. Maximum cl. 38 mm. Shelf, slope (150–1000 m). Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 4 species (Guinot and Richer de Forges 1995: rediagnosis, key to 3 species; Richer de Forges and Ng 2008: 4 species compared).

Homolomannia Ihle, 1912 Diagnosis. Dorsal carapace smooth; anterolateral margin without spine; posterolateral margin smooth; subhepatic region with triangular spine. Rostrum very short, triangular. Pseudorostral spine obsolete. Eyestalk basal segment short, distal segment and cornea short, cylindrical. Maxilliped 3 operculiform, almost completely closing buccal cavity. Maximum cl. 41 mm (Pl. 38f). Shelf, slope (50–1258 m). Western and Central Indo-Pacific. 2 species (Guinot and Richer de Forges 1995: rediagnosis, key to species).

Ihlopsis Guinot & Richer de Forges, 1995 Diagnosis. Dorsal carapace spinose; anterolateral margin with single strong spine as long as half carapace width; posterolateral margin with 2 pairs of spines. Pseudorostral spine longer than width of front but shorter than carapace width, antler-like, accessory spinules in several planes. Eyestalk basal segment thin, distal segment and cornea shorter, cylindrical. Male cheliped reaching end of pereopod 2 merus. Maximum cl. 24 mm (Pl. 38g). Slope (204–650 m). Central Indo-Pacific. 3 species (Guinot and Richer de Forges 1995: rediagnosis, key to 2 species; Naruse and Richer de Forges 2010: third species).

Lamoha Ng, 1998 Diagnosis. Dorsal carapace smooth or anteriorly tuberculate; anterolateral margin smooth or with few small spines; posterolateral margin smooth or with few small spines. Rostrum simple, or bifid. Pseudorostral spine minute, at base of rostrum. Eyestalk basal segment short, distal segment and cornea cylindrical, sitting in wide shallow space at anterior of carapace. Male cheliped with dark spot at base of fixed finger. Pereopod 5 thinner than pereopods 2–4, not reaching end of pereopod 4 merus. Maximum cl. 58 mm (Pl. 38h). Shelf–bathyal (35–2000 m). Tropical Atlantic, Indo-West Pacific, Temperate Australasia. 10 species (Guinot and Richer de Forges 1995: rediagnosis, key to 8 species as Hypsophrys; Richer de Forges and Ng 2020: new species). Lamoha is a replacement name for Hypsophrys (Ng 1998).

Homologenus A. Milne-Edwards in Henderson, 1888

Latreillopsis Henderson, 1888

Diagnosis. Dorsal carapace smooth, with prominent mesogastric spine; anterolateral margin with single strong spine as long as

Diagnosis. Dorsal carapace smooth or granulate; anterolateral margin smooth or with spine; posterolateral margin with broad

464

Marine Decapod Crustacea

horizontal spine; subhepatic region grossly swollen, with 2–4 spines. Pseudorostral spine simple, with 1 or 2 spinules. Eyestalk basal segment thin, distal segment and cornea shorter, cylindrical. Maxilliped 3 operculiform, almost completely closing buccal cavity. Pereopods 2–4 extremely long, thin, merus less than to slightly more than twice carapace length. Pereopod 5 merus reaching tip of rostrum. Maximum cl. 23 mm (Pl. 38i). Shelf, slope (20–500 m). Central and Eastern Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 10 species (Guinot and Richer de Forges 1995: rediagnosis, key to 7 species; Poore 2004: figure; Richer de Forges and Ng 2008: new records, new species).

Moloha Barnard, 1946 Diagnosis. Dorsal carapace spinose, tuberculate; anterolateral margin with 1 oblique spine; posterolateral margin with 1 long and 1 short spine. Pseudorostral spine twice as long as rostrum, longer than front, with lateral accessory spine. Eyestalk basal segment thin, distal segment and cornea shorter, cylindrical. Male cheliped reaching end of pereopod 2 dactylus. Maximum cl. 68 mm (Pl. 38j). Subtidal–slope (15–800 m). Temperate Northern Pacific, IndoWest Pacific, Tropical Eastern Pacific, Temperate Southern Africa. 7 species (Guinot and Richer de Forges 1995: rediagnosis, key to 6 species; Ng and Kumar 2015: new species).

Paromola Wood-Mason & Alcock, 1891 Diagnosis. Dorsal carapace with protogastric spine; anterolateral margin with 1 oblique spine; posterolateral margin spinose or granular. Pseudorostral spine little longer than rostrum, as long or little longer than front, with or without lateral accessory spine. Eyestalk basal segment thin, distal segment and cornea shorter, cylindrical. Maximum cl. 172 mm (Pl. 38k). Shelf, slope (80–1100 m). Temperate Northern and Tropical Atlantic, Indo-West Pacific, Temperate South America. 6 species (Guinot and Richer de Forges 1995: rediagnosis, key to species).

Paromolopsis Wood-Mason & Alcock, 1891 Diagnosis. Dorsal carapace smooth; anterolateral margin without spine; posterolateral margin smooth. Pseudorostral spine reduced, about as long as supraorbital spine. Eyestalk basal segment short, distal segment and cornea short, cylindrical. Maximum cl. 43 mm (Pl. 38l). Slope (284–1124 m). Western and Central Indo-Pacific, Temperate Australasia. 1 species (Guinot and Richer de Forges 1995: rediagnosis).

Yaldwynopsis Guinot & Richer de Forges, 1995 Diagnosis. Dorsal carapace spinose, tuberculate; anterolateral margin with 1 oblique spine; posterolateral margin with rows of spines, first largest. Pseudorostral spine simple, little shorter than

rostrum. Eyestalk basal segment short, distal segment and cornea longer, cylindrical. Male cheliped longer than pereopods 2–4; cheliped margins with row of long erect fine spines. Pereopod 5 thinner than pereopods 2–4, not reaching end of pereopod 4 merus. Maximum cl. 53 mm (Pl. 38m). Shelf, slope (30–293 m). Central and Eastern Indo-Pacific, Temperate Australasia. 4 species (Ahyong et al. 2009: key to 3 species; Guinot and Richer de Forges 1995: diagnosis; Ng and Eldredge 2012: new species; Richer de Forges and Ng 2007: new species). References Ahyong ST, Naruse T, Tan SH, Ng PKL (2009) Part II. Infraorder Brachyura: Sections Dromiacea, Raninoida, Cyclodorippoida. In Crustacean fauna of Taiwan: Brachyuran crabs, volume 1 – Carcinology in Taiwan and Dromiacea, Raninoida, Cyclodorippoida. (Eds Chan T-Y, Ng PKL, Ahyong ST, Tan SH) pp. 27–198. National Taiwan Ocean University, Keelung. Guinot D, Richer de Forges B (1995) Crustacea Decapoda Brachyura: révision de la famille des Homolidae De Haan, 1839. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 13. Mémoires du Muséum National d’Histoire Naturelle, Paris 163, 283–517. Guinot D, Wicksten MK (2015) Chapter 71–11. Camouflage: carrying behaviour, decoration behaviour, and other modalities of concealment in Brachyura. In Treatise on Zoology - Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 583–638. Brill, Leiden. Naruse T, Richer de Forges B (2010) A new species of Ihlopsis Guinot & Richer de Forges, 1995 (Decapoda, Brachyura, Homolidae) from Taiwan. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 221–228. doi:10.1163/ej.9789004170865.i-366.136 Ng PKL (1998) Lamoha, a replacement name for Hypsophrys WoodMason & Alcock, 1891 (Brachyura, Homolidae), a junior homonym of Hypsophrys Agassiz, 1859 (Pisces, Teleostei, Cichlidae). Crustaceana 71, 121–125. doi:10.1163/156854098X00833 Ng PKL (2016) On a new species of deep-sea crab of the genus Homologenus A. Milne-Edwards, in Henderson, 1888 (Crustacea: Decapoda: Brachyura: Homolidae) from Kaua’i, Hawai’i. Proceedings of the Biological Society of Washington 129, 48–55. doi:10.2988/0006-324X-129.Q2.48 Ng PKL, Eldredge LG (2012) A new species of Yaldwynopsis from O’ahu, Hawai’i (Crustacea: Decapoda: Brachyura: Homolidae). Bishop Museum Occasional Papers 112, 29–38. Ng PKL, Kumar AB (2015) The species of Moloha Barnard, 1946, from the western Indian Ocean, with the description of a new species from India (Crustacea: Brachyura: Homolidae). European Journal of Taxonomy 166, 1–25. doi:10.5852/ejt.2015.166 Ng PKL, Richer de Forges B (2016) A new species of Homola Leach, 1816 (Crustacea: Brachyura: Homolidae) from Palau, Western Pacific, with notes on H. mieensis Sakai, 1979. Crustacean Research 45, 1–13. doi:10.18353/crustacea.45.0_1 Ng PKL, Richer de Forges B (2017) On a collection of Homolidae from the South China Sea, with descriptions of two new species of Homologenus A. Milne-Edwards, in Henderson, 1888, and the identities of Homologenus malayensis Ihle, 1912, and Lamoha superciliosa (Wood-Mason, in Wood-Mason & Alcock, 1891). Raffles Bulletin of Zoology 65, 243–268. Ng PKL, Richer de Forges B (2020) A revision of the deep-sea porter crabs of the genus Gordonopsis Guinot & Richer de Forges, 1995 (Crustacea, Decapoda, Brachyura, Homolidae), with descriptions of five new species. Raffles Bulletin of Zoology 68, 267–307.

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Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Richer de Forges B, Ng PKL (2007) New records and new species of Homolidae De Haan, 1839, from the Philippenes and French Polynesia (Crustacea: Decapoda: Brachyura). Raffles Bulletin of Zoology Supplement 16, 29–45. Richer de Forges B, Ng PKL (2008) New western Pacific records of Homolidae De Haan, 1839, with descriptions of new species of Homolochunia Doflein, 1904, and Latreillopsis Henderson, 1888 (Crustacea: Decapoda: Brachyura). Zootaxa 1967, 1–35. doi:10.11646/zootaxa.1967.1.1 Richer de Forges B, Ng PKL (2020) The deep-water Homolidae of Papua New Guinea, (Crustacea: Decapoda: Brachyura), with description of a new species of Lamoha Ng, 1998. In: Corbari L, Chan T-Y and Ahyong ST (Eds) Deep-sea crustaceans from Papua New Guinea. Tropical Deep-Sea Benthos vol. 31. Mémoires du Muséum National d’Histoire Naturelle, Paris 213, 239–257.

Latreilliidae Stimpson, 1858 Figure 14.15, Plate 38n, o Latreilliids are notable for the anterior half of the carapace appearing as a long ‘neck’ and for their exceptionally long thin legs. As in most homoloids the last pairs of legs are carried aloft to carry algae, gorgonians or hydroids. They may wave these legs ‘when in danger’. In species of Latreillia the last pereopod is plumose and not always s­ ubchelate – these species are unlikely to carry camouflage (Castro et  al. 2003; ­Guinot and Wicksten 2015). Latreillia elegans Roux, 1830 has been observed climbing pennatulaceans (Porporato et  al. 2011). Latreilliids superficially resemble some majoids, especially some members of the Inachidae, such as Stenorhynchus. Monophyly of Latreillidae itself is not in doubt. The close relationship between latreilliids and homolids has been corroborated by cladistic analysis of sperm ultrastructure (Jamieson et al. 1995) and somatic morphology (Karasawa et  al. 2011). Molecular phylogenetic analyses, however, go further and place the group within the Homolidae indicating that latreilliids might even represent highly apomorphic homolids (Ahyong et  al. 2007; Tsang et  al. 2014). Castro

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et al. (2003), building on Williams (1982), revised the systematics of the family rediagnosing both genera and redescribing all seven species. The nature of the last pereopod distinguishes the genera. Diagnosis. Carapace triangular, with long neck occupying half of carapace length; linea homolica present. Eyestalk extremely long, basal segment much longer than terminal article, not included in orbit. Pereopod 5 less than one-third length of pereopod 4, subchelate. Eplumula Williams, 1982 Diagnosis. Pereopod 5 smooth, propodus half or less than length of carpus, without conspicuous setae (Fig. 14.15a). Maximum cl. 19 mm (Pl. 38o). Shelf, slope (30–300 m). Temperate Northern W Pacific, Central Indo-Pacific, Temperate Australasia. 2 species (Castro et  al. 2003: rediagnosis, key to species, distributions; Poore 2004: figure; ­Williams 1982).

Latreillia Roux, 1830 Diagnosis. Pereopod 5 propodus more than half length of carpus, with conspicuous setae along full length (Fig. 14.15b). Maximum cl. 16 mm (Pl. 38n). Shelf, slope (22–806 m). Temperate Northern and Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific, Temperate Southern Africa. 5 species (Castro et al. 2003: rediagnosis, key to species, distributions; Guinot and Wicksten 2015). References Ahyong ST, Lai JCY, Sharkey D, Colgan DJ, Ng PKL (2007) Phylogenetics of the brachyuran crabs (Crustacea: Decapoda): the status of Podotremata based on small subunit nuclear ribosomal RNA. Molecular Phylogenetics and Evolution 45, 576–586. doi:10.1016/j.ympev.2007.03.022 Castro P, Williams AB, Cooper LL (2003) Revision of the family Latreilliidae Stimpson, 1858 (Crustacea, Decapoda, Brachyura). Zoosystema 25, 601–634. Guinot D, Wicksten MK (2015) Chapter 71–11. Camouflage: carrying behaviour, decoration behaviour, and other modalities of concealment in

Fig. 14.15.  Latreilliidae. a, Eplumula australiensis (Henderson, 1888); b, Latreillia metanesa Williams, 1982.

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Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 583–638. Brill, Leiden. Jamieson BGM, Guinot D, Richer de Forges B (1995) Phylogeny of the Brachyura: evidence from spermatozoal ulstrastructure (Crustacea, Decapoda). In Advances in spermatozoal phylogeny and taxonomy. (Eds Jamieson BGM, Ausio J, Justine J-L) pp. 265–283. Muséum National d’Histoire Naturelle, Paris. Karasawa H, Schweitzer CE, Feldmann RM (2011) Phylogenetic analysis and revised classification of podotrematous Brachyura (Decapoda) Including extinct and extant families. Journal of Crustacean Biology 31, 523–565. doi:10.1651/10-3423.1 Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Porporato EMD, De Domenico F, Mangano MC, Spanò N (2011) Macropodia longirostris and Latreillia elegans (Decapoda, Brachyura) climbing on Mediterranean Pennatulidae (Anthozoa, Octocorallia): a preliminary note. Crustaceana 84, 1777–1780. doi:10.1163/156854011X612884 Tsang LM, Schubart CD, Ahyong ST, Lai JCY, Au EYC, et al. (2014) Evolutionary history of true crabs (Crustacea: Decapoda: Brachyura) and the origin of freshwater crabs. Molecular Biology and Evolution 31, 1173–1187. doi:10.1093/molbev/msu068 Williams AB (1982) Revision of the genus Latreillia Roux (Brachyura: Homoloidea). Quaderni del Laboratorio di Tecnologia della Pesca 3, 227–255.

Poupiniidae Guinot, 1991 Figure 14.16 The family Poupiniidae was erected for a single species from French Polynesia, Poupinia hirsuta Guinot, 1991, differing from other homoloids in lacking the linea homolica and having similar pereopods 2–5 but with the last pair held aloft as in others in the superfamily (Guinot 1991). Since then, the species has been found elsewhere in the Pacific. Diagnosis. Carapace barrel-shaped, covered in upright setae and small granules; linea homolica absent. Eyestalk short, basal segment shorter than terminal article, enclosed

within orbit. Pereopod 5 about as long as pereopod 4, dactylus linear, half as long as propodus. Poupinia Guinot, 1991 Maximum cl. 48 mm. Slope (366–600 m). Central and Eastern Indo-Pacific. 1 species (Guinot 1991; Poupin 1996: colour figure; Takeda and Suyama 2019: record, figures; Williams and Moffitt 1991: record, figure). References Guinot D (1991) Établissement de la famille des Poupiniidae pour Poupinia hirsuta gen. nov., sp. nov. de Polynésie (Crustacea Decapoda Brachyura Homoloidea). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 12, 577–605. Poupin J (1996) Atlas des Crustacés marins profonds de Polynésie Française Récoltes du navire Marara (1986/1996). Service Mixte de Surveillance Radiologique et Biologique, Montlhéry. Takeda M, Suyama N (2019) Five species of crabs (Crustacea: Decapoda: Brachyura) from the depths off Okino-Torishima, the southernmost Island in Japan. Bulletin of the National Science Museum, Tokyo, Series A (Zoology) 45, 45–59. Williams AB, Moffitt RB (1991) Crabs from the Mariana Archipelago: Bothromaia griffini new genus and species (Brachyura; Majidae), and remarks on Poupinia hirsuta Guinot (Homoloidea, Poupiniidae). Proceedings of the Biological Society of Washington 104, 569–582.

Section Raninoida Ahyong, Lai, Sharkey, Colgan & Ng, 2007 frog crabs, spanner crabs Raninoidans are unusual in having a carapace longer than wide, and the pleon not completely reflexed beneath the carapace as in most other brachyurans. This most uncrablike shape gives them the common name, frog crabs. Frog crabs live buried in sandy sediments on beaches or on the

Fig. 14.16.  Poupiniidae. Poupinia hirsuta Guinot, 1991: a, habitus; b, c, carapace, dorsal, lateral.

14 – Brachyura – crabs

shelf, the ‘walking legs’ often being paddle-like to aid digging. The group has more species in subtropical and tropical waters than in temperate seas (Lam-Gordillo and Ardisson 2019) and is well represented by a diverse fossil fauna (Karasawa et al. 2014; Schweitzer et al. 2018). Phylogenetic analysis of fossil and Recent genera of raninoid and allied superfamilies revealed extensive diversity that peaked in the late Cretaceous and steadily attennuated to the present (Karasawa et al. 2014; Van Bakel et al. 2012). The affinities of raninoidans have attracted considerable debate. The group has traditionally been discussed as a superfamily, Raninoidea De Haan, 1839, comprising one family, Raninidae De Haan, 1839. They are generally assumed to retain several plesiomorphic characters, such as a rostrum and oviduct openings on the coxae of pereopods 3. These features placed them in the Section Podotremata or ‘primitive’ crabs (Guinot et  al. 2013; Ng et  al. 2008). Podotremata, however, have been shown to be paraphyletic (Ahyong et al. 2007) with raninoids deserving of their own section Raninoida (De Grave et  al. 2009). The

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section name Raninoida takes its authorship from authors who first proposed it, not from that of the similarly named family. Guinot (1993) recognised six extant subfamilies of Raninidae s.l. but their status has been debated (Guinot and Ng 2020; Van Bakel et  al. 2012). Here, we treat the section as comprising a single superfamily with two families, Lyreididae and Raninidae, the latter with five subfamilies. The key to all Recent genera given by Dawson and Yaldwyn (2000) has been rearranged here to reflect the current classification. Schweitzer et al. (2018) diagnosed all genera and family-level taxa in detail, including fossils. Dawson and Yaldwyn (1994) listed all species then known; few have been described since. A key to all 19 species of Lyreididae and Raninidae from the Indo-West Pacific known at the time can be found in Goeke (1985). Diagnosis. Carapace longer than wide. Maxilliped 3 operculiform. Pereopods 2–4 or 2–5 propodus and dactylus flattened, often forming paddle, none subchelate. Male and female genital openings coxal.

Raninoidea De Haan, 1839 Key to families of Raninoidea 1. –

Carapace widest behind anterior one-third; front at most half as wide as greatest width; surface smooth. Pleon with median spine or tubercle on either or both pleonites 3, 4 (Fig. 14.17a, b). Thoracic sternum with pleonal locking mechanism (Fig. 14.17d, e)��������������������������������������������������������������������������������������������������Lyreididae Carapace usually widest across anterior one-quarter, usually at or near anterior margin; front usually more than half as wide as greatest width; surface smooth or granular to uniformly scabrous (Figs 14.18, 14.19). Pleon without median spines or tubercles. Thoracic sternum without pleonal locking mechanism����������� Raninidae

References Ahyong ST, Lai JCY, Sharkey D, Colgan DJ, Ng PKL (2007) Phylogenetics of the brachyuran crabs (Crustacea: Decapoda): the status of Podotremata based on small subunit nuclear ribosomal RNA. Molecular Phylogenetics and Evolution 45, 576–586. doi:10.1016/j.ympev.2007.03.022 Dawson EW, Yaldwyn JC (1994) The frog crabs (Crustacea: Brachyura: Raninidae) – a species list and bibliographic index. Occasional Papers of the Hutton Foundation of New Zealand 6, 1–60. Dawson EW, Yaldwyn JC (2000) Description and ecological distribution of a new frog crab (Crustacea, Brachyura, Raninidae) from northern New Zealand waters, with keys to Recent raninid genera and Notosceles species. Tuhinga 11, 47–71. De Grave S, Pentcheff ND, Ahyong ST, Chan T-Y, Crandall KA, et  al. (2009) A classification of living and fossil genera of decapod crustaceans. Raffles Bulletin of Zoology Supplement 21, 1–109. Goeke GD (1985) Decapod Crustacea: Raninidae. In: Résultats des Campagnes MUSORSTOM I et II – Philippines (1976, 1980). Tome 2. Mémoires du Muséum National d’Histoire Naturelle, Paris. Zoologie 133, 207–228. Guinot D (1993) Données nouvelles sur les Raninoidea De Haan, 1841 (Crustacea Decapoda Brachyura Podotremata). Comptes Rendus de l’Académie des Sciences, Paris, Sciences de la vie 316, 1324–1331.

Guinot D, Ng PKL (2020) On the nomenclature of Cyrtorhininae Guinot, 1993 (Brachyura, Raninoidea). Crustaceana 93, 71–76. doi:10.1163/​ 15685403-00003956 Guinot D, Tavares M, Castro P (2013) Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa. Zootaxa 3665, 1–414. doi:10.11646/zootaxa.3665.1.1 Karasawa H, Schweitzer CE, Feldmann RM, Luque J (2014) Phylogeny and classification of Raninoida (Decapoda: Brachyura). Journal of Crustacean Biology 34, 216–272. doi:10.1163/1937240X-00002216 Lam-Gordillo O, Ardisson P-L (2019) The global distribution and richness of frog crabs (Raninoidea). Crustaceana 92, 1–31. doi:10.1163/​ 15685403-00003832 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Schweitzer CE, Feldmann RM, Karasawa H, Luque J (2018) Part R, Revised, Volume 1, Chapter 8S: Systematic Descriptions: Section Raninoida. In Treatise Online. pp. 1–42. KU Paleontological Institute, Lawrence. Van Bakel BWM, Guinot D, Artal P, Fraaije RHB, Jagt JWM (2012) A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata). Zootaxa 3215, 1–216. doi:10.11646/zootaxa.3215.1.1

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Lyreididae Guinot, 1993 frog crabs Figure 14.17, Plate 39a, b The two genera of Lyreididae differ from other raninoidans by the more fusiform carapace, very narrow at the front and more strongly domed aspect. The end of the pleon is locked to the thoracic sternum by a pair of hooks. Guinot’s (1993) subfamily Lyreidinae is recognised at the family level here (Guinot and Ng 2020; Van Bakel et al. 2012). Diagnosis. Carapace widest behind anterior one-third; front at most half as wide as greatest width; surface smooth. Pleon with median spine or tubercle on either or both pleonites 3, 4 (Fig.  14.17a–c). Thoracic sternum with pleonal locking mechanism comprising a pairs of double-notched hooks on thoracic sternite 5 and sockets at the posterolateral margins of pleonite 6 (Fig. 14.17d, e). Pereopod 5 almost filiform (Fig. 14.17a, b). Lyreidus De Haan, 1841 Diagnosis. Rostrum as wide as or wider than long (Fig.  14.17a). Pereopod 4 propodus lower margin with acute point proximally; dactylus half as wide as long (Fig. 14.17f). Maximum cl. 48 mm (Pl. 39a). Shelf–slope (27–512 m). Temperate Northern Pacific, IndoWest Pacific, Temperate Australasia. 3 species (Feldmann 1992: systematics, biogeography, key to species including Lysirude).

Lysirude Goeke, 1985 Diagnosis. Rostrum longer than wide. Pereopod 4 propodus lower margin lobate, rounded; dactylus more than half as wide as long (Fig. 14.17b). Maximum cl. 34 mm (Pl. 39b). Shelf, slope (119–823 m). Temperate Northern and Tropical Atlantic, Indo-West Pacific. 4 species (Takeda et  al. 2021: most recent species).

References Feldmann RM (1992) The genus Lyreidus De Haan, 1839 (Crustacea, Decapoda, Raninidae): systematics and biogeography. Journal of Paleontology 66, 943–957. doi:10.1017/S002233600002103X Guinot D (1993) Données nouvelles sur les Raninoidea De Haan, 1841 (Crustacea Decapoda Brachyura Podotremata). Comptes Rendus de l’Académie des Sciences, Paris, Sciences de la vie 316, 1324–1331. Guinot D, Ng PKL (2020) On the nomenclature of Cyrtorhininae Guinot, 1993 (Brachyura, Raninoidea). Crustaceana 93, 71–76. doi:10.1163/​ 15685403-00003956 Takeda M, Ohtsuchi N, Komatsu H (2021) Crabs (Crustacea, Decapoda) from the sea off East and Southeast Asia collected by the RV Hakuhō Maru (KH-72–1 Cruise) 1. Sulu Sea and Sibutu Passage. Bulletin of the National Museum of Nature and Science. Series A, Zoology 47, 65–97. Van Bakel BWM, Guinot D, Artal P, Fraaije RHB, Jagt JWM (2012) A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata). Zootaxa 3215, 1–216. doi:10.11646/zootaxa.3215.1.1

Raninidae De Haan, 1839 frog crabs, spanner crabs Figures 14.18, 14.19, Plate 39c–k Raninidae is the larger of the two raninoidan families, more widespread in subtropical and tropical waters than in temperate seas (Lam-Gordillo and Ardisson 2019). It is well represented by a diverse fossil fauna (Karasawa et al. 2014; Schweitzer et al. 2018; Van Bakel et al. 2012). The largest raninid, called the spanner crab in Australia, Ranina ranina Linnaeus, 1758, is an edible species widespread in the Indo-West Pacific. This appears to be the only species whose biology has been studied, albeit mostly from a fisheries point of view (Brown et  al. 2008; Thomas et  al. 2013). One of the six extant subfamilies of Raninidae s.l. recognised by Guinot (1993) is here treated as the family Lyreididae (Guinot and Ng 2020; Van Bakel et al. 2012).

Fig. 14.17.  Lyreididae. Habitus: a, Lyreidus tridentatus De Haan, 1841; b, Lysirude griffini Goeke, 1985. c, pleonites 2–6, telson, Lysirude. d, thoracic sternum (sternites 4, 5 numbered), Lyreidus. e, thoracic sternite 5 hook, lateral view, Lyreidus. f, pereopod 4 propodus, dactylus, Lyreidus.

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Diagnosis. Carapace usually widest across anterior onequarter, usually at or near anterior margin; front usually more than half carapace width; surface smooth, granulate, minutely spinulate or uniformly scabrous (Figs 14.18, 14.19). Pleon without median spines or tubercles (Fig.  14.18).

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Thoracic sternum without pleonal locking mechanism. Pereopod 5 flattened. Implicit subfamily attributes. Maxilliped 3 ischium without oblique setose ridge. Pereopods 1 coxae widely separated; pereopods 2, 3 almost or not meeting in midline.

Key to subfamilies and genera of Raninidae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. – 9. – 10. –

Carapace widest across anterior margin; surface uniformly scabrous, shield-shaped, with 2 trifid processes on each anterolateral margin (Fig. 14.18f, Pl. 39g)���������������������������������������������������������������������������� Ranininae … Ranina Carapace widest at or behind anterior margin; surface generally smooth, granulate or minutely spinulate (Fig. 14.18a–e, g–i)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Maxilliped 3 ischium with oblique setose ridge (Fig. 14.19f)��������������������������������������������������������������������� Notopodinae … 3 Maxilliped 3 ischium without oblique ridge (Fig. 14.19g)���������������������������������������������������������������������������������������������������������6 Rostrum absent, front with median V-shaped incision. Carapace narrow anteriorly, dorsal surface with median ridge. Eyestalk elongate, length about half carapace width (Fig. 14.18b)����������������������������������������� Cosmonotus Rostrum present. Carapace widest anteriorly, dorsal surface evenly convex. Eyestalk stout, much less than one-third carapace width (Fig. 14.18c–e)��������������������������������������������������������������������������������������������������������������������������������4 Carapace with transverse spinose ridge between anterolateral spines (Fig. 14.18c)������������������������������������������������� Notopus Carapace without transverse spinose ridge between anterolateral spines (Fig. 14.18d, e)����������������������������������������������������5 Pereopod 4 dactylus acute (Fig. 14.19l)������������������������������������������������������������������������������������������������������������������������������Ranilia Pereopod 4 dactylus truncate (Fig. 14.19p)���������������������������������������������������������������������������������������������������������������������� Umalia Cheliped propodus with spine near base of fixed finger; dactylus with strong proximal spine on upper margin. Pereopods 1, 2 dactyli crescent-shaped; pereopods 3, 4 dactyli sickle-shaped (Fig. 14.18a). Pereopods 1–3 coxae meeting in midline���������������������������������������������������������������������������������������������� Cyrtorhininae … 7 Cheliped propodus without spine near base of fixed finger; dactylus without proximal spine on upper margin. Pereopods 3–5 dactyli sickle-shaped or not. Pereopods 1 coxae widely separated; pereopods 2, 3 almost or not meeting in midline���������������������������������������������������������������������������������������������������������������������������������������������8 Carapace anterolateral margin between exorbital tooth and first anterolateral spine granular, without spines. Eye slender, tapering, cornea subdistal (Fig. 14.19b). Pereopod 5 merus 2.2 times as long as wide, lower margin weakly convex (Fig. 14.19r)������������������������������������������������������������������������������������������������������������� Flaberhina Carapace anterolateral margin between exorbital spine and first anterolateral spine with two sharp spines. Eye stout, cornea distal (Fig. 14.19a). Pereopod 5 merus 2 times as long as wide, lower margin strongly convex (Fig. 14.19q)��������������������������������������������������������������������������������������������������������������������������������������������������� Cyrtorhina Rostrum extending well beyond exorbital spines. Fronto-orbital margin less than half carapace width (Fig. 14.18i). Cheliped swollen, fingers longer than palm (Fig. 14.19l). Pereopods 2–5 dactyli sickleshaped (Fig. 14.19o)��������������������������������������������������������������������������������������������������������������������������� Symethinae … Symethis Rostrum scarcely extending beyond exorbital spines. Fronto-orbital margin at least half carapace width (Fig. 14.18g, h). Cheliped palm flat, fingers about as long as palm. Pereopods 2–4 dactyli crescentshaped (Fig. 14.19n); pereopod 5 dactylus crescent-shaped or ovate�������������������������������������������������� Raninoidinae … 9 Carapace without anterolateral and lateral spine (Fig. 14.19c). Cheliped propodus upper margin without spines or carinae (Fig. 14.19i). Pereopods 2–5 subequal������������������������������������������������������������������������������������� Notopoides Carapace with anterolateral spine, without lateral spine (Fig. 14.18g, h). Cheliped propodus upper margin with spine or carinae (Fig. 14.19j, k). Pereopod 5 much shorter and more slender than pereopods 2–4 (Fig. 14.19m, n)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������10 Cheliped propodus upper margin with spine (Fig. 14.19k). Pleonite 1 narrower than carapace posterior margin (Fig. 14.18h)�������������������������������������������������������������������������������������������������������������������������������������������������� Raninoides Cheliped propodus upper margin with double carinae (Fig. 14.19j). Pleonite 1 as wide as carapace posterior margin (Fig. 14.181g)��������������������������������������������������������������������������������������������������������������������������������������������������Notosceles

470

Marine Decapod Crustacea

Fig. 14.18.  Raninidae. Habitus. Cyrtorhininae: a, Cyrtorhina granulosa Monod, 1956. Notopodinae: b, Cosmonotus grayii Adams in Belcher, 1848; c, Notopus dorsipes (Linnaeus, 1758); d, Ranilia muricata H. Milne Edwards, 1837; e, Umalia orientalis (Sakai, 1963). Ranininae: f, Ranina ranina (Linnaeus, 1758). Raninoidinae: g, Notosceles serratifrons (Henderson, 1893); h, Raninoides intermedius Dai & Xu, 1991. Symethinae: i, Symethis corallica Davie, 1989.

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471

Fig. 14.19.  Raninidae. Front: a, Cyrtorhina; b, Flaberhina; c, Notopoides; d, Notosceles; e, Raninoides. Maxilliped 3: f, Notopus; g, Notosceles. Cheliped, carpus–dactylus: h, Cyrtorhina; i, Notopoides; j, Notosceles; k, Raninoides. Pereopod 4, carpus/propodus–dactylus: l, Ranilia; m, o, Raninoides (with pereopod 5); o, Symethis; p, Umalia. Pereopod 5: q, Cyrtorhina; r, Flaberhina

Subfamily Cyrtorhininae Guinot, 1993 The subfamily was established on the unusual thoracic sternum (Guinot 1993) but its members, rarely collected, are more easily recognised by the spine on the dactylus of the cheliped. Diagnosis. Carapace widest behind anterior margin; surface granulate; fronto-orbital margin with rostrum and pairs of supraorbital and exorbital teeth. Pereopods 1–3 coxae meeting in midline. Cheliped propodus with spine near base of fixed finger; dactylus with strong spine on upper margin. Pereopods 2, 3 dactyli crescent-shaped, pereopods 4, 5 dactyli sickle-shaped. Cyrtorhina Monod, 1956 Diagnosis. Carapace anterolateral margin between exorbital spine and first anterolateral spine with two sharp spines. Eye stout, cornea distal. Pereopod 5 merus 2 times as long as wide, lower margin strongly convex. Penis an almost straight calcified tube projecting from edge of pereopod 5 coxa. Maximum cl. 47 mm. Subtidal (5–12 m). Tropical E Atlantic. 1 species (Monod 1956; Ng and Guinot 2020; Serène 1971: figures, redescription).

Flaberhina Ng & Guinot, 2020 Diagnosis. Carapace anterolateral margin between exorbital tooth and first anterolateral spine granular, without spines. Eye slender,

tapering, cornea subdistal. Pereopod 5 merus 2.2 times as long as wide, lower margin weakly convex. Penis a distinctly curved calcified tube projecting from edge of pereopod 5 coxa. Maximum cl. 52 mm (Pl. 39d). Shelf, slope (20–300 m). Central Indo-Pacific. 1 species (Ng and Guinot 2020; Serène 1971).

Subfamily Notopodinae Serène & Umali, 1972 Diagnosis. Carapace widest behind anterior margin; surface generally granulate or minutely spinulate; fronto-orbital margin without rostrum, or with 1 or 2 fissures each side of rostrum. Maxilliped 3 ischium with oblique setose ridge. Cheliped propodus upper margin smooth or with short teeth. Pereopods 2–5 dactyli broad, spatulate. Cosmonotus White, 1848 Diagnosis. Notopodinae. Carapace narrow anteriorly, V-shaped in cross-section; dorsal surface with median ridge; rostrum absent, front with median V-shaped incision. Eyestalk elongate, length about half carapace width. Pereopod 4 dactylus acute. Maximum cl. 14 mm (Pl. 39c). Shelf, slope (30–369 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 3 species (Tavares 2006: 3 species compared).

472

Marine Decapod Crustacea

Notopus De Haan, 1841 Diagnosis. Notopodinae. Carapace widest anteriorly, dorsal surface evenly convex, with transverse spinose ridge between anterolateral spines; rostrum present. Eyestalk stout, less than one-third carapace width. Pereopod 4 dactylus acute. Maximum cl. 27 mm (Pl. 39e). Intertidal–shelf (0–68 m). Temperate Northern W Pacific, Indo-West Pacific. 1 species (Ahyong et al. 2009: figures).

Ranilia H. Milne Edwards, 1837 Diagnosis. Notopodinae. Carapace widest anteriorly, dorsal surface evenly convex, granular; rostrum present. Eyestalk stout, less than one-quarter carapace width. Pereopod 4 dactylus acute. Maximum cl. 39 mm (Pl. 39f). Subtidal–slope (12–336 m). Temperate Northern and Tropical Atlantic, Tropical Eastern Pacific. 5 species (de Melo and Campos 1994: key to 3 species from Brazil).

Umalia Guinot, 1993 Diagnosis. Notopodinae. Carapace widest anteriorly, dorsal surface evenly convex, granular; rostrum present. Eyestalk stout, less than one-third carapace width. Pereopod 4 dactylus truncate. Maximum cl. 45 mm (Pl. 39k). Shelf, slope (50–400 m). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Australasia. 7 species (Davie and Short 1989: 2 species described as Ranilia; Guinot 1993: diagnosis).

Subfamily Ranininae De Haan, 1839 The subfamily includes a single species, the best known of all raninoidans. It is immediately recognised by its large size and scaly carapace. Diagnosis. Carapace widest across anterior margin; surface uniformly scabrous, shield-shaped, with 2 trifid processes on each anterolateral margin; fronto-orbital margin with 2 trifid processes on each anterolateral margin. Cheliped propodus with 2 spines on upper margin. Pereopods 2–5 dactyli broad, spatulate. Maximum cl. 150 mm. Ranina Lamarck, 1801 spanner crab Maximum cl. 150 mm (Pl. 39g). Intertidal, shelf (0–120 m). IndoWest Pacific, Temperate Australia (S to central NSW). 1 species (Ahyong et al. 2009).

Subfamily Raninoidinae Lörenthey, 1929 Diagnosis. Carapace widest behind anterior margin; surface smooth or granulate ; fronto-orbital margin at least half carapace width, rostrum scarcely extending beyond supraorbital and exorbital teeth. Cheliped propodus with or

without tooth on upper margin; dactylus without proximal tooth on upper margin. Pereopods 2–4 dactyli crescentshaped; pereopod 5 dactylus crescent-shaped or ovate. Notopoides Henderson, 1888 Diagnosis. Notopodinae. Carapace without anterolateral and lateral spine. Pleonite 1 as wide as carapace posterior margin. Cheliped propodus upper margin without spines or carinae. Pereopods 2–5 subequal. Maximum cl. 40 mm. Shelf, slope (148–645 m). Indo-West Pacific. 2 species (Goeke 1985: figures, key to species).

Notosceles Bourne, 1922 Diagnosis. Notopodinae. Carapace with anterolateral spine, without lateral spine. Pleonite 1 as wide as carapace posterior margin. Cheliped propodus upper margin with double carinae. Pereopod 5 much shorter and more slender than pereopods 2–4. Maximum cl. 23 mm. Shelf, slope (50–450 m). Temperate Northern W Pacific, IndoWest Pacific, Tropical Eastern Pacific, Temperate Australasia. 6 species (Dawson and Yaldwyn 2000: key to 6 species, distribution).

Raninoides H. Milne Edwards, 1837 Diagnosis. Ranininae. Carapace with anterolateral spine, without lateral spine. Pleonite 1 narrower than carapace posterior margin. Cheliped propodus upper margin with spine. Pereopod 5 much shorter and more slender than pereopods 2–4. Maximum cl. 39 mm (Pl. 39h, i). Subtidal–slope (18–366 m, possibly 677 m). Tropical Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific. 10 species (Chen and Türkay 2001: 4 species compared; Goeke 1980: species from Gulf of Mexico).

Subfamily Symethinae Goeke, 1981 Goeke (1981) erected Symethinae for a single genus with two American species on the basis of their greatly reduced eyes, inflated chelipeds and other features. A second species from the Central Pacific has been described since. Diagnosis. Carapace widest behind anterior margin; surface generally smooth, granulate or minutely spinulate; fronto-orbital margin less than half carapace width, rostrum extending well beyond supraorbital and exorbital teeth. Cheliped propodus swollen, unarmed; dactylus and fixed finger longer than palm. Pereopods 2–5 dactyli sickle-shaped. Symethis Weber, 1795 Subtidal, shelf (18–137 m). Tropical W Atlantic, Central IndoPacific, Eastern Indo-Pacific. Maximum cl. 35 mm. 3 species (Davie 1989: Central Pacific species; Goeke 1981: Gulf of Mexico, E Pacific species).

14 – Brachyura – crabs

References Ahyong ST, Naruse T, Tan SH, Ng PKL (2009) Part II. Infraorder Brachyura: Sections Dromiacea, Raninoida, Cyclodorippoida. In Crustacean fauna of Taiwan: Brachyuran crabs, volume 1 – Carcinology in Taiwan and Dromiacea, Raninoida, Cyclodorippoida. (Eds Chan T-Y, Ng PKL, Ahyong ST, Tan SH) pp. 27–198. National Taiwan Ocean University, Keelung. Brown IW, Scandol J, Mayer D, Campbell M, Kondyias S, et  al. (2008) ‘Reducing uncertainty in the assessment of the Australian spanner crab fishery. Fishing Industry project PR07–3314 – Final report’. Queensland Department of Primary Industries and Fisheries, Brisbane. Chen HL, Türkay M (2001) A new species of Raninoides (Crustacea: Decapoda: Raninidae) from off the southeastern coast of Hainan Island and Nansha Islands. Chinese Journal of Oceanology and Limnology 19, 332–336. doi:10.1007/BF02850737 Davie PJF (1989) Smethis [sic] corallica sp. nov. (Crustacea, Brachyura, Raninidae), the first member of the Smethinae [sic] to be recorded from the Indo-West Pacific region. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 11, 425–430. Davie PJF, Short JW (1989) Deepwater Brachyura (Crustacea: Decapoda) from southern Queensland, Australia with descriptions of four new species. Memoirs of the Queensland Museum 27, 157–187. de Melo GAS, Campos O, Junior (1994) O gênero Ranilia H. Milne Edwards, 1837, (Crustacea, Brachyura, Raninidae) no litoral brasileiro e descrição de uma nova espécie. Iheringia. Série Zoologia 77, 63–75. Goeke GD (1980) Range extensions of six western Atlantic frog crabs (Brachyura: Gymnopleura: Raninidae) with notes on the taxonomic status of Lyreidus bairdii. Proceedings of the Biological Society of Washington 93, 145–152. Goeke GD (1981) Symethinae, new subfamily, and Symethis garthi, new species, and the transfer of Raninoides ecuadorensis to Notosceles (Raninidae: Brachyura: Gymnopleura). Proceedings of the Biological Society of Washington 93, 971–981. Guinot D (1993) Données nouvelles sur les Raninoidea De Haan, 1841 (Crustacea Decapoda Brachyura Podotremata). Comptes Rendus de l’Académie des Sciences, Paris, Sciences de la vie 316, 1324–1331. Guinot D, Ng PKL (2020) On the nomenclature of Cyrtorhininae Guinot, 1993 (Brachyura, Raninoidea). Crustaceana 93, 71–76. doi:10.1163/​ 15685403-00003956 Karasawa H, Schweitzer CE, Feldmann RM, Luque J (2014) Phylogeny and classification of Raninoida (Decapoda: Brachyura). Journal of Crustacean Biology 34, 216–272. doi:10.1163/1937240X-00002216 Lam-Gordillo O, Ardisson P-L (2019) The global distribution and richness of frog crabs (Raninoidea). Crustaceana 92, 1–31. doi:10.1163/​ 15685403-00003832 Monod T (1956) Hippidea et Brachyura ouest-africains. Mémoires de I’Institut Français d’Afrique Noire 45, 1–674. Ng PKL, Guinot D (2020) Description of a new genus for Cyrtorhina balabacensis Serène, 1971, with notes on the Cyrtorhininae (Decapoda, Brachyura, Raninidae). Crustacean Research 49, 237–262. doi:10.18353/​ crustacea.49.0_237 Serène R (1971) Observations préliminaires sur des brachyoures nouveaux ou mal connus du Sud-Est Asiatique (Crustacea Decapoda). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 42, 903–918. Tavares M (2006) A new species of the crab genus Cosmonotus Adams & White in White, 1848 (Crustacea, Podotremata, Raninidae) from the Indo-West Pacific Ocean. Zoosystema 28, 533–537. Thomas LR, DiNardo GT, Lee H-H, Piner KR, Kahng SE (2013) Factors influencing the distribution of Kona crabs Ranina ranina (Brachyura: Raninidae) catch rates in the main Hawaiian islands. Journal of Crustacean Biology 33, 633–640. doi:10.1163/1937240X-00002171

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Van Bakel BWM, Guinot D, Artal P, Fraaije RHB, Jagt JWM (2012) A revision of the Palaeocorystoidea and the phylogeny of raninoidian crabs (Crustacea, Decapoda, Brachyura, Podotremata). Zootaxa 3215, 1–216. doi:10.11646/zootaxa.3215.1.1

Section Eubrachyura de Saint Laurent, 1980 The section name Eubrachyura was proposed by de Saint Laurent (1980) to distinguish ‘higher crabs’ from the ‘primitive crabs’ (Dromiacea, Homoloida, Raninoida, Cyclodorippoida). The female gonopores are on the thoracic sternum whereas in ‘primitive crabs’ they are on the coxae of pereopods 3 as in most other decapods. De Saint Laurent (1980) treated the primitive crabs collectively as the Podotremata whereas these are treated as four monophyletic sections. Eubrachyura includes two subsections, Heterotremata, probably paraphyletic, and the monophyletic Thoracotremata. References de Saint Laurent M (1980) Sur la classification et la phylogénie des Crustacés Décapodes Brachyoures. I. Podotremata Guinot, 1977 et Eubrachyura sect. nov. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 290, 1265–1268.

Subsection Heterotremata Guinot, 1977 This subsection of Eubrachyura was introduced by Guinot (1977) to include crabs with the female gonopore on the sternum and the male gonopore originating from the coxa of pereopod 5 but with the genital papilla sometimes displaced towards the sternum. Most families of brachyurans belong in this subsection. Inclusion of Hymenosomatoidea (Guinot 2011; Guinot et al. 2013) in which the male gonopore is on thoracic sternum 8 makes the group impossible to easily diagnose. Heterotremata are probably paraphyletic. Diagnosis. Male gonopores clearly coxal, with genital papilla protruding directly from coxa of pereopod 5, or male gonopores sternal, with genital papilla first emerging from surface of sternite of pereopod 5 (Hymenosomatidae only). References Guinot D (1977) Propositions pour une novelle classification des Crustacés, Décapodes, Brachyoures. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 285, 1049–1052. Guinot D (2011) The position of the Hymenosomatidae MacLeay, 1838, within the Brachyura (Crustacea, Decapoda). Zootaxa 2890, 40–52. doi:10.11646/zootaxa.2890.1.4 Guinot D, Tavares M, Castro P (2013) Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higherranked podotreme taxa. Zootaxa 3665, 1–414. doi:10.11646/zootaxa.​ 3665.1.1

474

Marine Decapod Crustacea

Aethroidea Dana, 1852 Aethroids are crabs with a ruggedly calcified carapace, mostly from rocky or coral reefs, although species of Hepatus and Hepatella favour sedimentary shorelines. The genera now included have had a convoluted taxonomic history having been variously placed in Calappidae, Leucosiidae and Parthenopidae owing to their diverse external features. The highly sculptured, eroded carapace of many aethrids closely resembles the reef rubble in which they occur, as well as several leucosiids and parthenopids from similar habitats. Guinot (1966, 1967) was first to recognise the close relationship between the aethrid genera, discuss their intrinsic connections and distinctions from other seemingly related groups. Aethrids have symmetrical chelipeds (asymmetrical in calappids) and usually a triangular buccal frame (subquadrate in parthenopids). Unlike leucosiids, with the inhalent respiratory channel under the exopod of maxilliped 3, the inhalent respiratory openings in aethrids are separated by maxillipeds 3 and located just above the base of the chelipeds. The exhalent channels on the endostome may be separated by a median septum (Fig.  14.20m). The group, as a superfamily comprising one family and seven genera, was recognised by Ng et al. (2008). Diagnosis. Carapace subovate to quadrate, as wide as or wider than long. Buccal cavity trapezoid-triangular or subquadrate, margins distinctly convergent anteriorly or subparallel. Inhalent respiratory openings lateral to maxilliped

above base of chelipeds. Maxilliped 3 merus triangular, tapering distally, rarely subquadrate–hexagonal; if maxilliped 3 subquadrate, retracted pereopods 1–5 completely concealed under carapace. Male pleonites 3–5 fused. Gonopod 1 stout. References Guinot D (1966) Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures. I. Les affinités des genres Aethra, Osachila, Hepatus, Hepatella et Actaeomorpha. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 38, 744–762. Guinot D (1967) Recherches préliminaires sur les groupements naturels chez les Crustacés. Décapodes Brachyoures I. Les affinités des genres Aethra, Osachila, Hepatus, Hepatella et Actaeomorpha (suite et fin). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 38, 828–845. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286.

Aethridae Dana, 1852 Figure 14.20, Plate 40a–f Two genera are restricted to the Indo-West Pacific, two genera occur only in the Americas and three span both regions. American species can be identified using the keys in Rathbun (1937), Hendrickx (1995) or Moscoso (2013). Implicit generic attributes. Orbit positioned anteriorly, eyes visible in dorsal view. Pereopods 2–5 smooth.

Key to genera of Aethridae 1. – 2. – 3. – 4. – 5. –

Pereopods 2–5 spinose, tuberculate or with pointed projections (Fig. 14.20o)����������������������������������������������������������������������2 Pereopods 2–5 smooth (Fig. 14.20a, n, e)�������������������������������������������������������������������������������������������������������������������������������������4 Carapace about as wide as long; regions swollen, divided by and surrounded by deep sharply-defined grooves; lateral margin with 4 blunt teeth, second most prominent (Fig. 14.20c); buccal frame subtriangular�������������������������������������������������������������������������������������������������������������������������������������������������������������� Drachiella Carapace wider than long; regions not surrounded by deep sharply-defined grooves; lateral margins irregularly dentate, serrated (Fig. 14.20b, g); buccal frame subquadrate (Fig. 14.20i, l)��������������������������������������������������3 Carapace front weakly projecting; lateral margins strongly expanded, upturned (Fig. 14.20b), fully concealing pereopods 1–5 when retracted. Maxilliped 3 merus distally truncate (Fig. 14.14i)���������������������������Aethra Carapace front prominently projecting; lateral margins weakly expanded, not upturned (Fig. 14.20g), only partially concealing pereopods 1–5 when retracted. Maxilliped 3 merus distally pointed (Fig. 14.14l)������������������������������������������������������������������������������������������������������������������������������������������������������������������������Sakaila Carapace anterolateral margins thick, not cristate or expanded; dorsal surface covered with pits and deep erosions (Fig. 14.20a, f)��������������������������������������������������������������������������������������������������������������������������������������������������������������5 Carapace anterolateral margins of carapace cristate, more or less expanded; dorsal surface smooth to granular, not pitted and eroded������������������������������������������������������������������������������������������������������������������������������������������������6 Carapace anterolateral margin with 2 obtuse angles dividing 3 straight sections (Fig. 14.20a)������������������Actaeomorpha Carapace anterolateral margin forming an arc of irregular teeth (Fig. 14.20f)������������������������������������������������������� Osachila

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Fig. 14.20.  Aethridae. Carapace, habitus: a, Actaeomorpha erosa Miers, 1877; b, Aethra scruposa (Linnaeus, 1764); c, Drachiella sculpta (Haswell, 1879); d, Hepatella peruviana Rathbun, 1933; e, Hepatus pudibundus (Herbst, 1785); f, Osachila tuberosa Stimpson, 1871; g, Sakaila africana Manning & Holthuis, 1981. Front, maxillipeds 3, ventral view: h, Actaeomorpha; i, Aethra; j, Hepatus; k, Osachila; l, Sakaila. Front, buccal frame, exhalent canals (maxillipeds 3 removed): m, Sakaila. Pereopod 5: n, Osachila; o, Sakaila.

6. –

Carapace evenly convex, usually with ~8 transverse arcs of tubercles (Pl. 40d); anterolateral margin tuberculate or with irregular low teeth; well marked suborbital depression (Fig. 14.20j); front ~4 or more times as wide as long (Fig. 14.20e)������������������������������������������������������������������������������������������������������������������������������� Hepatus Carapace evenly convex or with swollen regions; anterolateral margin widest at junction with posterolateral margin, with ~15 almost regular low teeth; weak suborbital depression; front 2–3 times wide as long (Fig. 14.20d)��������������������������������������������������������������������������������������������������������������������������������������������Hepatella Actaeomorpha Miers, 1877

Diagnosis. Carapace little wider than long, with regions divided by shallow ill-defined grooves; lateral margin thickened, not cristate or expanded, with 2 obtuse angles dividing 3 straight sections; rostrum half carapace width, square, with concave apex; buccal frame with convex lateral margins, triangular anterior margin. Maxilliped 3 merus triangular, with subacute apex. Maximum cl. 10 mm. Intertidal–shelf (0–60 m) (coral, reef). Indo-West Pacific, Temperate Australasia. 3 species (Guinot 1967: morphology; Miers 1877: description; Ng and Boyko 2017: distribution).

Aethra Latreille in Cuvier, 1816 Diagnosis. Carapace transversely elliptical; anterolateral and posterolateral margins continuous, upturned, with irregular short teeth; rostrum one sixth carapace width, tapering, weakly projecting; buccal frame rectangular. Maxilliped 3 merus subrectangular, with lateral lobe. Pereopods 1–5 fully concealed under

carapace when retracted; with acute tubercles and pointed projections. Maximum cl. 63 mm (Pl.40a). Intertidal–shelf (0–200 m). Temperate Northern Pacific, IndoWest Pacific, Tropical Eastern Pacific. 4 species (Garth 1958; Guinot 1966; Takeda 1975).

Drachiella Guinot in Serène & Soh, 1976 Diagnosis. Carapace little wider than long, with regions divided by and surrounded by deep sharply-defined grooves; lateral margin not cristate or expanded, with 4 blunt teeth, second most prominent; rostrum one-third carapace width, tapering; buccal frame with convex lateral margins, triangular anterior margin. Maxilliped 3 merus triangular, with subacute apex. Pereopods 1–5 with tubercles and spines. Maximum cl. 16 mm (Pl.40b). Subtidal (7–15 m; reef). Western and Central Indo-Pacific. 6 species (Edmondson 1935: key to 5 species as part of Actaeomorpha; Griffin 1972: figure; Serène and Soh 1976; Takeda and Tachikawa 1995: 2 species).

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Marine Decapod Crustacea

Hepatella Verrill, 1869 Diagnosis. Carapace transversely elliptical, evenly convex or with swollen upraised regions, with submarginal depression; anterolateral margin cristate, more or less expanded, with ~15 almost regular low teeth, widest at junction with posterolateral margin ; antero-posterolateral junction obtuse; rostrum one sixth carapace width, tapering, 2–3 times as wide as long; weak suborbital depression; buccal frame with convex lateral margins, triangular anterior margin. Maxilliped 3 merus triangular, with subacute apex. Maximum cl. 20 mm (Pl.40c). Subtidal, shelf (4–64 m). Tropical Eastern Pacific. 2 species (Rathbun 1911: figure; 1937: key to species).

Hepatus Latreille, 1802 Diagnosis. Carapace transversely elliptical, evenly convex, usually with ~8 transverse arcs of tubercles of tubercles; anterolateral margin cristate, more or less expanded, tuberculate or with irregular low teeth; antero- posterolateral junction obtuse; rostrum one sixth carapace width, tapering; well marked suborbital depression; buccal frame with convex lateral margins, triangular anterior margin. Maxilliped 3 merus triangular, with subacute apex. Maximum cl. 63 mm (Pl.40d). Intertidal–shelf (0–210 m). Tropical Atlantic, Temperate South America. 7 species (Holthuis 1959: 2 species; Martínez et al. 2009: distribution; Rathbun 1937: key to 5 species).

Osachila Stimpson, 1871 Diagnosis. Carapace with convex, irregularly dentate anterolateral margin, widest posteriorly; rostrum one-third carapace width, tapering; buccal frame with convex lateral margins, triangular anterior margin; endostome with exhalent respiratory channels not separated by median septum. Maxilliped 3 merus triangular, with subacute apex. Maximum cl. 25 mm (Pl.40e). Subtidal–slope (20–481 m). Tropical Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific. 11 species (Rathbun 1937: key to 7 species; Zimmerman and Martin 1999).

Sakaila Manning & Holthuis, 1981 Diagnosis. Carapace densely pitted or tuberculate, with convex anterolateral margin, widest posteriorly; rostrum one-third carapace width, tapering, projecting; orbit ventral, eyes not or scarcely visible in dorsal view; buccal frame rectangular; endostome exhalent respiratory channels separated by median septum. Maxilliped 3 merus distally pointed, with mesiodistal notch. Pereopods 2–5 spinose or tuberculate. Maximum cl. 60 mm (Pl.40f). Shelf, slope (35–367 m). Tropical Atlantic, Temperate Northern Pacific, Central and Eastern Indo-Pacific. 4 species (Manning and Holthuis 1981; Martin et al. 2009: key to species). References Edmondson CH (1935) New and rare Polynesian Crustacea. Occasional Papers of the Bernice P. Bishop Museum 10, 3–40.

Garth JS (1958) Brachyura of the Pacific coast of America. Oxyrhyncha. Allan Hancock Pacific Expeditions 21, 1–854. Griffin DJG (1972) Brachyura collected by Danish expeditions in southeastern Australia (Crustacea, Decapoda). Steenstrupia 2, 49–90. Guinot D (1966) Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures. I. Les affinités des genres Aethra, Osachila, Hepatus, Hepatella et Actaeomorpha. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 38, 744–762. Guinot D (1967) Recherches préliminaires sur les groupements naturels chez les Crustacés. Décapodes Brachyoures I. Les affinités des genres Aethra, Osachila, Hepatus, Hepatella et Actaeomorpha (suite et fin). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 38, 828–845. Hendrickx ME (1995) Cangrejos. In Guía FAO para la identificación de especes para los fines de la pesca. Pacifico centro-oriental. Volumen I. Plantas e invertebrados. (Eds Fischer W, F. K, Schneider W, Sommer C, Carpenter KE, Niem VH) pp. 565–636. FAO, Roma. Holthuis LB (1959) The Crustacea Decapoda of Suriname (Dutch Guiana). Zoologische Verhandelingen 44, 1–296, pls 1–16. Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Martin JW, Godwin S, Moffit R (2009) Additions to the decapod crustacean fauna of the Hawaiian Islands, I. A review of the genus Sakaila Manning and Holthuis, 1981 (Decapoda, Brachyura, Aethridae), with the description of a new species from the Northwestern Hawaiian Islands. Zootaxa 2018, 25–44. Martínez G, Scarabino F, Delgado E (2009) New records of the brachyuran crabs Hepatus pudibundus (Aethridae) and Persephona mediterranea (Leucosiidae) in their southernmost Western Atlantic distribution. Pan-American Journal of Aquatic Sciences 4, 279–282. Miers EJ (1877) On Actaeomorpha erosa, a new genus and species of Crustacea. Journal of the Linnean Society of London. Zoology 13, 183–185, pl. 14. Moscoso V (2013) Clave para identificación de crustáceos decápodos y estomatópodos del Perú. Boletín Instituto de Mar del Perú 28, 8–135. Ng PKL, Boyko CB (2017) New species and records of crabs of the families Dromiidae, Dynomenidae, Homolidae, Aethridae, Parthenopidae, Majidae, and Epialtidae (Crustacea: Decapoda: Brachyura) from Easter Island, with a review of the majid genus Schizophroida Sakai, 1933. Pacific Science 71, 197–227. doi:10.2984/71.2.8 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Rathbun MJ (1911) The stalk-eyed Crustacea of Peru and the adjacent coast. Proceedings of the United States National Museum 38, 531–620, pls 36–56. Rathbun MJ (1937) The oxystomatous and allied crabs of America. Bulletin of the United States National Museum 166, 1–278, pls 1–86. Serène R, Soh CL (1976) Brachyura collected during the Thai-Danish Expedition (1966). Phuket Marine Biological Center Research Bulletin 12, 1–37, figs 1–28, pls 1–8. Takeda M (1975) Preliminary description of a new crab of the Parthenopidae from the Seychelles Bank. Bulletin of the National Science Museum, Ser. A (Zoology) 1, 199–202, pl. 1. Takeda M, Tachikawa H (1995) Crabs of the Ogasawara Islands. VIII. Two species of the family Leucosiidae. Bulletin of the National Science Museum, Ser. Zoology 21, 211–218. Zimmerman TL, Martin JW (1999) Brachyuran crabs of Cocos Island (Isla Del Coco), Costa Rica: Leucosiidae, Calappidae, and Parthenopidae, with descriptions of two new species. Journal of Crustacean Biology 19, 643–668. doi:10.2307/1549267

14 – Brachyura – crabs

Bellioidea Dana, 1852 Bellioids are small compact shore crabs of mostly temperate Southern Hemisphere shorelines, noted for the antenna being reduced to a single article with a terminal seta or minute flagellum (Bouvier 1942; Guinot 1976). Salva and Feldmann (2001) treated the genera as members of Atelecyclidae, but seeing them as remote from other atelecyclids and figured representative species. The group has a complex

477

taxonomic history reflecting their uncertain affinities (Thompson and McLay 2005). One monotypic genus, Heterozius, is now placed in its own family (Števčić 2005). Diagnosis. Carapace longitudinally to transversely ovate. Antennal peduncle (excluding basal article) and flagellum minute, shorter than cornea (Figs 14.21d, e, 14.22b). Buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular. Male pleonites 3–5 fused (Figs 14.21f, 14.22c). Gonopod 1 stout.

Key to families of Bellioidea 1. –

Orbit and antennular fossa absent. Antennule not retractile, subparallel, directed forward (Fig. 14.21d, e)������� Bellidae Orbit totally closed, antennular fossa present. Antennule folding transversely (Fig. 14.22b)�������������������������Heteroziidae

References Bouvier E-L (1942) Les Crabes de la tribu des « Corystoïdea ». Mémoires de l’Académie des Sciences de l’Institut de France 65, 1–52. Castelnau F (1855) Expédition dans les parties centrales de l’Amérique du Sud, de Rio de Janeiro à Lima: et de Lima au Para. Septième Partie. Zoologies. Vol. 1. P. Bertrand, Paris. Guinot D (1976) Constitution de quelques groupes naturels chez les Crustacés Décapodes Brachyoures: I. La superfamille des Bellioidea et trois sous-familles de Xanthidae (Polydectinae Dana, Trichiinae de Haan, Actaeinae Alcock). Mémoires du Muséum National d’Histoire Naturelle. Nouvelle Série. Série A. Zoologie 97, 1–308. Salva EW, Feldmann RM (2001) Reevaluation of the family Atelecyclidae (Decapoda: Brachyura). Kirtlandia 52, 9–62. Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159.

Thompson GA, McLay CL (2005) Mating behaviour of Heterozius rotundifrons (Crustacea: Brachyura: Belliidae): is it a hard or soft shell mater? Marine and Freshwater Research 56, 1107–1116. doi:10.1071/MF04290

Belliidae Dana, 1852 Figure 14.21, Plate 40g Belliidae are now confined to three bellioid genera of shallow-water crabs without an orbit and antennular fossa. Diagnosis. Carapace longer than wide or slightly wider than long; front trilobed. Orbit and antennular fossa absent. Antennule not retractile, subparallel, directed forward. Chelipeds equal.

Key to genera of Belliidae 1. – 2. –

Carapace wider than long (Fig. 14.21a)����������������������������������������������������������������������������������������������������������������� Acanthocyclus Carapace as long as or longer than wide���������������������������������������������������������������������������������������������������������������������������������������2 Carapace widest anteriorly, with 4 triangular anterolateral teeth; pterygostomian region not visible in dorsal view (Fig. 14.21c)������������������������������������������������������������������������������������������������������������������������������������������ Corystoides Carapace widest near midlength, with 1 moderately prominent anterolateral tooth; pterygostomian region visible in dorsal view (Fig. 14.21b)������������������������������������������������������������������������������������������������������������������������������������Bellia Acanthocyclus Lucas, 1844

Diagnosis. Carapace ovate, convex, slightly wider than long; front trilobed; anterolateral margin with 4 teeth or lobes. Orbit semicircular. Pterygostomian region not visible in dorsal view. Maximum cl. 26 mm (Pl. 40g). Intertidal. Tropical Eastern Pacific, Temperate South America. 3 species (Meyer et al. 2013: COI differences; Navarrete and Castilla 1988: predatory feeding; Rathbun 1930: 3 species compared).

Bellia H. Milne Edwards, 1848 Diagnosis. Carapace as long as or longer than wide, widest near midlength, with 1 moderately prominent anterolateral tooth;

pterygostomian region visible in dorsal view. Maximum cl. 53 mm. Intertidal, sandy beaches. Tropical Eastern Pacific, Temperate South America (Pacific coast). 1 species (Rathbun 1930).

Corystoides Lucas, 1844 Diagnosis. Carapace longer than wide, widest anteriorly, with 4 triangular anterolateral teeth; pterygostomian region not visible in dorsal view. Maximum cl. 30 mm. Intertidal, subtidal (0–18 m). Temperate South America. 2 species (Guinot 1976: distribution; Rathbun 1930: figures; Scelzo et al. 2005: morphometry and growth).

478

Marine Decapod Crustacea

Fig. 14.21.  Belliidae. Carapace, habitus: a, Acanthocyclus; b, Bellia picta H. Milne Edwards, 1848; c, Corystoides. Right eye, antennule, antenna, epistome: d, Acanthocyclus; e, Bellia. f, male pleon, Bellia.

References Guinot D (1976) Constitution de quelques groupes naturels chez les Crustacés Décapodes Brachyoures: I. La superfamille des Bellioidea et trois sous-familles de Xanthidae (Polydectinae Dana, Trichiinae de Haan, Actaeinae Alcock). Mémoires du Muséum National d’Histoire Naturelle. Nouvelle Série. Série A. Zoologie 97, 1–308. Meyer R, Weis A, Melzer RR (2013) Decapoda of southern Chile: DNA barcoding and integrative taxonomy with focus on the genera Acanthocyclus and Eurypodius. Systematics and Biodiversity 11, 389–404. doi:10.1080/14772000.2013.833143 Navarrete SA, Castilla JC (1988) Foraging activities of Chilean intertidal crabs Acanthocyclus gayi Milne-Edwards et Lucas and A. hassleri Rathbun. Journal of Experimental Marine Biology and Ecology 118, 115–136. doi:10.1016/0022-0981(88)90235-3 Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Scelzo MA, Gavio MA, Martinez-Arca J (2005) Morphometry and relative growth of Corystoides abbreviatus (A. Milne Edwards, 1880) (Decapoda: Brachyura: Belliidae) in marine waters of Argentina. Nauplius 13, 137–147.

Heteroziidae Števčić, 2005 Figure 14.22 Heterozius rotundifrons A. Milne-Edwards, 1867 is a common endemic shore crab in New Zealand and the only member of its family (Števčić 2005). It is commonly called the big-handed crab because of the much enlarged cheliped on one side in males. Diagnosis. Carapace flat, wider than long; front sinuous, usually with shallow median notch; anterolateral margin unarmed. Orbit totally closed, antennular fossa present. Antennule folding transversely. Chelipeds significantly unequal in males, equal in females. Heterozius A. Milne-Edwards, 1867

Fig. 14.22.  Heteroziidae, Heterozius. a, habitus; b, right eye, antennule, antenna (shaded), epistome; c, male pleon.

References McLay CL (1988) Brachyura and crab-like Anomura of New Zealand [Crabs of New Zealand]. Leigh Laboratory Bulletin 22, 1–463. Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Thompson GA, McLay CL (2005) Mating behaviour of Heterozius rotundifrons (Crustacea: Brachyura: Belliidae): is it a hard or soft shell mater? Marine and Freshwater Research 56, 1107–1116. doi:10.1071/ MF04290

Bythograeoidea Williams, 1980

big-handed crab

vent crabs

Intertidal, buried in sand under stones. Maximum cl. 18 mm. Temperate Australasia (New Zealand endemic). 1 species (McLay 1988: identification, biology; Thompson and McLay 2005: mating behaviour).

Bythograeoidea includes one family, the only family of crabs truly endemic to deep-sea hydrothermal vents, where they can be abundant (Leignel et al. 2018; Martin and Haney

14 – Brachyura – crabs

2005). All species tend to be transversely oval with poorly defined carapace regions. In bythograeids, the orbits and eyes are strongly reduced, the cornea usually without pigment or ommatidial facets; they are blind or virtually blind as adults. The chelipeds are asymmetrical. Bythograeoids are naturally white or yellow, but specimens are frequently coated with red-brown to black deposits as a result of living in chemically reducing (low redox) environments. They feed on vent-associated bacteria (DeBevoise et al. 1990) or are predatory on other invertebrates (Voight 2002). The structural reduction and adaptations of bythograeids as result of living at depth on hydrothermal vents obscure recognition of their nearest brachyuran relatives. Their wider phylogenetic position is yet to be investigated in detail and most analyses have suggested placement among the xanthoid-like groups (Brösing et al. 2007, using foregut ossicles; Shen et al. 2013. using mitochondrial genomes; Tudge et al. 1998, using sperm ultrastructure). The most extensive, albeit preliminary, mitochondrial genome study to date (Mandon et al. 2017) placed bythograeids close to a clade of primarily portunoids, an affinity originally suggested by Williams (1980). The six genera and 18 species are confined to the Pacific where the most southerly species is from just north of

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New Zealand. An undescribed species has been recorded at 39°S (Stecher et al. 2002). Martin and Haney (2005) reviewed all records known at that time. Later, Leignel et  al. (2018) summarised type material and distribution of all species known at the time, and reviewed phylogenetics, biogeography, cryptic species, morphology and ecology. Diagnosis. Carapace subovate, wider than long, regions weakly to not defined. Orbit absent or reduced; eyestalk vestigial, reduced; cornea absent or if present, without pigment or facets. Antennule folding transversely. Buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular. Male pleonites 1–6 and telson free, or pleonites 4 and 5 partially fused. Gonopod 1 stout.

Bythograeidae Williams, 1980 Figure 14.23, Plate 40h, i Implicit generic attributes. Orbit ventrolateral concavity not filled with setae, with short suborbital area. Epistome strongly sinuous in frontal view. Eye slightly mobile, peduncle well developed. Maxilliped 3 coxa not obscured in ventral view, lateral extension visible.

Key to genera of Bythograeidae 1. – 2. – 3. – 4. – 5. –

Eye, antennule and antenna not deeply recessed. Orbits well circumscribed, complete (Fig. 14.23k, n)���������������������������2 Eye, antennule and antenna deeply recessed under front. Orbits open, incomplete (Fig. 14.23l, m, o, p)�������������������������3 Cornea unpigmented. Orbits ovate (Fig. 14.23k). Carapace flat, smooth, uniformly tomentose, widest at about midlength (Fig. 14.23a)������������������������������������������������������������������������������������������������������������������������������������Allograea Cornea pigmented. Orbits shallow, round (Fig. 14.23n). Carapace vaulted, with colour-tipped tubercles on frontolateral region, tomentose in patches, widest at about one-third length������������������������������������������������Cyanagraea Eye slightly mobile, peduncle well developed. Maxilliped 3 coxa not obscured in ventral view, lateral extension visible (Fig. 14.23f)����������������������������������������������������������������������������������������������������������������������������������������������������4 Eye immobile, peduncle vestigial, embedded in orbit wall. Maxilliped 3 coxa partially obscured in ventral view, lateral extension hidden by juncture of carapace ventral margin pterygostomian region with anterolateral sternal margin and cheliped coxa (Fig. 14.23e)�����������������������������������������������������������������������������������������������5 Ventrolateral edge of orbit with setose ventrolateral concave area, with short suborbital area (Fig. 14.23p)���������������������������������������������������������������������������������������������������������������������������������������������������������������� Segonzacia Ventrolateral edge of orbit without setose ventrolateral concave area, with prominent, wide, flat suborbital area (Fig. 14.23c, m)��������������������������������������������������������������������������������������������������������������������������������������������������Bythograea Epistome strongly sinuous in frontal view (Fig. 14.23l). Gonopod 2 shorter than gonopod 1 (Fig. 14.23g, h)�������������������������������������������������������������������������������������������������������������������������������������������������������� Austinograea Epistome gently sinuous in frontal view (Fig. 14.23o). Gonopod 2 about as long as gonopod 1 (Fig. 14.23i, j)��������������������������������������������������������������������������������������������������������������������������������������������������������������� Gandalfus Allograea Guinot, Hurtado & Vrijenhoek, 2002

Diagnosis. Carapace widest at about midlength, flat, smooth, uniformly tomentose. Eye, antennule and antenna not deeply recessed. Orbit ovate, well circumscribed, complete. Epistome

gently sinuous in frontal view. Cornea unpigmented. Maximum cl. 16 mm. Bathyal, seamount (2335 m). GOODS bathyal province: SE Pacific Ridges. 1 species (Guinot et al. 2002).

480

Marine Decapod Crustacea

Fig. 14.23.  Bythograeidae. Carapace: a, Allograea; b Austinograea. Habitus: c, Bythograea vrijenhoeki Guinot & Hurtado, 2003; d, Gandalfus yunohana (Takeda, Hashimoto & Ohta, 2000). Maxilliped 3 basis, ischium (dashed line), coxa (shaded), pterygostomian region (p), sternite 4 (s4), cheliped coxa (c1): e, Austinograea; f, Bythograea. Gonopods 1, 2: g, h, Austinograea; i, j, Gandalfus. Front, orbit, eyes, antennae, antennules, epistome: k, Allograea; l, Austinograea; m, Bythograea; n Cyanagraea; o, Gandalfus; p, Segonzacia.

Austinograea Hessler & Martin, 1989 Diagnosis. Carapace widest just anterior to midlength, flat. Eye, antennule and antenna deeply recessed under front. Orbit open, incomplete. Eye immobile, peduncle vestigial, embedded in orbit wall; cornea unpigmented or absent. Maxilliped 3 coxa partially obscured in ventral view, lateral extension hidden by juncture of carapace ventral margin pterygostomian region with anterolateral sternal margin and cheliped coxa. Gonopod 2 shorter than gonopod 1. Maximum cl. 38 mm. Slope–abyssal (1450–3660 m; hydrothermal vents). GOODS bathyal provinces: SE Pacific Ridges, Indian Ocean, South China Sea. 5 species (Guinot and Segonzac 2018: key to species; Hessler and Martin 1989: diagnosis).

Bythograea Williams, 1980 Diagnosis. Carapace widest at or just anterior to midlength, flat. Eye, antennule and antenna deeply recessed under front, with wide, flat suborbital plate demarcated by rows of granules, divided by uncalcified fissure. Orbit open, incomplete. Cornea unpigmented or pigmented. Gonopod 2 longer than gonopod 1. Maximum cl. 40 mm (Pl. 40i). Bathyal (2333–2832 m; hydrothermal vents). GOODS bathyal provinces: SE Pacific Ridges, Cocos Plate. 6 species (Guinot and

Hurtado 2003: rediagnosis; Guinot and Segonzac 2018: 2 species; McLay 2007: distributions, key to species; Williams 1980: diagnosis).

Cyanagraea de Saint Laurent, 1984 Diagnosis. Carapace widest at anterior one-third length, vaulted, with colour-tipped tubercles on frontolateral region, tomentose in patches. Eye, antennule and antenna not deeply recessed. Orbit shallow, round. Eye immobile; cornea pigmented. Gonopod 2 about as long as gonopod 1. Maximum cl. 75 mm. Bathyal (2535–2630 m; hydrothermal vents). GOODS bathyal province: Cocos Plate (East Pacific Rise). 1 species (Guinot and Segonzac 1997: distribution; Hessler and Martin 1989: redescription; de Saint Laurent 1984).

Gandalfus McLay, 2007 Diagnosis. Carapace widest just anterior to midlength, flat. Eye, antennule and antenna deeply recessed under front. Orbit elongate transversely, incomplete. Epistome gently sinuous in frontal view. Eye immobile, peduncle vestigial, embedded in orbit wall; cornea unpigmented. Maxilliped 3 coxa partially obscured in ventral view, lateral extension hidden by juncture of carapace ventral margin pterygostomian region with anterolateral sternal

14 – Brachyura – crabs

margin and cheliped coxa. Gonopod 2 about as long as gonopod 1. Maximum cl. 33 mm (Pl. 40h). Slope (239–1647 m; seamounts). Central Indo-Pacific, Temperate Australasia (Kermadec Is). 2 species (McLay 2007: diagnosis; Takeda et al. 2000: species description as Austinograea).

Segonzacia Guinot, 1989 Diagnosis. Carapace widest just anterior to midlength, flat. Eye, antennule and antenna deeply recessed under front. Orbit open, incomplete; with ventrolateral concavity filled with setae. Cornea unpigmented. Gonopod 2 longer than gonopod 1. Maximum cl. 22 mm. Slope–bathyal (850–3478 m; hydrothermal vents). GOODS bathyal province: North Atlantic Ocean (Mid-Atlantic). 1 species (Guinot 1989: diagnosis; Williams 1988: species description). References Brösing A, Richter S, Scholtz G (2007) Phylogenetic analysis of the Brachyura (Crustacea, Decapoda) based on characters of the foregut with establishment of a new taxon. Journal of Zoological Systematics and Evolutionary Research 45, 20–32. doi:10.1111/j.1439-0469.2006.00367.x de Saint Laurent M (1984) Crustacés Décapodes d’un site hydrothermal actif de la dorsale du Pacifique oriental (13° Nord), en provenance de la campagne française Biocyatherm. Comptes Rendus de l’Académie des Sciences 299, 355–360. DeBevoise AE, Childress JJ, Withers NW (1990) Carotenoids indicate differences in diet of the hydrothermal vent crab Bythograea thermydron (Brachyura). Marine Biology 105, 109–115. doi:10.1007/BF01344276 Guinot D (1989) Description de Segonzacia gen. nov. et remarques sur Segonzacia mesatlantica (Williams): campagne HYDROSNAKE 1988 sur la dorsale médio-Atlantique (Crustacea Decapoda Brachyura). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série 11, 347–385, pls 1–3. Guinot D, Hurtado LA (2003) Two new species of hydrothermal vent crabs of the genus Bythograea from the southern East Pacific Rise and from the Galapagos Rift (Crustacea Decapoda Brachyura Bythograeidae). Comptes Rendus Biologies 326, 423–439. doi:10.1016/S1631-0691(03)​00126-4 Guinot D, Segonzac M (1997) Description d’un crabe hydrothermal nouveau du genre Bythograea (Crustacea Decapoda Brachyura) et remarques sur les Bythograeidae de la dorsale du Pacifique oriental. Zoosystema 19, 121–150. Guinot D, Segonzac M (2018) A review of the brachyuran deep-sea vent community of the western Pacific, with two new species of Austinograea Hessler & Martin, 1989 (Crustacea, Decapoda, Brachyura, Bythograeidae) from the Lau and North Fiji Back-Arc Basins. Zoosystema 40, 75–107. doi:10.5252/zoosystema2018v40a5 Guinot D, Hurtado LA, Vrijenhoek RC (2002) New genus and species of brachyuran crab from southern East Pacific Rise (Crustacea Decapoda Brachyura Bythograeidae). Comptes Rendus Biologies 325, 1143–1152. doi:10.1016/S1631-0691(02)01520-2 Hessler RR, Martin JW (1989) Austinograea williamsi, new genus, new species, a hydrothermal vent crab (Decapoda: Bythograeidae) from the Mariana Back-Arc Basin, western Pacific. Journal of Crustacean Biology 9, 645–661. doi:10.2307/1548594 Leignel V, Hurtado LA, Segonzac M (2018) Ecology, adaptation and acclimatisation mechanisms of Bythograeidae Williams, 1980, a unique endemic hydrothermal vent crabs family: current state of knowledge. Marine and Freshwater Research 69, 1–15. doi:10.1071/MF17007 Mandon P, Aznar-Cormano L, Hourdez S, Samadi S (2017) Assembly of the mitochondrial genome of the hydrothermal vent crab Segonzacia mesatlantica and detection of potential nuclear pseudogenes.

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­ itochondrial DNA. Part B, Resources 2, 291–293. doi:10.1080/23802 M 359.2017.1318674 Martin JW, Haney TA (2005) Decapod crustaceans from hydrothermal vents and cold seeps: a review through 2005. Zoological Journal of the Linnean Society 145, 445–522. doi:10.1111/j.1096-3642.2005.00178.x McLay CL (2007) New crabs from hydrothermal vents of the Kermadec Ridge submarine volcanoes, New Zealand: Gandalfus gen. nov. (Bythograeidae) and Xenograpsus (Varunidae) (Decapoda: Brachyura). Zootaxa 1524, 1–22. doi:10.11646/zootaxa.1524.1.1 Shen H, Braband A, Scholtz G (2013) Mitogenomic analysis of decapod crustacean phylogeny corroborates traditional views on their relationships. Molecular Biology and Evolution 66, 776–789. Stecher J, Türkay M, Borowski C (2002) Faunal assemblages on the Pacific-Antarctic Ridge near the Foundation Seamount Chain (37°30′ S, 110°30′ W). Cahiers de Biologie Marine 43, 271–274. Takeda M, Hashimoto J, Ohta S (2000) A new species of the family Bythograeidae (Crustacea, Decapoda, Brachyura) from the hydrothermal vents along volcanic front of the Philippine Sea Plate. Bulletin of the National Science Museum, Ser. Zoology 26, 159–172. Tudge CC, Jamieson BGM, Segonzac M, Guinot D (1998) Spermatozoal ultrastructure in three species of hydrothermal vent crab, in the genera Bythograea, Austinograea and Segonzacia (Decapoda, Brachyura, Bythograeidae). Invertebrate Reproduction and Development 34, 13–23. doi:10.1080/07924259.1998.9652349 Voight JR (2002) A review of predators and predation at deep-sea hydrothermal vents. Cahiers de Biologie Marine 41, 155–166. Williams AB (1980) A new crab family from the vicinity of submarine thermal vents on the Galapagos Rift (Crustacea: Decapoda: Brachyura). Proceedings of the Biological Society of Washington 93, 443–472. Williams AB (1988) New marine decapod crustaceans from waters influenced by hydrothermal discharge, brine, and hydrocarbon seepage. Fishery Bulletin 86, 263–287.

Calappoidea De Haan, 1833 box crabs, moon crabs The family name Calappidae was previously used to include several genera, some of which are now treated as matutids, orithyiids and aethrids. Bellwood (1996, 1998) reviewed the usage of the name and concluded on the basis of a phylogenetic analysis that the family, as understood at that time, was polyphyletic just as earlier authors had suspected (Števčić 1983). The superfamily Calappoidea is retained for just two families, Calappidae and Matutidae but these might not be closely related (Davie 2002; Ng et al. 2008). On the other hand, a limited study comparing the complete mitogenome of two representatives found them to be sister taxa (Lu et al. 2020). Calappoids are heavily armoured crabs of tropical and temperate reefs and sandy beaches worldwide. Diagnosis. Carapace circular to subcircular or transversely ovate. Antennule folding transversely, obliquely or vertically. Buccal cavity approximately trapezoid–triangular. Inhalent respiratory openings lateral to maxilliped above base of chelipeds. Maxilliped 3 merus triangular, tapering distally, rarely subquadrate-hexagonal. Male pleonites 3–5 fused. Gonopod 1 stout. Female gonopores fully concealed by pleon, not exposed.

482

Marine Decapod Crustacea

Key to families of Calappoidea 1.



Chelipeds asymmetrical. Major chela (usually right) with blunt cutting tooth at base of dactylus (Fig. 14.25e–g). Minor chela fingers long, forceps-like. Pereopods 2–5 propodi and dactyli linear, slender. Carapace posterolateral margin sometimes expanded as wing-like process or series of teeth, or with posterolateral tooth or spine (Fig. 14.24)������������������������������������������������������������������������������������������������������Calappidae Chelipeds symmetrical, dactylus without blunt cutting tooth. Pereopods 2–5 propodi and dactyli paddle-like, flat. Carapace lateral margin with single tooth, usually prominent (Fig. 14.26)����������������������� Matutidae

References Bellwood O (1996) A phylogenetic study of the Calappidae H. Milne Edwards 1837 (Crustacea: Brachyura) with a reappraisal of the status of the family. Zoological Journal of the Linnean Society 118, 165–193. doi:10.1111/j.1096-3642.1996.tb00225.x Bellwood O (1998) The phylogeny of box crab genera (Crustacea: Brachyura: Calappidae) with notes on their fossil record, biogeography and depth distribution. Journal of Zoology 244, 459–471. doi:10.1111/j.1469-7998.​ 1998.tb00051.x Davie PJF (2002) Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells A, Houston WWK, Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing, Melbourne. Lu X, Gong L, Zhang Y, Chen J, Liu L, et al. (2020) The complete mitochondrial genome of Calappa bilineata: The first representative from the family Calappidae and its phylogenetic position within Brachyura. Genomics 112, 2516–2523. doi:10.1016/j.ygeno.2020.02.003 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Števčić Z (1983) Revision of the Calappidae. Australian Museum Memoir 18, 165–171. doi:10.3853/j.0067-1967.18.1984.382

Calappidae De Haan, 1833 box crabs, shame-faced crabs Figures 14.24, 14.25, Plate 41a–i The Calappidae, or box crabs, are a distinctive family with highly modified chelipeds. The carapace of species of Calappa, in particular have wing-like lateral expansions of the carapace forming a cavity underneath. This hides the walking legs while the crab is buried in sand and gives them the name ‘box crabs’. The tall flat chelipeds are held in front

of the carapace, a behaviour that gives them the alternative name of ‘shame-faced crabs’. Calappids are mostly shallow-water crabs of sandy bottoms around coral reefs but some occur to slope depths. They burrow into sand using their chelipeds and walking legs. The most distinctive feature is a curved accessory tooth on the outside of the dactylus of the flat major cheliped which opposes teeth on the finger (Fig. 14.25d–e). These are used to peel open gastropod mollusc shells to prey on the snail or hermit crabs housed inside (Shoup 1968). Because most gastropods coil clockwise when viewed from above, this predatory tool favours right-handedness and virtually all calappids have the larger modified chela on the right (Ng and Tan 1985). The genera appear well supported on the basis of morphology (Bellwood 1998) and molecular data (EwersSaucedo et al. 2016). Diagnosis. Carapace posterolateral margin sometimes expanded as wing-like process or series of teeth, or with posterolateral tooth or spine. Maxilliped 3 exopod narrow, with flagellum. Chelipeds asymmetrical, dactylus of one chela (usually right side) with curved tooth on outer proximal surface of dactylus occluding with shallow cavity on fixed finger; curved tooth absent on other cheliped dactylus (Fig.  14.25e–g). Pereopods 2–5 slender, with pointed dactyli. Implicit generic attributes. Supraorbital border with 1 fissure; branchial regions without ridges. Buccal cavity with anterior median septum. Pleonite 2 carina entire. Cheliped merus distal outer surface without spines. Pereopod 5 ischium-merus flexor margin smooth.

Key to genera of Calappidae 1. – 2. –

Carapace posterolateral region not expanded into wing-like projection, with 1 or more teeth or long spine (Figs 14.24a, e, f, h, i, 14.25c)������������������������������������������������������������������������������������������������������������������������������������������������������2 Carapace posterolateral region expanded into wing-like projection or series of teeth (Figs 14.24b–d, g, j, 14.25a, b)����������������������������������������������������������������������������������������������������������������������������������������������������6 Cheliped merus distal outer surface with 1 spine as long as half carapace width, usually with second more proximal shorter spine (Fig. 14.24a)���������������������������������������������������������������������������������������������������������������� Acanthocarpus Cheliped merus distal outer surface without 1 very long spine (Mursia with 2 or 3 distal spines shorter than carapace width)���������������������������������������������������������������������������������������������������������������������������������������������������������������������������3

14 – Brachyura – crabs

483

Fig. 14.24.  Calappidae. Habitus, carapace: a, Acanthocarpus bispinosus A. Milne-Edwards, 1880; b, Calappa hepatica (Linnaeus, 1758); c, Calappa philargius (Linnaeus, 1758; d, Calappula tortugae (Rathbun, 1933); e, Cryptosoma balguerii (Desbonne in Desbonne & Schramm, 1867); f, Cycloes granulosa De Haan, 1837; g, Cyclozodion angustum (A. Milne-Edwards, 1880); h, Mursia aspera Alcock, 1899; i, Mursia bicristimana Alcock & Anderson, 1894; j, Paracyclois milneedwardsii Miers, 1886.

3. – 4. – 5. – 6. – 7. –

Carapace subcircular, with negligible or small tooth at lateral angle; front as wide as orbit, bidentate, slightly projecting; branchial regions with 3 ridges subparallel to anterolateral margin, furrows bordering cardiac region most pronounced (Fig. 14.24e, f)�������������������������������������������������������������������������������������������������4 Carapace subcircular to transversely oval, with spine (usually prominent) at lateral angle; front wider than orbit, trilobate, median lobe projecting; branchial regions without ridges (Fig. 14.24h, i)���������������������������������������������5 Carapace wider than long; lateral tooth small (Fig. 14.24e). Cheliped carpus upper margin smooth (Fig. 14.25f), anterior angle smooth on inner face; palm upper margin with second and third teeth smooth on inner face; dactylus with milled band on inner surface having more than 25 ridges�������������� Cryptosoma Carapace as wide as long; lateral tooth absent or indistinct (Fig. 14.24f). Cheliped carpus upper margin dentate (Fig. 14.25g), anterior angle granulate on inner face; palm upper margin with second and third teeth granulate on inner face; dactylus with milled band on inner surface having fewer than 15 ridges������������ Cycloes Cheliped dactylus inner surface irregularly granulate. Pleonite 2 carina entire����������������������������������������������������Platymera Cheliped dactylus inner surface with milled ridge. Pleonite 2 carina trilobed (Fig. 14.25i)������������������������������������� Mursia Pereopod 5 ischium-merus flexor margin with row of spines (Fig. 14.25h)��������������������������������������������������������� Paracyclois Pereopod 5 ischium-merus flexor margin smooth���������������������������������������������������������������������������������������������������������������������7 Carapace dentate wings not exceeding anterolateral margins (Fig. 14.24g)������������������������������������������������������� Cyclozodion Carapace dentate wings exceeding anterolateral margins (Figs 14.24b–d, 14.25a, b)�����������������������������������������������������������8

484

Marine Decapod Crustacea

Fig. 14.25.  Calappidae. Carapace: a, Calappa woodmasoni Alcock, 1896 ; b, Calappa japonica Ortmann, 1892; c, Platymera gaudichaudii H. Milne Edwards, 1837. Orbits, buccal cavity: d, Calappa. Major pereopod, outer view: e, Calappa; f, Cryptosoma; g, Cycloes. Pereopod 5: h Paracyclois. Pleonites 2–6, telson: i, Mursia.

8. –

Carapace supraorbital border with 2 fissures (Fig. 14.24c); buccal cavity with median septum anteriorly (Fig. 14.25d) (sometimes concealed). Chela without transverse milled ridges on inner surface�����������������������Calappa Carapace supraorbital border with 1 fissure (Fig. 14.24d); buccal cavity without median septum anteriorly. Chela with 2 transverse milled ridges on inner face��������������������������������������������������������������������������������������������� Calappula Acanthocarpus Stimpson, 1871

Calappula Galil, 1997

Diagnosis. Carapace subcircular, with oblique posterolateral tooth; front wider than orbit, trilobate, median lobe projecting. Cheliped merus distal outer surface with 2 spines, more distal spine stronger, extending laterally about half carapace width (proximal sometimes obsolete); dactylus inner surface with milled ridge. Maximum cl. 43 mm (Pl. 41a). Shelf, slope (20–1034 m). Temperate Northern and Tropical Atlantic, Tropical Eastern Pacific, Temperate South America (Atlantic coast). 5 species (Alves-Júnior et al. 2018: figure, distribution; Garth 1973: new species; Haefner 1981: biology; MañéGarzón 1979: new species; Rathbun 1937: key to 2 species).

Diagnosis. Carapace posterolateral region expanded into wing-like projection exceeding mid-carapace width, usually dentate, with ventral cavity for walking legs; front narrower than orbit, tridentate; buccal cavity without anterior median septum. Cheliped palm with transverse milled ridge on inner face. Maximum cl. 38 mm. Subtidal–slope (13–238 m). Temperate Northern and Tropical W Atlantic, Eastern Indo-Pacific. 2 species (Galil 1997: diagnosis; Williams and Child 1989: C. tortugae (Rathbun, 1933) figured).

Calappa Weber, 1795 Diagnosis. Carapace posterolateral region expanded into winglike projection exceeding mid-carapace width, usually dentate, with ventral cavity for walking legs; front narrower than orbit, tridentate; supraorbital border with 2 fissures. Cheliped palm without transverse milled ridge on inner face. Maximum cl. 104 mm (Pl. 41b–d). Intertidal–slope (0–1246 m). Cosmopolitan except Temperate South America and polar seas. 41 species (Galil 1997: rediagnosis, 4 species groups defined, key to 20 Indo-Pacific species; Lai et al. 2006: descriptions, list of species, phylogeny and key to 4 species of C. lophos-group). The genus contains twice as many species as in Galil’s (1997) key but Atlantic species have not been reviewed.

Cryptosoma Brullé, 1837 Diagnosis. Carapace subcircular, with small tooth at lateral angle; front as wide as orbit, bidentate, slightly projecting. Cheliped carpus upper margin smooth, anterior angle smooth on inner face; palm upper margin with second and third teeth smooth on inner face; dactylus with milled band on inner surface having more than 25 ridges. Maximum cl. 39 mm (Pl. 41e). Subtidal–slope (3–230 m). Temperate Northern and Tropical Atlantic, Eastern Indo-Pacific. 4 species (Galil and Clark 1996: rediagnosis, key to species; Tamburus et al. 2019: distribution).

Cycloes De Haan, 1837 Diagnosis. Carapace subcircular, with small tooth at lateral angle; front as wide as orbit, bidentate, slightly projecting. Cheliped carpus upper margin dentate, anterior angle granulate on inner face;

14 – Brachyura – crabs

palm upper margin with second and third teeth granulate on inner face; dactylus with milled band on inner surface having fewer than 15 ridges. Maximum cl. 38 mm (Pl. 41f). Subtidal, shelf (15–110 m). Western and Central Indo-Pacific, Eastern Indo-Pacific. 2 species (Galil and Clark 1996: rediagnosis, key to species).

Cyclozodion Williams & Child, 1989 Diagnosis. Carapace posterolateral margin with dentate ridge, not reaching maximum carapace width; front narrower than orbit, tridentate. Cheliped palm with transverse milled ridge vertically on inner face. Maximum cl. 26 mm (Pl. 41g). Shelf, slope (31–640 m). Temperate Northern and Tropical W Atlantic. 2 species (Williams and Child 1989: diagnosis, key to species).

Mursia Desmarest, 1823 Diagnosis. Carapace subcircular to transversely oval, with lateral spine, usually prominent; front wider than orbit, trilobate, median lobe projecting; supraorbital border with 1 or 2 fissures; branchial regions without ridges. Pleonite 2 carina trilobed. Cheliped dactylus inner surface with milled ridge. Maximum cl. 58 mm (Pl. 41h). Subtidal–slope (9–732 m). Temperate Northern Pacific, Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific, Temperate South America, Temperate Southern Africa. 29 species (Galil 1993: rediagnosis, key to 15 species). Twice as many species are now known as appeared in Galil’s (1993) key (e.g. Crosnier 1997; Galil 2001; Kumar et al. 2013).

Paracyclois Miers, 1886 Diagnosis. Carapace subcircular, posterolateral region expanded as triangular teeth, not reaching maximum carapace width; front narrower than orbit, tridentate. Cheliped palm with transverse milled ridge vertically on inner face. Pereopod 5 ischium-merus flexor margin with row of spines. Maximum cl. 62 mm (Pl. 41i). Shelf, slope (137–365 m). Tropical Atlantic, Temperate Northern Pacific, Central Indo-Pacific. 2 species (Williams and Child 1989: rediagnosis, key to species).

Platymera H. Milne Edwards, 1837 Diagnosis. Carapace transversely ovate, with posterolateral spine one-quarter carapace width; front narrower than orbit, tridentate; branchial regions without ridges. Cheliped dactylus inner surface irregularly granulate. Maximum cl. 64 mm. Shelf, slope (50–550 m). Tropical Eastern Pacific, Temperate South America (E Pacific). 1 species (Galil 1993: rediagnosis; Retamal et al. 2013: figure, distribution). References Alves-Júnior Fd A, Pereira ÂF, Câmara de Araújo Md SL (2018) New records of two species of Acanthocarpus Stimpson, 1871 (Crustacea Decapoda: Calappidae) in northeastern Brazil. Pesquisa e Ensino em Ciências Exatas e da Natureza 2, 60–66.

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Bellwood O (1998) The phylogeny of box crab genera (Crustacea: Brachyura: Calappidae) with notes on their fossil record, biogeography and depth distribution. Journal of Zoology 244, 459–471. doi:10.1111/j.1469-7998.​ 1998.tb00051.x Crosnier A (1997) Une espèce nouvelle de Mursia de Nouvelle-Calédonie (Crustacea, Decapoda, Brachyura, Calappidae). Zoosystema 19, 151–158. Ewers-Saucedo C, Wares JP, Hanel R, Brandis D (2016) Evolution of male copulatory organs in box crabs (Decapoda: Eubrachyura: Calappidae De Haan, 1833). Journal of Crustacean Biology 36, 804–814. doi:10.1163/1937240X-00002479 Galil BS (1993) Crustacea Decapoda: a revision of the genus Mursia Desmarest, 1823 (Calappidae). In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 10. Mémoires du Muséum National d’Histoire Naturelle, Paris 156, 347–379. Galil BS (1997) Crustacea Decapoda: a revision of the Indo-Pacific species of the genus Calappa Weber, 1795 (Calappidae). In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 18. Mémoires du Muséum National d’Histoire Naturelle, Paris 176, 271–335. Galil B (2001) The Calappidae of the Marquesas Islands with a description of a new species of Mursia (Crustacea, Decapoda, Brachyura). Zoosystema 23, 499–505. Galil BS, Clark PF (1996) A revision of Cryptosoma Brullé, 1837 and Cycloes De Haan, 1837 (Crustacea: Brachyura: Calappidae). Zoological Journal of the Linnean Society 117, 175–204. doi:10.1111/j.1096-3642.​ 1996.tb02155.x Garth JS (1973) New taxa of brachyuran crabs from deep water off western Peru and Costa Rica. Bulletin of the Southern California Academy of Sciences 72, 1–12. Haefner PA (1981) Morphometry, reproductive biology, and diet of Acanthocarpus alexandri Stimpson, 1871 (Decapoda, Brachyura) in the Middle Atlantic Bight. Journal of Crustacean Biology 1, 348–357. doi:10.2307/1547967 Kumar BA, Kumar MS, Galil BS (2013) Calappid and leucosiid crabs (Crustacea: Decapoda: Brachyura) from Kerala, India, with the description of a new species of Mursia Desmarest, 1823, from the Arabian Sea and redescription of M. bicristimana Alcock & Anderson, 1894. Zootaxa 3746, 529–551. doi:10.11646/zootaxa.3746.4.2 Lai JCY, Chan WK, Ng PKL (2006) Preliminary molecular and morphological study of the Calappa lophos species group (Decapoda:Brachyura:Calap pidae). Journal of Crustacean Biology 26, 193–205. doi:10.1651/​C-2571.1 Mañé-Garzón F (1979) Un nuevo Decapoda Brachyura Oxystomata de la costa oceanica del Uruguay: Acanthocarpus meridionalis n. sp. Revista de Biologia del Uruguay 7, 29–38. Ng PKL, Tan LWH (1985) ‘Right handedness’ in heterochelous calappoid and xanthoid crabs: suggestion for a functional advantage. Crustaceana 49, 98–100. doi:10.1163/156854085X00288 Rathbun MJ (1937) The oxystomatous and allied crabs of America. Bulletin of the United States National Museum 166, 1–278, pls 1–86. Retamal MA, Ernst B, Chamorro J, Manríquez P (2013) Descripción y distribución geográfica de una nueva especie para el archipiélago Juan Fernández: Platymera gaudichaudii (H. Milne-Edwards, 1837) (Decapoda, Calappidae). Latin American Journal of Aquatic Research 41, 1010–1012. doi:10.3856/vol41-issue5-fulltext-21 Shoup JB (1968) Shell opening by crabs of the genus Calappa. Science 160, 887–888. doi:10.1126/science.160.3830.887 Tamburus AF, França NFC, Mantelatto FL (2019) First record of the box crab Cryptosoma balguerii (Desbonne in Desbonne and Schramm, 1867) (Brachyura: Calappidae) from São Paulo, Brazil revealed by DNA markers. Nauplius 27, e2019022. doi:10.1590/2358-2936e2019022 Williams AB, Child CA (1989) Comparison of some genera and species of box crabs (Brachyura: Calappidae), aouthwestern North Atlantic, with description of a new genus and species. Fishery Bulletin 87, 105–121.

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Matutidae De Haan, 1835 moon crabs Figure 14.26, Plate 41j–l Matutids, moon crabs, are noted for the expanded, flattened, ends of the legs which aid the crabs to swim and burrow. The carapace is oval or round with a laterally-directed spine on the lateral margin. The crabs live in the surf zone on tropical beaches, where they are active predators (Perez and Bellwood 1988), or on deeper sandy bottoms. Long considered a subfamily of Calappidae, matutids are now ranked as a separate family, possibly not closely

related to calappids. Galil and Clark (1994) created the current classification by dividing the morphologically diverse genus Matuta. Our key relies mainly on the ornamentation of the face of the chela, features of pereopod 4 and the development of the anterolateral spine on the carapace. Diagnosis. Carapace anterolateral margin with 1 tooth. Maxilliped 3 exopod triangular, without flagellum. Chelipeds symmetrical; dactyli of both chelipeds similar. Pereopods 2–5 forming swimming legs, propodus and dactylus flattened, paddle-like (Fig. 14.26a).

Key to genera of Matutidae 1. – 2. – 3. –

Chela outer surface smooth (Fig. 14.26e). Carapace pterygostomian surface with stridulating organ of 1 row of tubercles; setae margined channel descending from lateral end of orbit (inhalent canal) straight���������Mebeli Chela outer surface sculptured (Fig. 14.26b–d). Carapace pterygostomian region granulose or with several rows of tubercles; setae margined channel descending from lateral end of orbit (inhalent canal) curved�������������������2 Chela outer surface with midpalmar milled stridulating ridge (Fig. 14.26c). Carapace lateral spine rudimentary; antennular fossa excluded from orbit; pterygostomian region evenly granulose. Pleonite 2 carinate����������������������������������������������������������������������������������������������������������������������������������������������������������������������������Izanami Chela outer surface with midpalmar ridge not milled (Fig. 14.26b, d), palmar teeth interiorly striate. Carapace lateral spine well developed; antennular fossa continuous with orbit; pterygostomian region with several rows of elongate tubercles. Pleonite 3 carinate�������������������������������������������������������������������������������������������������3 Chela outer surface with ridge oblique; dactylus outer surface ridge in male milled throughout (Fig. 14.26d). Pereopod 4 carpus unicarinate������������������������������������������������������������������������������������������������������������� Matuta Chela outer surface with midpalmar ridge of tubercles parallel to lower margin; dactylus outer surface ridge in male distally milled or smooth (Fig. 14.26b). Pereopod 4 carpus bicarinate�����������������������������������������Ashtoret Ashtoret Galil & Clark, 1994

Diagnosis. Carapace lateral spine well developed; antennular fossa continuous with orbitpterygostomian region with several rows of elongate tubercles; inhalant canal curved. Chela outer

surface with midpalmar ridge of tubercles parallel to lower margin; dactylus outer surface ridge in male distally milled or smooth. Pereopod 4 carpus upper margin bicarinate. Maximum cl. 66 mm (Pl. 41j).

Fig. 14.26.  Matutidae. a, Matuta victor (Fabricius, 1781). Chela, outer face: b, Ashtoret; c, Izanami; d, Matuta; e, Mebeli.

14 – Brachyura – crabs

Intertidal–shelf (0–50 m). Temperate Northern W Pacific, Indo-West Pacific. 8 species (Galil and Clark 1994: diagnosis, key to species). One species has invaded the eastern Mediterranean Sea (Galil and Mendelson 2013).

Izanami Galil & Clark, 1994 Diagnosis. Carapace lateral spine rudimentary; antennular fossa excluded from orbit; pterygostomian region evenly granulose; inhalant canal curved. Chela outer surface with milled midpalmar stridulating ridge, palmar teeth interiorly granulose. Pereopod 4 carpus upper margin unicarinate. Maximum cl. 33 mm. Shelf, slope (35–217 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 2 species (Galil and Clark 1994: diagnosis, key to species).

Mebeli Galil & Clark, 1994 Diagnosis. Carapace lateral spine well developed; antennular fossa continuous with orbit; pterygostomian region with stridulating organ consisting of 1 row of tubercles. Inhalant canal straight. Chela outer surface smooth. Pereopod 4 carpus upper margin unicarinate. Maximum cl. 21 mm. Intertidal–shelf (0–30 m). Tropical E Atlantic. 1 species (Galil and Clark 1994).

Matuta Weber, 1795 Diagnosis. Carapace inhalant canal curved; antennular fossa continuous with orbit; pterygostomian region with several rows of elongate tubercles; inhalant canal curved. Chela outer surface with ridge oblique; dactylus outer surface ridge in male milled throughout. Pereopod 4 carpus upper margin unicarinate. Maximum cl. 70 mm (Pl. 41k, l). Intertidal–shelf (0–40 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate E Australia. 4 species (Galil and Clark 1994: rediagnosis, key to 3 species; Lai and Galil 2007: fourth species; Romimohtarto 1972: key to 5 species). Miers (1877) published excellent illustrations of ten species and varieties, not all now recognised. Matuta victor (Fabricius, 1781) has invaded the eastern Mediterranean Sea (Galil and Mendelson 2013).

487

References Galil BS, Clark PF (1994) A revision of the genus Matuta Weber, 1795 (Crustacea: Brachyura: Calappidae). Zoölogische Verhandelingen 294, 1–55. Galil B, Mendelson M (2013) A record of the moon crab Matuta victor (Fabricius, 1781) (Crustacea; Decapoda; Matutidae) from the Mediterranean coast of Israel. BioInvasions Records 2, 69–71. doi:10.3391/ bir.2013.2.1.12 Lai JCY, Galil BS (2007) A new species of Matuta Weber, 1795 (Crustacea, Decapoda, Matutidae) from Java, Indonesia. Raffles Bulletin of Zoology Supplement 16, 75–77. Miers EJ (1877) Notes upon oxystomatous Crustacea. Transactions of the Linnean Society of London, Series 2, Zoology 2, 235–49, pls 38–40. Perez O, Bellwood D (1988) Ontogenetic changes in the natural diet of the sandy shore crab, Matuta lunaris (Forskål) (Brachyura: Calappidae). Marine and Freshwater Research 39, 193–199. doi:10.1071/MF9880193 Romimohtarto K (1972) Five species of Matuta (Calappidae, Brachyura, Decapoda) from Indonesia. Marine Research in Indonesia 12, 3–23. doi:10.14203/mri.v12i0.338

Cancroidea Latreille, 1802 Cancroids include the archetypal crab of the genus Cancer. All have simple walking legs and an ovate carapace, but differ from many superfamilies in having the antennae folding longitudinally. Cancridae and Atelecyclus were placed in the one superfamily largely on the basis of shared features of the thoracic sternum (Guinot et  al. 2008). Two genera now comprise Atelecyclidae (Ng et  al. 2008). The two families are separated on the level of setation of the antennal flagellum. Cancrids are free-ranging crabs on rocky coasts while atelecyclids are burrowers on soft sediments. Diagnosis. Carapace semicircular to broadly oval; front divided into 3 or more angular lobes or spines. Antennule folding longitudinally or almost so. Buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular. Cheliped dactylus occlusal margin with teeth of similar size. Male pleonites 3–5 fused; male telson semicircular or elongated, pointed. Gonopod 1 stout.

Key to families of Cancroidea 1. –

Carapace subcircular to longitudinally ovate. Antennal flagellum densely setose, appearing ‘furry’ (Fig. 14.27d). Maxilliped 3 ischium-merus ~3 times as long as wide (Fig. 14.27e)�������������������������������������� Atelecyclidae Carapace transversely ovate. Antennal flagellum sparsely, minutely setose. Maxilliped 3 ischium-merus about twice as long as wide (Fig. 14.28q)���������������������������������������������������������������������������������������������������������������� Cancridae

References Guinot D, De Angeli A, Garassino A (2008) Marocarcinidae, a new eubrachyuran family, and Marocarcinus pasinii n. gen., n. sp. from the Upper Cretaceous (Cenomanian-Turonian) of Gara Sbaa, southeastern Morocco (Crustacea, Decapoda, Brachyura). Atti della Società

Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano 149, 25–36. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286.

488

Marine Decapod Crustacea

Fig. 14.27.  Atelecyclidae. Carapace: a, Atelecyclus rotundatus (Olivi, 1792); b, A. undecimdentatus (Herbst, 1783); c, Pseudocorystes sicarius (Poeppig, 1836). d, front, antennae, Atelecyclus. e, maxilliped 3, Atelecyclus. f, gonopod 1, Atelecyclus.

Atelecyclidae Ortmann, 1893 Figure 14.27, Plate 42a Atelecyclidae, sensu Salva and Feldmann (2001) and Ng et al. (2008), included a diverse array of mostly Indo-Pacific and western Atlantic deep-water ‘non-cancroid’ genera united with Atelecyclus largely by carapace and antennular similarities, now considered to be superficial. Tavares and Cleva (2010) transferred these ‘non-cancroid’ atelecyclids to Trichopeltarioidea. Pseudocorystes, a genus of fluid taxonomic placement and not considered by Salva and Feldmann (2001) or Tavares and Cleva (2010) is herein considered an atelecyclid following Ng et al. (2008). Atelecyclidae is now a family of just three species of burrowing crabs, restricted to temperate waters of the north Atlantic and south-east Pacific (Salva and Feldmann 2001). Diagnosis. Carapace subcircular to longitudinally ovate. Eyestalk longer than wide, cornea shorter than half length of peduncle. Antenna flagellum densely setose, appearing ‘furry’. Maxilliped 3 merus elongate, much longer than wide, almost as long as ischium.

median tooth triangular, distinctly longer and wider than lateral teeth (Fig. 14.27c). Maximum cl. 52 mm. Intertidal–slope (0–269 m). Temperate South America (Pacific coast). 1 species (Rathbun 1930: diagnosis, figure). References Forest J (1958) Sur la validité et le nom des deux espèces d’Atelecyclus (Crustacea Decapoda Brachyura). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 29, 469–474. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Salva EW, Feldmann RM (2001) Reevaluation of the family Atelecyclidae (Decapoda: Brachyura). Kirtlandia 52, 9–62. Tavares M, Cleva R (2010) Trichopeltariidae (Crustacea, Decapoda, Brachyura), a new family and superfamily of eubrachyuran crabs with description of one new genus and five new species. Papéis Avulsos de Zoologia 50, 97–157. doi:10.1590/S0031-10492010000900001 Taylor AC (1984) Branchial ventilation in the burrowing crab, Atelecyclus rotundatus. Journal of the Marine Biological Association of the United Kingdom 64, 7–20. doi:10.1017/S0025315400059609

Atelecyclus Leach, 1814 Diagnosis. Carapace subcircular or almost semicircular, slightly wider than long; anterolateral margin coarsely dentate; front with 3 teeth of subequal width (Fig.  14.27a, b). Maximum cl. 50 mm (Pl. 42a). Intertidal–slope (0–300 m). Temperate Northern E Atlantic (including Mediterranean). 2 species (Forest 1958: differentiation of species, figures; Taylor 1984: burrowing behaviour, physiology).

Pseudocorystes H. Milne Edwards, 1837 Diagnosis. Carapace oval, longer than wide; anterolateral margin with 2 small teeth, otherwise smooth, evenly curved; front with

Cancridae Latreille, 1802 Figure 14.28, Plate 42b–g The genus name Cancer was introduced by Linnaeus (1758) for all species of decapods but since then the name became restricted to a small group of species related to the European edible crab or brown crab, C. pagurus Linnaeus, 1758. This species is the target of a pot fishing industry and frequently seen for sale in European and British markets. Cancrid crabs have a robust carapace and chelipeds and are common in the fossil record. For this reason cancrid taxonomy has attracted the wide attention of palaeontologists

14 – Brachyura – crabs

(Nations 1975; Schram and Ng 2012; Schweitzer and ­Feldmann 2000). Their diagnoses and keys are the basis of those here. The key to species of Cancer from the Americas by Rathbun (1930) is useful to distinguish species but some she dealt with are now junior synonyms and others are now in other genera. The origin of the family is disputed, in the North Pacific according to Harrison and Crespi (1999) or the Southern Hemisphere according to Schweitzer and F ­ eldmann (2000).

489

Diagnosis. Carapace transversely ovate. Eyestalk retracted into orbit, stout. Antenna flagellum sparsely, minutely setose (Fig. 14.28f). Maxilliped 3 merus subquadrate, shorter than ischium. Implicit generic attributes. Carapace oblong, antero- and posterolateral margins separated by tooth or rounded angle. Carapace margins and pereopods with scattered setae, never obscuring margins and/or surfaces. Chela outer surface not setose. Male telson longer than wide, acute.

Key to genera of Cancridae 1. – 2. – 3. – 4. – 5.



Carapace roundish, antero- and posterolateral margins poorly separated; regions well demarcated by deep grooves or with strong granulose ornaments. Carapace margins, pereopods often prominently setose, often obscuring margins and/or surfaces (Fig. 14.28d, f)�����������������������������������������������������������������������������������������������������2 Carapace oblong, antero- and posterolateral margins separated by tooth or rounded angle; regions indistinct, not densely covered with granules or large ornaments. Carapace margins, pereopods with scattered setae, never obscuring margins and/or surfaces (Fig. 14.28a–c, e, g)����������������������������������������������������������������3 Carapace regions well defined, evenly granulated, setose, separated by deep grooves (Fig. 14.28f). Chela outer surface setose (Fig. 14.28l)���������������������������������������������������������������������������������������������������������������Platepistoma Carapace regions poorly demarcated, without prominent grooves or setae, sometimes with large granular patches (Fig. 14.28d). Chela outer surface not setose (Fig. 14.28j)�����������������������������������������������Glebocarcinus Carapace posterolateral margin with distinct tooth or tubercle on anterior one-third (Fig. 14.28b, g, arrowed)�������������������������������������������������������������������������������������������������������������������������������������������������������������4 Carapace posterolateral margin unarmed (Fig. 14.28a, c, e)�����������������������������������������������������������������������������������������������������5 Carapace smooth, regions poorly demarcated; anterior anterolateral teeth separated by fissures, posterior ones separated by prominent clefts (Fig. 14.28g). Adult chelae elongate (Fig. 14.28m)�����������������������������������Romaleon Carapace regions demarcated by patches of granules, grooves indistinct; all anterolateral teeth separated by fissures, some appearing paired (Fig. 14.28b). Adult chelae short (Fig. 14.28h)����������������������������������������������Anatolikos Male telson elongated, acute, or as long as wide, with rounded apex (Fig. 14.28o). Chela wider distally than proximally, fingers twice as long as basal width, cutting edges with 8–10 teeth (Fig. 14.28k). Carapace anterolateral teeth distinct, bluntly pointed, sharp, or sometimes spiniform; separated by short fissures or clefts, first tooth narrower than second (Fig. 14.28a, e)��������������������������������������������������������������������������������Metacarcinus Male telson longer than wide, acute (Fig. 14.28n). Chela of similar width throughout length, fingers little longer than basal width, cutting edges with 2–4 blunt teeth (Fig. 14.28i). Carapace anterolateral teeth low, lobiform, separated by deep fissures, first tooth wider than second (Fig. 14.28c)����������������������������������������� Cancer Anatolikos Schweitzer & Feldmann, 2000

Diagnosis. Carapace regions demarcated by patches of granules, grooves indistinct; anterolateral margin with all teeth separated by fissures, some appearing paired; posterolateral margin with distinct tooth or tubercle on anterior third. Adult chelae short. Maximum cl. 54 mm (Pl. 42b). Shelf (30–150 m). Temperate Northern W Pacific. 2 species (Sakai 1976: key to species; Schweitzer and Feldmann 2000).

Cancer Linnaeus, 1758 Diagnosis. Carapace regions smooth, regions poorly demarcated; anterolateral margin low, lobiform, separated by deep fissures; posterolateral margin unarmed. Chela of similar width

throughout length, fingers little longer than basal width, cutting edges with 2–4 teeth. Male telson longer than wide, acute. Maximum cl. 150 mm (Pl. 42c). Intertidal–slope (0–800 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Tropical Eastern Pacific, Temperate South America, Temperate Southern Africa. 8 species (Rathbun 1930: key to American species; Schram and Ng 2012: rediagnosis).

Glebocarcinus Nations, 1975 Diagnosis. Carapace roundish, antero- and posterolateral margins poorly separated; regions poorly demarcated, without prominent grooves or setae, sometimes with large granular patches; anterolateral margin with 9 teeth separate to bases, directed forward, sharp,

490

Marine Decapod Crustacea

Fig. 14.28.  Cancridae. a, Metacarcinus novaezelandiae (Hombron & Jacquinot, 1846). Carapace: b Anatolikos; c, Cancer; d, Glebocarcinus; e, Metacarcinus; f, Platepistoma guezei (Crosnier, 1976); g, Romaleon. Chela: h, Anatolikos; i, Cancer; j, Glebocarcinus; k, Metacarcinus; l, Platepistoma; m, Romaleon. Male pleonites 4–6, telson: n, Cancer; o, Metacarcinus. p, gonopod 1, Platepistoma. q, maxilliped 3, Platepistoma.

with granules or spinelets, teeth 2–3, 4–5 sometimes paired. Carapace margins and pereopods prominently setose, often obscuring margins and/or surfaces. Maximum cl. 30 mm (Pl. 42d). Intertidal–slope (0–435 m). Temperate Northern Pacific, Tropical Eastern Pacific. 3 species (Marin et  al. 2018: species compared).

Metacarcinus A. Milne-Edwards, 1862 Diagnosis. Carapace regions smooth, regions poorly demarcated; anterolateral margin teeth distinct, bluntly pointed, sharp, or sometimes spiniform; separated by short fissures or clefts; posterolateral margin unarmed. Chela wider distally than proximally, fingers twice as long as basal width, cutting edges with 6–10 teeth. Maximum cl. 85 mm (Pl. 42e). Intertidal–slope (0–327 m). Temperate Northern Pacific, Tropical Eastern Pacific, Temperate South America, Temperate

Australasia. 5 species (Rathbun 1930: key to species as Cancer). Metacarcinus novaezelandiae (Hombron & Jacquinot, 1846), popularly known as the pie-crust crab (Fig. 14.28a) is native to New Zealand but has become established in south-eastern Australia (Poore 2004). It is not a typical species of Metacarcinus and way outside the biogeographic range of other species and of species of Cancer, which it also somewhat resembles.

Platepistoma Rathbun, 1906 Diagnosis. Carapace roundish, antero- and posterolateral margins poorly separated; regions well defined, evenly granulated, setose, separated by deep grooves; anterolateral margin with 10 short broad teeth separated to bases, last smaller. Carapace margins and pereopods prominently setose, often obscuring margins and/or surfaces. Chela outer surface setose. Maximum cl. 51 mm (Pl. 42f).

14 – Brachyura – crabs

Shelf, slope (50–720 m). Temperate Northern Pacific, IndoWest Pacific, Temperate Southern Africa, Temperate Australasia. 8 species (Crosnier 1976: figure; Davie 1991: key to 7 species; Davie and Ng 2012).

Schweitzer CE, Feldmann RM (2000) Re-evaluation of the Cancridae Latreille, 1802 (Decapoda: Brachyura) including three new genera and three new species. Contributions to Zoology 69, 223–250. doi:10.1163/18759866-06904002

Carpilioidea Ortmann, 1893

Romaleon Gistel, 1848 Diagnosis. Carapace regions smooth, regions poorly demarcated; anterolateral margin with anterior teeth separated by fissures, posterior ones separated by prominent clefts; posterolateral margin with distinct tooth or tubercle on anterior third. Adult chelae elongate. Maximum cl. 80 mm (Pl. 42g). Intertidal–slope (0–280 m). Temperate Northern Pacific, Tropical Eastern Pacific, Temperate South America. 7 species (Rathbun 1930: key as species of Cancer; Schweitzer and Feldmann 2000: rediagnosis). Romaleon setosum (Molina, 1782) is the basis of an artisanal fishery in southern Chile (Fischer and Thatje 2016). References Crosnier A (1976) Données sur les Crustacés Décapodes capturés par M. Paul Guézé a l’île de La Réunion lors d’essais de pêche en eau profonde. In Biologie marine et exploitation des resources de l’océan Indien occidental. Travaux et Documents ORSTOM. pp. 225–254. Davie PJF (1991) Crustacea Decapoda: the genus Platepistoma Rathbun, 1906 (Cancridae) with the description of three new species. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 9. Mémoires du Muséum National d’Histoire Naturelle, Paris 152, 493–514. Davie PJF, Ng PKL (2012) A new species of Platepistoma Rathbun, 1906 (Decapoda: Brachyura: Cancridae) from deep water off South Africa. Zootaxa 3522, 73–80. doi:10.11646/zootaxa.3522.1.5 Fischer S, Thatje S (2016) Temperature effects on life-history traits cause challenges to the management of brachyuran crab fisheries in the Humboldt Current: A review. Fisheries Research 183, 461–468. doi:10.1016/j.fishres.2016.07.008 Harrison MK, Crespi BJ (1999) Phylogenetics of Cancer crabs (Crustacea: Decapoda: Brachyura). Molecular Phylogenetics and Evolution 12, 186–199. doi:10.1006/mpev.1998.0608 Linnaeus C (1758) Systema Naturae per Regna tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio Decima, reformata. Vol. 1 Animalia. 10th edn. Laurentii Salvii, Holmiae. Marin IN, Maiorova AS, Korn OM (2018) Cryptic diversity of the rocky crab genus Glebocarcinus Nations, 1975 (Crustacea: Decapoda: Cancridae): description of a new species from Russian coastal waters of the Sea of Japan based on morphology, DNA and distribution. Zootaxa 4415, 473–497. doi:10.11646/zootaxa.4415.3.4 Nations D (1975) The genus Cancer (Crustacea: Brachyura): systematics, biogeography and the fossil record. Natural History Museum of Los Angeles County. Science Series 23, 1–104. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo. Schram FR, Ng PKL (2012) What is Cancer? Journal of Crustacean Biology 32, 665–672. doi:10.1163/193724012X640650

491

The superfamily contains only three species all in one family and one genus. They are large brightly coloured crabs living in tropical coral reefs and adjacent habitats. Their large size and distinctive coloration attracted early biologists and all species were described in the 18th century. Carpilius has long been considered as one of the many xanthoid-like genera (Guinot 1968) and molecular studies of relationships find the genus well embedded in this clade, probably nearest to the eriphioids (Lai et al. 2014; Wetzer et al. 2003). The genus is distinguished from xanthoids by the combination of fused male pleonites 3 and 4 (Fig. 14.29b) and the long coiled gonopod 2 (Fig. 14.29c, d). Diagnosis. Carapace transversely ovate, prominently convex, smooth or finely rugose, without spines, almost glabrous; anterolateral margin evenly curved, terminating in rounded lobe. Buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular. Cheliped surfaces and margins smooth, without spines; fixed finger of larger chela with massive molariform tooth proximally on occlusal margin. Male pleonites 3–4, immovably fused, 5 partially movable. Gonopod 1 stout. Gonopod 2 distally looped, when straightened, exceeding gonopod 1 length.

Carpiliidae Ortmann, 1893 Figure 14.29, Plate 42h Carpilius Desmarest, 1823 Maximum cl. 115 mm (Pl. 42h). Intertidal, subtidal (0–50 m; associated with coral reefs). Tropical W Atlantic, Temperate Northern W Pacific, Indo-West Pacific. 3 species. (Crosnier 1984: keys to 2 Indo-West Pacifc species; Sakai 1976). Carpilius corallinus (Herbst, 1783) is a Caribbean species (Chace et al. 1986). References Chace FA, McDermott JJ, McLaughlin PA, Manning RB (1986) Order Decapoda (shrimps, lobsters and crabs). In Marine Fauna and Flora of Bermuda. (Ed. Sterrer W) pp. 312–358. Wiley-Interscience, New York. Crosnier A (1984) Addendum: Familles de Carpiliidae et des Menippidae. In: Serène, R. Crustacés Décapodes Brachyoures de l’Océan Indien Occidental et de la Mer Rouge, Xanthoidea: Xanthidae et Trapeziidae. Faune Tropicale 24, 299–313. Guinot D (1968) Recherches préliminaires sur les groupements naturels chez les crustacés décapodes brachyoures. VI. Les Carpilinae. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 40, 320–334. Lai JCY, Thoma BP, Clark PF, Felder DL, Ng PKL (2014) Phylogeny of eriphioid crabs (Brachyura, Eriphioidea) inferred from molecular and

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Marine Decapod Crustacea

Fig. 14.29.  Carpiliidae. a, Carpilius maculatus (Linnaeus, 1758). Carpilius: b, male pleon (pleonites 3–4 fused); c, d, gonopods 1, 2. morphological studies. Zoologica Scripta 43, 52–64. doi:10.1111/ zsc.12030 Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo. Wetzer R, Martin JW, Trautwein SE (2003) Phylogenetic relationships within the coral crab genus Carpilius (Brachyura, Xanthoidea, Carpiliidae) and of the Carpiliidae to other xanthoid crab families based on molecular sequence data. Molecular Phylogenetics and Evolution 27, 410–421. doi:10.1016/S1055-7903(03)00021-6

Intertidal–slope (0–400 m). Temperate Northern Pacific. 1 species (Armetta and Stevens 1987: biology, fishery; Azuma et al. 2008: genetic variation).

Cheiragonoidea Ortmann, 1893

Diagnosis. Carapace broader than long, pentagonal; lateral margins with compound teeth. Ambulatory legs without spines (Fig. 14.30b). Maximum cl. 83 mm. Intertidal–shelf (0–180 m). Temperate Northern Pacific. 2 species (keys to species: Rathbun 1930; Sakai 1976).

The three species of cheiragonoids are crabs from muddy sediments in the North Pacific. All are large enough to be fished commercially, especially in Asia. For this reason, there has been considerable research on their biology and fishery. The taxonomy of the two genera and their species has been long understood (Benedict 1892) but their family placement has moved from Corystidae to Atelecylidae (Rathbun 1930; Sakai 1976) to Cheiragonidae (Salva and Feldmann 2001). All these authors diagnose the genera in more detail than below. The type genus, Cheiragonus Brandt, 1851, from which the family name is derived, is a junior synonym of Telmessus. Diagnosis. Carapace subhexagonal to longitudinally subrectangular. Antennule folding longitudinally or almost so. Antennae shorter than half carapace length. Buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular. Male pleonites 1–6 and telson free; male telson elongated, pointed. Female vulva exposed, not concealed by pleon. Gonopod 1 stout.

Cheiragonidae Ortmann, 1893 Figure 14.30, Plate 42i Erimacrus Benedict, 1892 hair crab Diagnosis. Carapace longer than broad, elongate oval; lateral margins arcuate, with 7 teeth. Ambulatory legs spinous (Fig. 14.30a). Maximum cl. 110 mm (Pl. 42i).

Telmessus White, 1846 helmet crab

References Armetta TM, Stevens BG (1987) Aspects of the biology of the hair crab, Erimacrus isenbeckii, in the eastern Bering Sea. Fishery Bulletin 85, 523–544. Azuma N, Kunihiro Y, Sasaki J, Mihara E, Mihara Y, et al. (2008) Genetic variation and population structure of hair crab (Erimacrus isenbeckii) in Japan inferred from mitochondrial DNA sequence analysis. Marine Biotechnology 10, 39–48. doi:10.1007/s10126-007-9033-1 Benedict JE (1892) Corystoid crabs of the genera Telessemus and Erimacrus. Proceedings of the United States National Museum 15, 223–230, pls 25–27. Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo. Salva EW, Feldmann RM (2001) Reevaluation of the family Atelecyclidae (Decapoda: Brachyura). Kirtlandia 52, 9–62.

Corystoidea Samouelle, 1819 masking crabs Corystoids formerly included a suite of elongated burrowing crabs that hold their long, setose antennae together to form a snorkel, allowing them to breathe whilst fully buried under the sand (Zimmermann 1912). Several of these have

14 – Brachyura – crabs

493

Fig. 14.30.  Cheiragonidae. a, Erimacrus isenbeckii (Brandt, 1848); b, Telmessus cheiragonus (Tilesius, 1815). c, antennule, antenna, orbit, Telmessus. d, male pleon, Telmessus. e, female pleon, vulvae on sternum, Telmessus.

been moved to other families: Nautilocorystes to Nautilocorystidae (Portunoidea), Podocatactes to Trichopeltariidae (Trichopeltarioidea) and Pseudocorystes to Atelecyclidae (Cancroidea). The remaining three genera, constituting the single family and superfamily, form a group closely related to the cancroids. Corystes is confined to the North-east Atlantic while the other two genera have an overlapping distribution in the Western Pacific and may be easily confused with each other (Ng et al. 2000).

Diagnosis. Carapace longitudinally ovate, longer than wide, convex, regions poorly defined. Antennule folding longitudinally or almost so. Antenna as long as or longer than carapace. Buccal cavity approximately quadrate. Male pleonites 3–5 fused. Gonopod 1 stout.

Corystidae Samouelle, 1819 Figure 14.31, Plate 42j–l

Key to genera of Corystidae 1. – 2. –

Carapace anterolateral margin with 1 sharp spine, posterolateral margin with 2 small well spaced spines (Fig. 14.31a)���������������������������������������������������������������������������������������������������������������������������������������������������������������������Corystes Carapace anterolateral and posterolateral margins with 9 or more spines�����������������������������������������������������������������������������2 Carapace widest near midlength, strongly convex; anterolateral margin with 3 small subequal teeth, posterolateral margin with larger teeth diminishing posteriorly (Fig. 14.31b)���������������������������������������������������� Gomeza Carapace widest in anterior half, gently convex; anterolateral margin with 3 large teeth, posterolateral margin with smaller teeth (Fig. 14.31c)��������������������������������������������������������������������������������������������������������������������������� Jonas Corystes Bosc, 1801

Gomeza Gray, 1831

Diagnosis. Carapace oval, 1.3 times as long as wide; anterolateral margin with 1 sharp spine, posterolateral margin with 2 small well spaced spines. Male chelipeds twice as long as carapace. Pereopod 5 dactylus slender. Maximum cl. 40 mm (Pl. 42j). Intertidal–shelf (0–115 m). Temperate Northern Atlantic (E Atlantic, Baltic, Mediterranean). 1 species (Hartnoll 1972: distribution, biology).

Diagnosis. Carapace widest near midlength, strongly convex; anterolateral margin with 3 small subequal teeth, posterolateral margin with larger teeth diminishing posteriorly. Male chelipeds about as long as carapace. Pereopod 5 dactylus slender, styliform. Maximum cl. 21 mm (Pl. 42k). Intertidal–shelf (0–123 m). Temperate Northern Atlantic, Western and Central Indo-Pacific, Temperate South America

494

Marine Decapod Crustacea

Fig. 14.31.  Corystidae. a, Corystes cassivelaunus (Pennant, 1777); b, Gomeza bicornis Gray, 1831. Jonas,: c, carapace; d, pereopod 4.

(Pacific coast), Temperate Australasia. 2 species (Poore 2004: Indo-West Pacific species; Rathbun 1930: eastern S American species).

Jonas Hombron & Jacquinot, 1846 Diagnosis. Carapace widest in anterior half, gently convex; anterolateral margin with 3 large teeth, posterolateral margin with smaller teeth. Male chelipeds about as long as carapace. Pereopod 5 dactylus spatuliform (Fig. 14.31d). Maximum cl. 41 mm (Pl. 42l). Shelf (20–90 m). Temperate Northern Pacific, Western and Central Indo-Pacific. 7 species (Barathkumar et al. 2016: species compared, distribution; Ng et al. 2000: 3 species compared). References Barathkumar S, Das NP, Satpathy KK (2016) A new species of sand crab Jonas Hombron & Jacquinot, 1846 (Crustacea: Decapoda: Brachyura: Corystidae) from the southeastern coast of India. Zootaxa 4079, 480–486. doi:10.11646/zootaxa.4079.4.7 Hartnoll RG (1972) The biology of the burrowing crab, Corystes cassivelaunus. Bijdragen tot de Dierkunde 42, 139–155. doi:10.1163/​2666064404202003 Ng PKL, Chan T-Y, Wang C-H (2000) The crabs of the families Dromiidae, Raninidae, Corystidae and Palicidae (Crustacea: Decapoda: Brachyura) of Taiwan. In: Proceedings of the International Symposium on Marine Biology in Taiwan – Crustacea and zooplankton taxonomy, ecology and living resources, 26–27 May, 1998, Taiwan. (Eds Hwang J-S, Wang C-H, Chan T-Y) pp. 155–180. National Taiwan Museum, Taipei. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne.

Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Zimmermann KE (1912) Notes on the respiratory mechanism of Corystes cassivelaunus. Journal of the Marine Biological Association of the United Kingdom 9, 288–291. doi:10.1017/S0025315400048323

Dairoidea Serène, 1984 In addition to Dairidae, the superfamily has variously also included Dairoididae and Dacryopilumnidae (Ng et  al. 2008; Števčić 2005), but the latter two families have since been shown to have parthenopoid and eriphioid affinities, respectively (Lai et  al. 2014). As such, Dairoidea currently includes a single family, Dairidae, and its type genus, Daira. Dairoids are xanthid-like but have unusual cuticular ornamentation, presumably convergent on that of dairoidids, in which the carapace and cheliped surfaces are covered with perforated, interlocking, wart-like, tubercles that form a pavement-like surface (Guinot 1967). Diagnosis. Carapace and chelipeds covered in large, nodular, wart-like, perforated tubercles, often margined with tufts of short setae. Carapace transversely ovate; antennule folding obliquely; endostomial ridges present; buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular. Male pleonites 3–5 fused, sutures visible. Gonopod 1 stout. Gonopod 2 as long as gonopod 1, distal portion strongly recurved, even looped.

14 – Brachyura – crabs

Dairidae Serène, 1984 Figure 14.32, Plate 43a Daira De Haan, 1833 Maximum cl. 34 mm (Pl. 43a). Intertidal, subtidal (0–10 m; coral reefs). Indo-West Pacific. Mediterranean Sea (introduced). 2 species, Daira perlata (Herbst, 1790) in the Indo-West Pacific and Mediterranean Sea, and D. americana Stimpson, 1860, from the eastern Pacific (Milne-Edwards 1865). The crab is toxic to humans (Llewellyn 2001). References Guinot D (1967) Recherches préliminaires sur les groupements naturels chez les Crustacés décapodes brachyoures. III. A propos des affinités des genre Dairoides Stebbing et Daira De Haan. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 39, 540–563. Lai JCY, Thoma BP, Clark PF, Felder DL, Ng PKL (2014) Phylogeny of eriphioid crabs (Brachyura, Eriphioidea) inferred from molecular and morphological studies. Zoologica Scripta 43, 52–64. doi:10.1111/zsc.12030 Llewellyn LE (2001) Human fatalities in Vanuatu after eating a crab (Daira perlata). Medical Journal of Australia 175, 343–344. doi:10.5694/j.1326-5377.2001.tb143612.x Milne-Edwards A (1865) Études zoologiques sur les Crustacés Récents de la famille des Cancériens. Nouvelles Archives du Muséum d’Histoire naturelle, Paris 1, 179–308. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159.

495

Dorippoidea MacLeay, 1838 carrier crabs The superfamily comprises two families (13 genera) of crabs renowned for carrying objects such as bivalve shells, sea urchins, sea anemones and plant leaves with the last two pairs of specialised legs. These legs are either subchelate or hooked to grasp their chosen object. Separation of two groups, as subfamilies of Dorippidae or as separate families, long recognised on morphological grounds (Ng et al. 2008) has been confirmed using mitochondrial genes (Sin et al. 2009). The two families are ecologically segregated, with most dorippids living in shallow water and most ethusids in waters deeper than 200 m. Dorippoids and cyclodorripoids, especially Cymonomidae, have a similar habitus and were classified together until separated on the basis of differences in reproductive morphology (Garth and Haig 1971). Although the phylogenetic position of the dorippoids is yet to be settled (Tsang et  al. 2014), some results place dorippoids close to the base of the Eubrachyura, suggesting carrying behaviour may have been inherited from ‘lower’ clades such as cyclodorippoids (Ahyong et al. 2007). Diagnosis. Carapace as long as or slightly wider than long, subquadrate to rounded, often widest posteriorly. Antennule folding obliquely. Buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular. Pereopods 4–5 inserted obliquely on body, held dorsally. Pleonites 1, 2 not folded ventrally, visible from dorsal view.

Key to families of Dorippoidea 1. –

Pereopods 4, 5 dactyli hook-like (Fig. 14.34j). Inhalent branchial opening circular or oval, placed against bases of cheliped. Maxilliped 3 epipod completely occluding inhalent branchial opening, exopod with flagellum����������������������������������������������������������������������������������������������������������������������������������������������������������������������� Ethusidae Pereopods 4, 5 dactyli subchelate (Fig. 14.33e, g, i). Inhalent branchial opening elongate, adjacent to and separated from cheliped by carapace wall. Maxilliped 3 epipod not occluding inhalent branchial opening, exopod without flagellum���������������������������������������������������������������������������������������������������������������������� Dorippidae

Fig. 14.32.  Dairidae, Daira perlata (Herbst, 1790). a, habitus; b, front, antennae, antennae, orbit; c, d, gonopods 1, 2.

496

Marine Decapod Crustacea

References Ahyong ST, Lai JCY, Sharkey D, Colgan DJ, Ng PKL (2007) Phylogenetics of the brachyuran crabs (Crustacea: Decapoda): the status of Podotremata based on small subunit nuclear ribosomal RNA. Molecular Phylogenetics and Evolution 45, 576–586. doi:10.1016/j.ympev.2007.03.022 Garth JS, Haig J (1971) Decapod crustacea (Anomura and Brachyura) of the Peru-Chile Trench. Anton Bruun Report 6, 3–20. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Sin YW, Lai JCY, Ng PKL, Chu KH (2009) Phylogeny of Dorippoidea (Crustacea: Decapoda: Brachyura) inferred from three mitochondrial genes. Invertebrate Systematics 23, 223–230. doi:10.1071/IS09020 Tsang LM, Schubart CD, Ahyong ST, Lai JCY, Au EYC, et al. (2014) Evolutionary history of true crabs (Crustacea: Decapoda: Brachyura) and the origin of freshwater crabs. Molecular Biology and Evolution 31, 1173–1187. doi:10.1093/molbev/msu068

Dorippidae MacLeay, 1838 carrier crabs Figure 14.33, Plate 43b–d Distinguishing features of dorippids are the thinner, shorter, last two pairs of legs, held dorsally and with hooked terminal articles. These enable the crabs to carry objects over the carapace, hence the common name, carrier crabs. The objects carried include sponges, bivalve shells, sea anemones, medusae and other fragments (Guinot and Wicksten 2015).

Species of Neodorippe, the leaf-porter crabs, are perhaps unique in carrying plant leaves, and swim upside down near the water surface (Ng and Rahayu 2002). Most species are tropical or subtropical, usually on soft sediments from the intertidal to the edge of the continental shelf. The taxonomy of the family was substantially dealt with in two major papers (Holthuis and Manning 1990; Manning and Holthuis 1981). These works deal in detail with the East Atlantic and Indo-West Pacific genera and species respectively. Only three species have been described since (Manning 1993; Ng and Rahayu 2002). Heikegani, Heikeopsis japonica (von Siebold, 1824), is a dorippid reknowned in Japanese folklore. The carapace displays a pattern resembling a human face. According to legend, Heikegani contain the souls of slain Heike samurai warriors. Diagnosis. Opening for inhalent respiratory current narrow, elongate, adjacent to and separated from cheliped by carapace wall. Maxilliped 3 epipod not occluding inhalent branchial opening, exopod without flagellum. Pereopods 4, 5 subchelate, dactylus occluding with tubercle on propodus. Implicit generic attributes. Front extending to or slightly beyond exorbital teeth; carapace lateral margin without branchial tubercle; inner suborbital tooth smaller than exorbital tooth; orbital margin unarmed. Pleonites of male and female without sublateral tubercles. Pereopods dactyli lacking fringes, naked or nearly so; pereopod 2, 3 meri without spines.

Key to genera of Dorippidae 1. – 2. – 3. – 4. – 5. – 6. – 7.

Carapace lateral margin with branchial tubercle or spine (Fig. 14.33a, d, i)��������������������������������������������������������������������������2 Carapace lateral margin without branchial tubercle or spine (Fig. 14.33b, c, e–h)����������������������������������������������������������������4 Carapace cardiac region smooth. Pereopods 2, 3 dactyli with fringe of setae on basal half of extensor margin (Fig. 14.33i)���������������������������������������������������������������������������������������������������������������������������������������������� Phyllodorippe Carapace cardiac region with V-shaped median ridge (Fig. 14.33a, d). Pereopods 2, 3 dactyli lacking fringes, naked or nearly so���������������������������������������������������������������������������������������������������������������������������������������������������������3 Carapace with small tubercles (Fig. 14.33d). Pereopods 2, 3 meri extensor margin with row of spines (Fig. 14.33m). Pleonites of male and female without sublateral tubercles���������������������������������������������������������Medorippe Carapace rough, with deep grooves and tubercles (Fig. 14.33a). Pereopods 2, 3 meri without spines (Fig. 14.33l). Pleonites 2–4 of male, 3 and 4 of female with sublateral tubercles (Fig. 14.33n)��������������������������� Dorippe Pereopods 3, 4 dactyli lacking fringes, naked or nearly so. Carapace wider than long (Fig. 14.33b, g, h)�������������������������5 Pereopods 3, 4 dactyli with fringes of setae on extensor and flexor margins. Carapace longer than wide or barely wider than long (Fig. 14.33c, e, f)����������������������������������������������������������������������������������������������������������������������������������7 Carapace with inner suborbital tooth smaller than exorbital tooth (Fig. 14.33j)����������������������������������������������� Paradorippe Carapace with inner suborbital tooth strong, about as large as exorbital tooth (Fig. 14.33k)����������������������������������������������6 Carapace cardiac region with X-shaped granular ridge (Fig. 14.33h). Male pleonite 3 with sublateral lobes visible in dorsal view (Fig. 14.33p)��������������������������������������������������������������������������������������Philippidorippe Carapace cardiac region smooth (Fig. 14.33b). Male pleonite 3 most prominent medially in dorsal view (Fig. 14.33o)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Dorippoides Carapace usually longer than wide; front reaching well beyond exorbital teeth. Pereopods 2, 3 propodi, dactyli with fringe of long setae along extensor and flexor margins (Fig. 14.33e)���������������������������������������� Neodorippe

14 – Brachyura – crabs

497

Fig. 14.33.  Dorippidae. Carapace/habitus: a, Dorippe; b, Dorippoides; c, Heikeopsis; d, Medorippe lanata (Linnaeus, 1767); e, Neodorippe; f, Nobilum; g, Paradorippe australiensis (Miers, 1884); h, Philippidorippe armata (Miers, 1881); i, Phyllodorippe . Right orbit, exorbital, suborbital teeth (ventral view): j, Paradorippe; k, Philippidorippe. Pereopod 2: l, Dorippe; m, Medorippe. Male pleonites 1–3 (dorsal view): n, Dorippe; o, Dorippoides; p, Philippidorippe.

– 8. –

Carapace usually slightly wider than long (Fig. 14.33c, f); front extending to or beyond exorbital teeth. Pereopods 2, 3 dactyli with fringe of short setae along extensor and flexor margins������������������������������������������������������8 Carapace gastric region with 2 pairs plus 1 median tubercles; orbital margin with low tooth lateral to fissure (Fig. 14.33f)�������������������������������������������������������������������������������������������������������������������������������������������������������� Nobilum Carapace mesogastric region prominent; orbital margin unarmed (Fig. 14.33c)������������������������������������������������� Heikeopsis Dorippe Weber, 1795

Diagnosis. Carapace wider than long, or as long as wide, rough, uneven, with deep grooves and distinct tubercles; cardiac region with V-shaped median ridge; lateral margin with branchial tubercle; inner suborbital tooth strong, about as large as exorbital tooth. Pleonites 2–4 of male, 3 and 4 of female with sublateral tubercles. Maximum cl. 42 mm (Pl. 43b). Intertidal–shelf (0–120 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia. 7

species (Galil 2005: invasion into Mediterranean; Holthuis and Manning 1990: key to 5 species; Manning 1993).

Dorippoides Serène & Romimohtarto, 1969 Diagnosis. Carapace wider than long, smooth; inner suborbital tooth strong, about as large as exorbital tooth. Maximum cl. 37 mm (Pl. 43c). Subtidal, shelf (2–80 m). Western and Central Indo-Pacific. 2 species (Holthuis and Manning 1990: key to species).

498

Marine Decapod Crustacea

Heikeopsis Ng, Guinot & Davie, 2008

Phyllodorippe Manning & Holthuis, 1981

Diagnosis. Carapace wider than long, or as long as wide, moderately sculptured; cardiac region smooth; mesogastric region prominent. Pereopods 2, 3, propodi, dactyli with fringes of setae on extensor and flexor margins. Maximum cl. 30 mm (Pl. 43d). Subtidal, shelf (15–90 m). Temperate Northern W Pacific, Central Indo-Pacific. 2 species (Holthuis and Manning 1990: key to species as Heikea).

Diagnosis. Carapace wider than long, smooth; lateral margin with branchial tubercle. Pereopods 2, 3 dactyli with fringe of setae on basal half of extensor margin. Maximum cl. 25 mm. Subtidal, shelf (7–77 m). Tropical W Atlantic. 1 species (Manning and Holthuis 1981).

Medorippe Manning & Holthuis, 1981 Diagnosis. Carapace wider than long, with small tubercles; cardiac region with V-shaped median ridge; lateral margin with branchial tubercle; inner suborbital tooth strong, about as large as exorbital tooth. Pereopod 2, 3 meri extensor margin with row of spines. Maximum cl. 28 mm. Shelf (25–100 m). Temperate Northern and Tropical Atlantic, Western Indo-Pacific, Temperate Southern E Africa. 2 species (Holthuis and Manning 1990; Manning and Holthuis 1981).

Neodorippe Serène & Romimohtarto, 1969 Diagnosis. Carapace as long as wide, or longer than wide; smooth; front reaching well beyond exorbital teeth. Pereopods 2, 3, propodi, dactyli with fringes of setae on extensor and flexor margins. Maximum cl. 15 mm. Intertidal, shelf (0–46 m). Western and Central Indo-Pacific. 2 species (Holthuis and Manning 1990; Ng and Rahayu 2002).

Nobilum Serène & Romimohtarto, 1969 Diagnosis. Carapace as long as wide, deeply sculptured, grooves distinct; gastric region with 2 pairs plus 1 median tubercles; orbital margin with tooth lateral to fissure. Pereopods 2, 3, propodi, dactyli with fringes of setae on extensor and flexor margins. Maximum cl. 23 mm. Intertidal–shelf (0–46 m). Central Indo-Pacific. 1 species (Holthuis and Manning 1990).

Paradorippe Serène & Romimohtarto, 1969 Diagnosis. Carapace wider than long, smooth, or finely granular. Maximum cl. 33 mm. Intertidal–shelf (0–154 m). Temperate Northern Atlantic, Western and Central Indo-Pacific, Temperate Australasia. 4 species (Holthuis and Manning 1990: key to species).

Philippidorippe Chen, 1986 Diagnosis. Carapace wider than long, finely granular; cardiac region with X-shaped granular ridge; inner suborbital tooth strong, about as large as exorbital tooth. Pleonite 3 of male with rounded, sublateral lobes visible in dorsal view. Maximum cl. 30 mm. Shelf (96–200 m). Western and Central Indo-Pacific. 1 species (Holthuis and Manning 1990).

References Galil BS (2005) Dorippe quadridens (Decapoda, Dorippidae): first record of an alien dorippid in the Mediterranean Sea. Crustaceana 78, 497–498. doi:10.1163/1568540054473576 Guinot D, Wicksten MK (2015) Chapter 71–11. Camouflage: carrying behaviour, decoration behaviour, and other modalities of concealment in Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 583–638. Brill, Leiden. Holthuis LB, Manning RB (1990) Crabs of the subfamily Dorippinae MacLeay, 1838, from the Indo-West Pacific region (Crustacea: Decapoda: Dorippidae). Crustacean Research Special Number 3, 1–151. Manning RB (1993) Two new dorippid crabs from Australia (Crustacea; Decapoda: Dorippidae). Records of the Australian Museum 45, 1–4. doi:10.3853/j.0067-1975.45.1993.124 Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Monod T (1933) Sur quelques crustacés de l’Afrique occidentale (liste des décapodes mauritaniens et des Xanthides ouest-africains). Bulletin du Comité d’Études Historiques et Scientifiques de I’Afrique Occidentale Française 15, 456–548. Monod T (1956) Hippidea et Brachyura ouest-africains. Mémoires de I’Institut Français d’Afrique Noire 45, 1–674. Ng PKL, Rahayu DL (2002) A new species of leaf porter crab of the genus Neodorippe Serène & Romimohtarto, 1969 (Crustacea, Decapoda, Brachyura, Dorippidae) from Irian Jaya, Indonesia. Proceedings of the Biological Society of Washington 115, 754–759.

Ethusidae Guinot, 1977 Figure 14.34, Plate 43e–g Ethusids are small crabs adapted to carry a bivalve shell, piece of wood or other similar object on their back with the help of the hooked dactyli of their last two pairs of pereopods. Most inhabit deep water, rarely shallower than 100 m and as deep as 5046 m in the case of Ethusina abyssicola Smith, 1884, in the Caribbean Sea, the deepest confirmed record for a brachyuran crab (Gore 1983). The genera largely segregate bathymetrically with species of Serpenthusa, Parethusa and most species of Ethusa occurring at slope depths above 500 m, and species of Ethusina largely between 500 and 1000 m (Castro 2005). Most of more than one hundred species occur in the Indo-West Pacific region but are rarely captured. These were well documented by Castro (2005) who outlined ethusid anatomy, diagnosed three genera and provided keys to all Indo-West Pacific species then known. Few species

14 – Brachyura – crabs

(Ahyong 2008; Castro 2013; Komatsu and Komai 2009) and one genus have been described since (Naruse et  al. 2009). For the species of the Eastern Pacific see Hendrickx (1989) and for the Eastern Atlantic see Manning and Holthuis (1981). Diagnosis. Opening for inhalent respiratory current semioval or semi-circular, placed against base of cheliped. Maxilliped 3 epipod completely occluding inhalent branchial

499

opening, exopod with flagellum. Pereopods 4, 5 simple, dactylus not occluding with propodus. Implicit generic attributes. Inter-antennular septum formed of converging lateral margins of median frontal teeth. Eyestalks short, stout. Antennular article 1 enlarged, swollen, adpressing eyestalk. Pereopods 2, 3 slender, long, pereopod 2 merus extending over frontal teeth when legs folded.

Key to genera of Ethusidae 1. – 2. – 3. –

Antennular article 1 not reaching eyestalk. Eyestalks slender, visible dorsally (Fig. 14.34a, f). Pereopods 2, 3 short, pereopod 2 merus not usually extending well over frontal teeth when legs folded (Fig. 14.34a)������������������������������������������������������������������������������������������������������������������������������������������������������������������������Ethusa Antennular article 1 enlarged, swollen, adpressing eyestalk (Fig. 14.34g–i). Eyestalks short, stout, not or barely visible dorsally (Fig. 14.34b–e). Pereopods 2, 3 slender, long, pereopod 2 merus extending over frontal teeth when legs folded (Fig. 14.34b)����������������������������������������������������������������������������������������������������������������������������2 Carapace granular, regions separated by grooves, with setae or tomentum (Fig. 14.34b)��������������������������������������Ethusina Carapace smooth, regions indistinct, without setae or tomentum (Fig. 14.34c–e)���������������������������������������������������������������3 Eyestalks movable, visible dorsally. Inter-antennular septum absent, antennular articles 1 contiguous (Fig. 14.34e, i)����������������������������������������������������������������������������������������������������������������������������������������������������������� Serpenthusa Eyestalks immovable, not or barely visible dorsally (Fig. 14.34c, d). Inter-antennular septum formed of converging lateral margins of median frontal teeth (Fig. 14.34h)����������������������������������������������������������������������� Parethusa Ethusa Roux, 1830

Diagnosis. Carapace granular, often tomentose, regions separated by grooves; with 4 frontal teeth. Eyestalks slender, movable, visible dorsally. Antennular article 1 not reaching eyestalk. Pereopods 2,

3 short, pereopod 3 merus not usually extending well over frontal teeth when folded. Maximum cl. 22 mm (Pl. 43e). Shelf–bathyal (15–1315 m). Cosmopolitan, except polar seas. 43 species (Castro 2005: diagnosis, key to 28 Indo-West Pacific species; Castro 2020: species redescriptions).

Fig. 14.34.  Ethusidae. Habitus: a, Ethusa; b, Ethusina. Carapace: c, Parethusa glabra Chen, 1997; d, P. glabra Castro, 2005; e, Serpenthusa brucei Naruse, Castro & Ng, 2009. Orbit, eyestalk, antennular articles 1, 2, antennal peduncle, basal articles, ventral view: f, Ethusa; g, Ethusina; h, Parethusa; i, Serpenthusa. Pereopod 5 dactylus: j, Ethusina.

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Ethusina Smith, 1884 Diagnosis. Carapace granular, often tomentose, regions usually separated by grooves; with 4 frontal teeth. Eyestalks immovable, just visible dorsally. Maximum cl. 26 mm (Pl. 43f). Slope–abyssal (360–5046 m). Cosmopolitan, except polar seas. 35 species (Castro 2005: diagnosis, key to 25 Indo-West Pacifc species; Castro 2020: species redescriptions).

Parethusa Chen, 1997 Diagnosis. Carapace smooth, regions indistinct; with 2 or 4 frontal teeth. Eyestalks immovable, just or not visible dorsally. Maximum cl. 11 mm (Pl. 43g). Slope (390–700 m). Central Indo-Pacific. 2 species (Castro 2005: diagnosis, key to species).

Serpenthusa Naruse, Castro & Ng, 2009 Diagnosis. Carapace smooth, regions indistinct; with 4 frontal lobes. Inter-antennular septum absent, antennular articles 1 contiguous. Eyestalks movable, visible dorsally. Maximum cl. 12 mm. Slope (275–290 m). Central Indo-Pacific. 1 species (Naruse et al. 2009). References Ahyong ST (2008) Deepwater crabs from seamounts and chemosynthetic habitats off eastern New Zealand (Crustacea: Decapoda: Brachyura). Zootaxa 1708, 1–72. doi:10.11646/zootaxa.1708.1.1 Castro P (2005) Crabs of the subfamily Ethusinae Guinot, 1977 (Crustacea, Decapoda, Brachyura, Dorippidae) of the Indo-West Pacific region. Zoosystema 27, 499–600. Castro P (2013) Brachyuran crabs (Crustacea, Brachyura: Crossotonotidae, Ethusidae, Euryplacidae, Goneplacidae, Latreilliidae, Palicidae, Tetraliidae, Trapeziidae) of the MAINBAZA, MIRIKI, and ATIMO VATAE expeditions to the Mozambique Channel and Madagascar. In: Ahyong ST, Chan T-Y, Corbari L and Ng PKL (Eds) Tropical Deep-Sea Benthos vol. 27. Mémoires du Muséum National d’Histoire Naturelle, Paris 2041, 437–466. Castro P (2020) Brachyuran crabs (Crustacea: Brachyura) of eleven families of Dorippoidea, Goneplacoidea, Homoloidea, Palicoidea, Pilumnoidea, and Trapezioidea from Papua New Guinea. In: Corbari L, Chan T-Y and Ahyong ST (Eds) Deep-sea crustaceans from Papua New Guinea. Tropical Deep-Sea Benthos vol. 31. Mémoires du Muséum National d’Histoire Naturelle, Paris 213, 141–205. Gore RH (1983) Notes on rare species of Munidopsis (Anomura: Galatheidae) and Ethusina (Brachyura: Dorippidae) collected by the USNS Bartlett in the Venezuela Basin, Caribbean Sea. Proceedings of the Academy of Natural Sciences of Philadelphia 135, 200–217. Hendrickx M (1989) Notes on the genus Ethusa Roux, 1828, and description of Ethusa steyaerti n. sp. (Crustacea, Decapoda, Dorippidae), from the continental shelf of the Gulf of California, Mexico. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 11, 407–423. Komatsu H, Komai T (2009) Thalassinidea, Anomura and Brachyura (Crustacea: Decapoda) from northeastern Japan collected during the “Research on Deep-sea Fauna and Pollutants off Pacific Coast of Northern Japan” Project. In: Fujita T (Ed.) Deep-sea fauna and pollutants off Pacific coast of northern Japan. National Museum of Nature and Science Monograph 39, 581–613.

Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Naruse T, Castro P, Ng PKL (2009) A new genus and new species of Ethusidae (Decapoda, Brachyura) from Vanuatu, Western Pacific. Crustaceana 82, 931–938. doi:10.1163/156854009X427450

Eriphioidea MacLeay, 1838 Eriphioids are a group of usually heavy-bodied crabs with compact, sometimes massive chelipeds, the larger having crushing teeth or a peeling tooth at the base of the dactylus. The smaller chela often has slender evenly-toothed fingers. The male pleon is narrow and all pleonites free (Ng et al. 2008). The superfamily includes almost 70 species occupying diverse habitats from intertidal rocky shores, mangrove swamps, coral reefs and the continental slope to depths below 500 m. Most are of moderate size but species of Hypothalassia and Menippe may reach 15 cm in carapace width and are of commercial interest. The superfamily and several of its families, in particular Eriphiidae, Oziidae and Menippidae, were found not to be monophyletic on preliminary molecular and larval evidence with members of other ‘xanthoid’ s.l. families nested among eriphioids (Lai et al. 2014). Some genera too were found not to be monophyletic. These authors concluded that many of the adult morphological features used to define the superfamily are convergent, reflecting common feeding strategies, or plesiomorphies. Indeed, we could not identify consistent distinctions between Oziidae and Menippidae according to previously used characters (Davie et  al. 2015; Ng et  al. 2008; Ng et  al. 2009); here we focus on aspects of thoracic sternites 3 and 4, and the anterolateral teeth on the carapace. Notably, Lai et al. (2014) found the Tasmanian Giant Crab Pseudocarcinus gigas (Lamarck, 1818) to lie well outside of Menippidae based on both molecular and larval data and, according the latter, with clear affiliations to oziids. It is now placed in its own family, Pseudocarcinidae, and superfamily, Pseudocarinoidea (Ng and Davie 2020). Based on the results of Lai et  al. (2014), Ng and Davie (2020) recognised Dacryopilumnidae as eriphioid, and the former eriphioid family, Dairoididae, as a member of the Parthenopoidea. Diagnosis. Carapace transversely ovate to hexagonal, as wide as long to wider than long; frontal width of adults onequarter carapace width or greater. Antennules folding transversely or obliquely. Antenna longer than cornea width; flagellum well developed. Cheliped palm ovate in cross-section; adult male major chela length not exceeding carapace width. Pereopods 2–5 inserted laterally on body, similar, as walking legs, dactylus slender, pointed. Mouthfield approximately quadrate. Maxilliped 3 merus pentagonal to subquadrate. Male pleonites 1–6 freely articulating;

14 – Brachyura – crabs

pleonite 3 width less than or equal to twice telson width, without transverse ridge; narrowly subrectangular or linguiform, with lateral margins of pleonites 3–6 gradually

501

converging towards telson. Gonopod 1 straight, not sigmoid. Gonopod 2 shorter than to longer than gonopod 1, straight except for curved or looped distal end.

Key to families of Eriphioidea 1. – 2.

– 3. – 4. – 5. –

Carapace with orbits completely closed, antenna widely separated from and excluded from inner margin of orbit; orbits widely separated, usually placed close lateral edges of anterior margin (Fig. 14.36c, d)���������������������������2 Carapace with orbits not completely closed, antenna positioned at inner edge of orbit; orbits well inward of lateral edge of carapace (Figs 14.35, 14.37–14.40)�������������������������������������������������������������������������������������������������������������������3 Eyes positioned dorsally, above anterolateral edge of carapace; entire margin of orbit visible in dorsal view when eyes retracted. Carapace posterolateral margins smooth, strongly converging, posterior margin strongly narrowed, distance between coxae of pereopods 5 less than 20% carapace width (Fig. 14.35a)�������������������������������������������������������������������������������������������������������������������������������������������������� Dacryopilumnidae Eyes positioned anteriorly, inside anterolateral edge of carapace; entire margin of orbit not visible in dorsal view when eyes retracted. Carapace posterolateral margins serrate, moderately converging, posterior margin, with distance between coxae of pereopods 5 25–30% carapace width (Fig. 14.36a, b)�������������������� Eriphiidae Carapace subhexagonal, anterior surface covered in sharp, curved spines. Chelipeds and legs covered with numerous sharp spines all over dorsal and lateral surface; larger chela with indistinct cutting/crushing tooth (Fig. 14.37)��������������������������������������������������������������������������������������������������������������������������������������������Hypothalassiidae Carapace subhexagonal to transversely ovate; surface rugose smooth, without upright spines; anterolateral teeth low, angular or with short points. Chelipeds and legs smooth to tuberculate not spinose; larger chela usually with distinct molariform crushing teeth���������������������������������������������������������������������������������������������������������4 Endostomial ridges absent (Fig. 14.40i). Male telson half pleonite 3 width (Fig. 14.40h)����������������������������Platyxanthidae Endostomial ridges present on floor of buccal cavity (Fig. 14.38g). Male telson two-thirds pleonite 3 width or more (Figs 14.38j, 14.39l, m)�������������������������������������������������������������������������������������������������������������������������������������������������5 Thoracic sternite 3 wider than anterior width of sternite 4 (Fig. 14.38k, l). Carapace anterolateral margin (excluding exorbital tooth) with 4 broad, angular teeth or lobes, apices often pointed (Fig. 14.38a–d)��������������������������������������������������������������������������������������������������������������������������������������������������������� Menippidae Thoracic sternite 3 narrower than to as wide as anterior width of sternite 4 (Fig. 14.39n, o). Carapace anterolateral margin (excluding exorbital tooth) with 5 (rarely 4) broad, angular teeth or lobes, or margins uneven to irregularly nodulose or spinose (Fig. 14.39a–g)�����������������������������������������������������������������������Oziidae

References Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. Lai JCY, Thoma BP, Clark PF, Felder DL, Ng PKL (2014) Phylogeny of eriphioid crabs (Brachyura, Eriphioidea) inferred from molecular and morphological studies. Zoologica Scripta 43, 52–64. doi:10.1111/zsc.12030 Ng PKL, Davie PJF (2020) A new family and superfamily for the southern giant crab of Australia, Pseudocarcinus gigas (Lamarck, 1818) (Decapoda: Brachyura). Journal of Crustacean Biology 40, 607–626. doi:10.1093/jcbiol/ruaa058 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Ng PKL, Shih H-T, Tan SH, Ahyong ST, Ho P-H (2009) Part I. Carcinology in Taiwan. In Crustacean fauna of Taiwan: Brachyuran crabs, volume 1 – Carcinology in Taiwan and Dromiacea, Raninoida, Cyclodorippoida. (Eds

Chan T-Y, Ng PKL, Ahyong ST, Tan SH) pp. 1–26. National Taiwan Ocean University, Keelung.

Dacryopilumnidae Serène, 1984 Figure 14.35 Controversy has surrounded the taxonomic position of the two species comprising Dacryopilumnus, which until recently were retained within a family in Dairoidea (Ng et al. 2008). A study of larvae and genetic similarities suggested they were closest to Oziidae, a member of the superfamily Eriphioidea (Lai et  al. 2014) but deserving of their own family. They are recognised by the widely separated dorsally positioned orbits and strongly converging posterolateral carapace margins. Diagnosis. Carapace subtrapezoid, posterolateral margins smooth, strongly converging, posterior margin strongly

502

Marine Decapod Crustacea

morphological studies. Zoologica Scripta 43, 52–64. doi:10.1111/ zsc.12030 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo.

Eriphiidae MacLeay, 1838

Fig. 14.35.  Dacryopilumnidae. Dacryopilumnus eremita Nobili, 1906. a, b, Carapace, dorsal, anterior. c, cheliped.

narrowed, distance between coxae of pereopods 5 less than one-fifth carapace width; orbits completely closed, antenna widely separated from and excluded from inner margin of orbit. Orbits placed close lateral edges of anterior margin. Eyes positioned dorsally, above anterolateral edge of carapace; entire margin of orbit visible in dorsal view when eyes retracted. Dacryopilumnus Nobili, 1906 Maximum cl. 7 mm. Intertidal, subtidal (0–2 m). Indo-West Pacific. 2 species (Sakai 1976: key to species) References Lai JCY, Thoma BP, Clark PF, Felder DL, Ng PKL (2014) Phylogeny of eriphioid crabs (Brachyura, Eriphioidea) inferred from molecular and

Figure 14.36, Plate 43h Eriphiids are essentially shallow-water species of rocky shores or reefs. Crabs of the genus Eriphia are said to be the most common and often largest intertidal crabs along many temperate and tropical rocky shores. They are algal grazers and nocturnal predators (Pérez-Miguel et al. 2017; Vannini et  al. 1989). Some Indo-West Pacific species of Eriphia are toxic when eaten by humans (Garth and Alcala 1977). Toxins in reef crabs are often transient and unpredictable (Llewellyn et  al. 2002), so it is wise advice not to indulge in culinary experiments with any small reef crabs in tropical waters. Diagnosis. Carapace subtrapezoid, posterolateral margins serrate, moderately converging, posterior margin narrowed, distance between coxae of pereopods 5 25–30% carapace width; orbits completely closed, antenna widely separated from and excluded from inner margin of orbit. Orbits placed close lateral edges of anterior margin, positioned anteriorly, lower margin of orbit not visible in dorsal view when eyes retracted. Endostomial ridges present. Thoracic sternite 3 narrower than to as wide as anterior width of sternite 4. Chelipeds and legs smooth to tuberculate, at most with several blunt spines on walking leg meri; larger chela usually with distinct molariform crushing teeth. Male telson two-thirds or more as wide as pleonite 3.

Fig. 14.36.  Eriphiidae. a, Eriphia squamata (Stimpson, 1859); b, Eriphides hispida (Stimpson, 1860). Right front carapace, orbit, antenna: c, Eriphia; d, Eriphides. e, f, major, minor chelae, Eriphia. g, h, gonopods 1, 2, Eriphia.

14 – Brachyura – crabs

503

Eriphia Latreille, 1817 Diagnosis. Carapace hexagonal, posterior margin half greatest width, posterolateral margin straight to convex, regions distinct, tuberculate anteriorly; frontal margin between antennae more than one-quarter carapace width; antennae closer to orbit than to midline (Fig.  14.36a, c). Minor chela fingers acute (Fig.  14.36g). Maximum cl. 57 mm (Pl. 43h). Intertidal, subtidal (0–6 m). Cosmopolitan except polar seas. 8 species (Koh and Ng 2008: diagnosis, key to species). Some species are called red-eyed crabs on account of the colour of their cornea in life.

Eriphides Rathbun, 1897 Diagnosis. Carapace hexagonal, posterior margin half greatest width, posterolateral margin straight to convex, regions indistinct, tuberculate anteriorly; frontal margin between antennae less than quarter carapace width; antennae closer to midline than to orbit (Fig. 14.36b, d). Minor chela fingers with hollowed, spoonshaped tips (Fig. 14.36b). Maximum cl. 44 mm. Intertidal (rocky shore, in burrows in sandstone). Tropical Eastern Pacific. 1 species (Rathbun 1930: redescription). References Garth JS, Alcala A (1977) Poisonous crabs of the Indo-West Pacific coral reefs, with special reference to the genus Demania Laurie. In Proceedings of the the Third International Coral Reef Symposium, Rosential School of Marine and Atmospheric Science. pp. 645–651, figs 1–7. University of Miami, Miami. Koh S, Ng PKL (2008) A revision of the shore crabs of the genus Eriphia (Crustacea: Brachyura: Eriphiidae). Raffles Bulletin of Zoology 56, 327–355. Llewellyn LE, Dodd MJ, Robertson A, Ericson G, de Koning C, et  al. (2002) Post-mortem analysis of samples from a human victim of a fatal poisoning caused by the xanthid crab, Zosimus aeneus. Toxicon 40, 1463–1469. doi:10.1016/S0041-0101(02)00164-2 Pérez-Miguel M, Drake P, Cuesta JA (2017) Experimental predatory behavior of the stone crab Eriphia verrucosa (Forskål, 1775) (Decapoda, Brachyura, Eriphiidae). Nauplius 25, e2017033. doi:10.1590/23582936e2017033 Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Vannini M, Chelazzi G, Gherardi F (1989) Feeding habits of the pebble crab Eriphia smithi (Crustacea, Brachyura, Menippidae). Marine Biology 100, 249–252. doi:10.1007/BF00391965

Hypothalassiidae Karasawa & Schweitzer, 2006 champagne crabs Figure 14.37 The changing family placement of the only genus, Hypothalassia, was reviewed by Koh and Ng (2000) who placed it in Eriphiidae, but it has since been placed in its own family

Fig. 14.37.  Hypothalassiidae. Hypothalassia acerba Koh & Ng, 2000

(Karasawa and Schweitzer 2006). Hypothalassiids are distinctive eriphioids in their great size and prominent, curved black-brown corneous spines covering the chelipeds, walking legs and carapace. Hypothalassia acerba Koh & Ng, 2000 is the basis of a small pot fishery in Western Australia (Smith et al. 2004) Diagnosis. Carapace subhexagonal, anterior surface and margins covered in sharp, curved spines; orbits well inward of lateral edge of carapace, not completely closed, antenna positioned at inner edge of orbit. Thoracic sternite 3 narrower than to as wide as anterior width of sternite 4. Chelipeds and legs covered with numerous sharp spines all over dorsal and lateral surface; larger chela with indistinct cutting/crushing tooth. Male telson two-thirds or more as wide as pleonite 3. Hypothalassia Gistel, 1848 Shelf, slope (50–600 m). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Australasia. Maximum cl. 140 mm. 2 species (diagnoses, keys to species: Koh and Ng 2000; Poore 2004). References Karasawa H, Schweitzer CE (2006) A new classification of the Xanthoidea sensu lato (Crustacea: Decapoda: Brachyura) based on phylogenetic analysis and traditional systematics and evaluation of all fossil Xanthoidea sensu lato. Contributions to Zoology 75, 23–73. doi:10.1163/18759866-0750102002 Koh SK, Ng PKL (2000) A revision of the spiny crabs of the genus Hypothalassia Gistel, 1848 (Crustacea: Decapoda: Brachyura: Eriphiidae). Raffles Bulletin of Zoology 48, 123–141. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Smith KD, Hall NG, Potter IC (2004) Relative abundances and size compositions of champagne crabs, Hypothalassia acerba, on two coasts and in different water depths and seasons. Marine and Freshwater Research 55, 653–661. doi:10.1071/MF04063

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Menippidae Ortmann, 1893 Figure 14.38, Plate 43i, j Menippids are oval, relatively smooth crabs with small eyes close together. Differences from oziids and platyxanthids are slight. The family includes some of the largest crabs, including stone crabs harvested for their claws in North America. Phylogenetic relationships among the menippids and allies are yet to be clearly resolved, particularly in Menippidae and Oziidae, both of which are polyphyletic according to Lai et al. (2014). Menippidae is here restricted to three genera; two others that have traditionally been included in Menippidae are excluded: Pseudocarcinus H. Milne Edwards, 1834, was recently referred to Pseudocarcinoidea (Ng and Davie

2020) and Ruppellioides A. Milne-Edwards, 1867 is herein referred to Oziidae as a synonym of Ozius. Diagnosis. Carapace subhexagonal to transversely ovate, anterior margin arcuate, surface rugose or tuberculate to smooth; anterolateral margin (excluding exorbital tooth) with 4 broad, angular teeth or lobes, apices often pointed; endostomial ridges present (Fig. 14.38g); orbits well inward of lateral edge of carapace, not completely closed, antenna positioned at inner edge of orbit. Thoracic sternite 3 wider than anterior width of sternite 4. Chelipeds and legs smooth to granulate, never prominently spinose; larger chela with distinct molariform crushing teeth (Fig. 14.38a). Male telson two-thirds or more as wide as pleonite 3 (Fig. 14.38j).

Key to genera of Menippidae 1. – 2. –

Orbital hiatus absent, supra- and suborbital angles in contact (Fig. 14.38f). Carapace regions moderately well marked (Fig. 14.38a, c)���������������������������������������������������������������������������������������������������������������������������������� Myomenippe Orbital hiatus present, supra- and suborbital angles not meeting (Fig. 14.38e). Carapace regions obscure (Fig. 14.38b, d)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Front with 6 lobes, distance between antennae one-seventh carapace width. Carapace anterolateral margin about as long as posterolateral margin, 3 broad asymmetrical lobes, all rounded anteriorly, and blunt epibranchial tooth (Fig. 14.38b)���������������������������������������������������������������������������������������������������������������������������������� Menippe Front with 2 lobes, distance between antennae one-quarter carapace width. Carapace anterolateral margin shorter than posterolateral margin, 3 asymmetrical, wide, anteriorly pointed or angular teeth, third largest, and sharp epibranchial tooth (Fig. 14.38d)��������������������������������������������������������������������������������������������Sphaerozius Menippe De Haan, 1833

stone crabs Diagnosis. Carapace smooth, regions obscure; front with 6 lobes, distance between antennae seventh carapace width; anterolateral

margin about as long as posterolateral margin, with exorbital tooth plus 3 broad asymmetrical lobes, all rounded anteriorly, epibranchial tooth blunt; supra- and suborbital angles not in contact, with orbital hiatus. Antennal basal article excluded from orbit. Maximum cl. 86 mm (Pl. 42i).

Fig. 14.38.  Menippidae. a, Myomenippe fornasinii (Bianconi, 1851). Carapace: b, Menippe; c, Myomenippe; d, Sphaerozius. Left front: e, Menippe; f, Myomenippe. Endostome with ridges: g, Menippe. Gonopods 1, 2: h, i, Sphaerozius. Male pleon: j, Menippe. Thoracic sternites 1–4: k, Menippe; l, Sphaerozius.

14 – Brachyura – crabs

Intertidal–shelf (0–80 m). Tropical Atlantic, Western IndoPacific, Tropical Eastern Pacific. 8 species (Rathbun 1930: key to 4 species; Schneider-Broussard et  al. 1998: phylogeography of two species; Williams and Felder 1986). Stone crabs, particularly M. mercenaria (Say, 1818) are prized for their claws and are fished throughout their range in the south-eastern USA and the C ­ aribbean (Williams and Felder 1986).

Myomenippe Hilgendorf, 1879 thunder crab Diagnosis. Carapace smooth, regions moderately well marked; front with 6 lobes, distance between antennae one-fifth carapace width; anterolateral margin longer than posterolateral margin, with exorbital tooth plus 3 lobes, square anteriorly, third largest, and sharp epibranchial tooth; supra- and suborbital angles in contact, without orbital hiatus. Antennal basal article excluded from orbit. Maximum cl. 60 mm (Pl. 43j). Intertidal. Western and Central Indo-Pacific. 2 species (­Bianconi 1851: figure; Crosnier 1984: key to species, figures; De Man 1888: figure).

Sphaerozius Stimpson, 1858 Diagnosis. Carapace smooth, regions obscure; front with 2 low lobes, distance between antennae one-quarter carapace width; anterolateral margin shorter than posterolateral margin, with exorbital tooth plus 3 asymmetrical wide, anteriorly pointed or angular teeth, third largest, and sharp epibranchial tooth; supra- and suborbital angles not in contact, with orbital hiatus. Antennal basal article very short, not in contact with front. Maximum cl. 17 mm. Intertidal–shelf (0–35 m). Temperate Northern Pacific, IndoWest Pacific, Temperate Southern Africa, Mediterranean Sea (Brockerhoff and McLay 2011: S. nitidus Stimpson, 1858 introduced). 2 species (Crosnier 1984: key to species; Miers 1886: description; Sakai 1976: figure). References Bianconi JJ (1851) Specimina zoologica Mosambicana. Fasciculus V. Memorie della Accademia delle Scienze dell’Istituto di Bologna 3, 91–112, pls 8–11. Brockerhoff A, McLay C (2011) Human-mediated spread of alien crabs. In In the Wrong Place – Alien Marine Crustaceans: Distribution, Biology and Impacts. (Eds Galil BS, Clark PF, Carlton JT) pp. 27–106. Springer, Dordrecht. Crosnier A (1984) Addendum: Familles de Carpiliidae et des Menippidae. In: Serène, R. Crustacés Décapodes Brachyoures de l’Océan Indien Occidental et de la Mer Rouge, Xanthoidea: Xanthidae et Trapeziidae. Faune Tropicale 24, 299–313. De Man JG (1888) Report on the podophthalmous Crustacea of the Mergui Archipelago, collected for the Trustees of the Indian Museum, Calcutta, by Dr. John Anderson, F.R.S., Superintendent of the Museum. Journal of the Linnean Society of London. Zoology 22, 1–312, pls 1–19. Lai JCY, Thoma BP, Clark PF, Felder DL, Ng PKL (2014) Phylogeny of eriphioid crabs (Brachyura, Eriphioidea) inferred from molecular and morphological studies. Zoologica Scripta 43, 52–64. doi:10.1111/zsc.12030

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Miers EJ (1886) Zoology Part XLIX. Report on the Brachyura collected by H.M.S. Challenger during the years 1873–76. Report of the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873–76 17, i–l, 1–362, pls 1–29. Ng PKL, Davie PJF (2020) A new family and superfamily for the southern giant crab of Australia, Pseudocarcinus gigas (Lamarck, 1818) (Decapoda: Brachyura). Journal of Crustacean Biology 40, 607–626. doi:10.1093/jcbiol/ruaa058 Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo. Schneider-Broussard R, Felder DL, Chlan CA, Neigel JE (1998) Tests of phylogeographic models with nuclear and mitochondrial DNA sequence variation in the stone crabs, Menippe adina and Menippe mercenaria. Evolution 52, 1671–1678. doi:10.1111/j.1558-5646.1998. tb02246.x Williams AB, Felder DL (1986) Analysis of stone crabs: Menippe mercenaria (Say), restricted, and a previously unrecognized species described (Decapoda: Xanthidae). Proceedings of the Biological Society of Washington 99, 517–543.

Oziidae Alcock, 1898 Figure 14.39, Plate 43k, l Oziids are usually oval, having a somewhat swollen to flattened carapace, usually with 4 or 5 anterolateral teeth. Some are called reef crabs associated with coral or rocky reefs but others live in muddy mangrove environments where they are known predators (Cannicci et  al. 1998; Sivaguru and McLay 2010). Like menippids, oziids have molar-like teeth on the major chela and are subtly differentiated from that family primarily on the relative proportions of the anterior thoracic sternites. The two families have been treated as synonymous in the past – molecular evidence has suggested that neither family is monophyletic (Lai et al. 2014). Most genera and species were described during the nineteenth century. Only one large genus has been reviewed since. Diagnosis. Carapace subhexagonal to transversely ovate, anterior margin arcuate; surface rugose or tuberculate to smooth; anterolateral margin (excluding exorbital tooth) with 5 (rarely 4) broad, angular teeth or lobes, or margins uneven to irregularly nodulose or spinose; orbits well inward of lateral edge of carapace, not completely closed, antenna positioned at inner edge of orbit. Endostomial ridges present. Thoracic sternite 3 narrower than to as wide as anterior width of sternite 4. Chelipeds and legs smooth to tuberculate, never prominently spinose; larger chela with distinct molariform crushing teeth (Fig. 14.39a, d, g). Walking legs smooth to weakly tuberculate, without spines on merus extensor margin. Male telson about two-thirds as wide as pleonite 3 (Fig. 14.39l, m).

506

Marine Decapod Crustacea

Key to genera of Oziidae 1. – 2. – 3. – 4. – 5. – 6. –

Orbit closed, supra- and suborbital angles in contact. Antenna basal article in contact with front (Fig. 14.39h)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Orbit open, supra- and suborbital angles not in contact. Antenna basal article not reaching front (Fig. 14.39i)���������������� 5 Minor cheliped dactylus shorter than upper margin of palm (Fig. 14.39b, f)������������������������������������������������������������������������3 Minor cheliped dactylus longer than upper margin of palm (Fig. 14.39a, c)��������������������������������������������������������������������������4 Carapace anterolateral margin shorter than posterolateral margin, with 3 small, blunt, well separated, spines (Fig. 14.39b)����������������������������������������������������������������������������������������������������������������������������������������������������� Bountiana Carapace anterolateral margin longer than posterolateral margin, with 5 lobes or teeth (Fig. 14.39f)���������������������Lydia Major cheliped dactylus shorter than upper margin of palm. Carapace anterolateral teeth broad, closely spaced, angular lobes (Fig. 14.39a)��������������������������������������������������������������������������������������������������������������������������� Baptozius Chelipeds similar, dactylus longer than upper margin of palm. Carapace anterolateral teeth narrow, pointed, widely spaced projections (Fig. 14.39c)���������������������������������������������������������������������������������������������Epixanthoides Minor cheliped dactylus longer than upper margin of palm. Carapace anterolateral margin longer than posterolateral margin, regularly lobate or lobes indistinct (Fig. 14.39d)���������������������������������������������������������Epixanthus Minor cheliped dactylus shorter than upper margin of palm. Carapace anterolateral margin about as long or indistinct from posterolateral margin��������������������������������������������������������������������������������������������������������������������������������6 Carapace little wider than long, setose. Cheliped palm covered with setose, conical tubercles (Fig. 14.39e)Eupilumnus Carapace 1.5 times as wide as long, not setose. Cheliped palm usually smooth (Fig. 14.39g), rarely tuberculate, always glabrous����������������������������������������������������������������������������������������������������������������������������������������������Ozius

Fig. 14.39.  Oziidae. Carapace, some with chelipeds, ambulatory legs: a, Baptozius vinosus (H. Milne Edwards, 1834); b, Bountiana norfolcensis (Grant & McCulloch, 1907); c, Epixanthoides anomalus Balss, 1935; d, Epixanthus; e, Eupilumnus fragaria (Yang, Dai & Ng, 1998); f, Lydia; g, Ozius. Left orbit, antenna: h, Bountiana; i, Ozius. Gonopods 1, 2: j, k, Eupilumnus. Male pleon: l, m, Eupilumnus. Thoracic sternites 1–4: n, Epixanthus; o, Eupilumnus.

14 – Brachyura – crabs

Baptozius Alcock, 1898 Diagnosis. Carapace broad, transversely oval, moderately convex; regions not indicated; frontal margin 40% of carapace width, obliquely deflexed, straight; anterolateral margin shorter than posterolateral margin, with 4 sharply defined teeth. Orbit closed, supra- and suborbital angles in contact. Antenna basal article in contact with front. Minor cheliped dactylus longer than upper margin of palm. Maximum cl. 8 mm. Intertidal, mudflats. Western and Central Indo-Pacific. 1 species (Alcock 1898; 1899: illustration).

507

posterolateral margin, with 5 or more teeth, spines or granules. Orbit open, supra- and suborbital angles not in contact. Antenna basal article not reaching front. Major cheliped palm swollen, covered with conical, setose tubercles. Minor cheliped dactylus about as long as upper margin of palm. Maximum cl. 11 mm. Intertidal–slope (0–624 m). Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific. Temperate Australasia. 9 species (Yang et  al. 1998: key to species as Globopilumnus). The genus was for many years known as Globopilumnus Balss, 1933. Eupilumnus is a senior synonym. Three species from the Atlantic and one from the Indo-West Pacific have stridulatory organs on the chelipeds (Ng and Tan 1985; Yang et al. 1998).

Bountiana Davie & Ng, 2000 Lydia Gistel, 1848

Diagnosis. Carapace transversely oval, strongly vaulted anteriorly; regions poorly defined; frontal margin 40% of carapace width, obliquely deflexed, with median notch; anterolateral margin shorter than posterolateral margin, with 3–5 small, blunt, well separated, granules. Orbit closed, supra- and suborbital angles in contact. Antenna basal article in contact with front. Minor cheliped dactylus shorter than upper margin of palm. Maximum cl. 12 mm. Intertidal, reefs. Temperate Australia. 1 species (Davie and Ng 2000).

Diagnosis. Carapace wider than long; anterior regions well demarcated; frontal margin about one-third carapace width, with 4 lobes, separated from orbit by notch; anterolateral margin longer than posterolateral margin, with 5 lobes or teeth. Orbit closed, supra- and suborbital angles in contact. Antenna basal article in contact with front. Minor cheliped dactylus shorter than upper margin of palm. Maximum cl. 31 mm. Intertidal, subtidal (0–1 m). Indo-West Pacific. 4 species (Crosnier 1984: discussion, figures).

Epixanthoides Balss, 1935

Ozius H. Milne Edwards, 1834

Diagnosis. Carapace broad, transversely oval, moderately convex; regions poorly defined; frontal margin about one-quarter carapace width, margin broadly bilobate, separated from orbit by notch; anterolateral margin curved, slightly longer than posterolateral margin, with 4 short, pointed, widely spaced teeth. Orbit closed, supra- and suborbital angles not in contact. Antenna basal article reaching front. Chelipeds subequal, dactylus longer than upper margin of palm. Maximum cl. 5 mm. Intertidal, subtidal (0–6 m; coral rubble). Indo-West Pacific. 1 species (Ng et al. 2008: colour figure).

Epixanthus Heller, 1861 Diagnosis. Carapace broad, transversely oval, moderately convex; regions poorly defined, with anterolateral sculpture; frontal margin about one-quarter carapace width, obliquely deflexed, margin irregularly transverse to 4 or 5 lobate, separated from orbit by notch; anterolateral margin curved, longer than posterolateral margin, regularly lobate. Orbit open, supra- and suborbital angles not in contact. Antenna basal article not reaching front. Minor cheliped dactylus longer than upper margin of palm. Maximum cl. 35 mm (Pl. 43k). Intertidal. Indo-West Pacific. 6 species (Crosnier 1984: key to 3 Indo-West Pacific species, figures).

Eupilumnus Kossmann, 1877 Diagnosis. Carapace little wider than long, round or squarish, moderately convex, setose; regions poorly defined; frontal margin with 2 broadly convex lobes; anterolateral margin continuous with

iron crab Diagnosis. Carapace broad, transversely oval, moderately convex; regions poorly defined; frontal margin about one-quarter carapace width, obliquely deflexed, with 4 weak lobes, separated from orbit by notch; anterolateral margin about as long as posterolateral margin, with 5 teeth or lobes. Orbit open, supra- and suborbital angles not in contact. Antenna basal article not reaching front. Minor cheliped dactylus about as long as upper margin of palm. Maximum cl. 50 mm (Pl. 43l). Intertidal, subtidal (0–6 m). Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Tropical Eastern Pacific, Temperate Southern Africa, Temperate Australasia. 12 species (Sakai 1976: key to 3 Japanese species; Sivaguru and McLay 2010: biology). McNeill (1968) synonymised the type species of Ruppellioides A. MilneEdwards, 1867 with O. tuberculosus H. Milne Edwards, 1834, the type species of Ozius. Accordingly, Ruppellioides is a synonym of Ozius. References Alcock A (1898) Materials for a carcinological fauna of India. No. 3. The Brachyura Cyclometopa. Part I. The family Xanthidae. Journal of the Asiatic Society of Bengal 67, 67–233. Alcock A (1899) Illustrations of the zoology of the Royal Indian marine surveying steamer, Investigator, under the command of Commander T.H. Heming, R.N. Crustacea. – Part VII, Plates XXXVI–XLV. Indian Museum, Calcutta. Cannicci S, Ruwa RK, Giuggioli M, Vannini M (1998) Predatory activity and spatial strategies of Epixanthus dentatus (Decapoda: Oziidae), an ambush predator among the mangroves. Journal of Crustacean Biology 18, 57–63. doi:10.2307/1549520

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Crosnier A (1984) Addendum: Familles de Carpiliidae et des Menippidae. In: Serène, R. Crustacés Décapodes Brachyoures de l’Océan Indien Occidental et de la Mer Rouge, Xanthoidea: Xanthidae et Trapeziidae. Faune Tropicale 24, 299–313. Davie PJF, Ng PKL (2000) Bountiana, a new genus for Eriphia norfolcensis Grant & McCulloch, 1907 (Crustacea: Brachyura: Eriphiidae). Memoirs of the Queensland Museum 45, 267–272. Lai JCY, Thoma BP, Clark PF, Felder DL, Ng PKL (2014) Phylogeny of eriphioid crabs (Brachyura, Eriphioidea) inferred from molecular and morphological studies. Zoologica Scripta 43, 52–64. doi:10.1111/zsc.12030 McNeill F (1968) Crustacea, Decapoda & Stomatopoda. Great Barrier Reef Expedition 1928–29 Scientific Reports 7, 1–98, pls 1, 2. Ng PKL, Tan LWH (1985) Globopilumnus multituberosus Garth & Kim, 1983 – a new record for Australia (Decapoda: Crustacea: Menippidae). Records of the Australian Museum 36, 127–129. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo. Sivaguru K, McLay CL (2010) Population dynamics and distribution of Ozius truncatus H. Milne Edwards, 1834 (Brachyura, Oziidae) on Echinoderm Reef, Leigh Marine Reserve, New Zealand. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 301–317. Yang S-L, Dai A-Y, Ng PKL (1998) Globopilumnus fragaria, a new species of eriphiid crab (Crustacea: Decapoda: Brachyura: Xanthoidea) from the South China Sea, with a key to the genus Globopilumnus Balss, 1933. Raffles Bulletin of Zoology 46, 93–99.

Platyxanthidae Guinot, 1977 Figure 14.40 The family of five genera and six species, all large crabs from Atlantic and Pacific coasts of Central and South America, has been reviewed in detail (Thoma et al. 2012). Their paper described and illustrated all species, tabulated characteristics of the genera and provide a key. Our key and diagnoses are simplified from their work. Distributions were derived from Thoma et al. (2012), Rathbun (1911, 1930) and Garth (1940). Unlike other eriphioids, platyxanthids lack endostomial ridges. Diagnosis. Carapace subhexagonal to transversely ovate; surface rugose or smooth; anterolateral margin (excluding exorbital tooth) without or with up to 8 teeth; orbits well inward of lateral edge of carapace, not completely closed, antenna positioned at inner edge of orbit. Endostomial ridges absent (Fig. 14.40i). Thoracic sternite 3 narrower than to as wide as anterior width of sternite 4. Chelipeds and legs smooth to tuberculate, at most with several blunt spines on walking leg meri (Fig.  14.40a); larger chela usually with distinct molariform crushing teeth. Male telson half as wide as pleonite 3 (Fig. 14.40h).

Key to genera of Platyxanthidae 1. – 2. – 3. – 4. –

Carapace front straight with slight median sulcus; anterolateral margin with 3 broad, weakly divided lobes (excluding epibranchial tooth). Inner supraorbital tooth absent or weak (Fig. 14.40d)�������������������������������Homalaspis Carapace front subdivided into 4 prominences; anterolateral margin with 3 or more well separated lobes or teeth (excluding epibranchial tooth). Inner supraorbital tooth present (Fig. 14.40b, c, e–g)������������������������������������������2 Carapace anterolateral margin with 8 or more teeth (Fig. 14.40g). Maxilliped 3 merus covering nephridiopore operculum (Fig. 14.40n)������������������������������������������������������������������������������������������������������������� Platyxanthus Carapace anterolateral margin with 4 teeth (Fig. 14.40b, c, e, f). Maxilliped 3 merus not covering nephridiopore operculum (Fig. 14.40h–k)������������������������������������������������������������������������������������������������������������������������������3 Anterolateral margin with 3 blunt well separated teeth plus epibranchial tooth (Fig. 14.40a, e). Antennular fossa inclined at ~50° angle from horizontal (Fig. 14.40l). Gonopod 2 less than one-third length of gonopod 1�����������������������������������������������������������������������������������������������������������������������������������������������������������Otmaroxanthus Anterolateral margin with 3 teeth, first 2 or all lobiform, subdivided or reduced to tubercles, plus epibranchial tooth (Fig. 14.40b, c, f). Antennular fossa inclined at ~25° angle from horizontal (Fig. 14.40j, n). Gonopod 2 as long or longer than gonopod 1���������������������������������������������������������������������������������������������4 Carapace strongly convex. Inner supraorbital tooth about as long as outer supraorbital tooth (Fig. 14.40f). Maxilliped 3 merus longer than wide (Fig. 14.40m)���������������������������������������������������������������������������������������������������Peloeus Carapace moderately convex. Inner supraorbital tooth strong, reaching anteriorly well beyond outer supraorbital tooth (Fig. 14.40b, c). Maxilliped 3 merus as wide as long (Fig. 14.40j)������������������������������������Danielethus Danielethus Thoma, Ng & Felder, 2012

Diagnosis. Carapace moderately convex; front with 4 lobes, deep median notch; anterolateral margin with 3 broad, well divided lobes, each with lobulate margin, sharp epibranchial tooth. Inner supraorbital tooth longer than outer supraorbital tooth.

Maxilliped 3 merus distal margin as long as wide, not covering nephridiopore operculum when retracted. Antennular fossa inclined at ~25° angle from horizontal. Gonopod 2 as long as gonopod 1. Maximum cl. 50 mm. Shelf (46–119 m). Temperate South America (Uruguay–Peru). 2 species (Thoma et al. 2012).

14 – Brachyura – crabs

509

Fig. 14.40.  Platyxanthidae. a, Otmaroxanthus balboai (Garth, 1940). Carapace: b, Danielethus crenulatus (A. Milne-Edwards, 1879); c, D. patagonicus (A. Milne-Edwards, 1879); d, Homalaspis plana (H. Milne Edwards, 1834); e, Otmaroxanthus balboai (Garth, 1940); f, Peloeus armatus Eydoux & Souleyet, 1842; g, Platyxanthus orbignyi (H. Milne Edwards & Lucas, 1843). Pleon: h, Otmaroxanthus. i, epistome, Danielethus. Front, orbit, maxilliped 3: j, Danielethus; k, Homalaspis; l, Otmaroxanthus; m, Peloeus; n, Platyxanthus.

Homalaspis A. Milne-Edwards, 1863 Diagnosis. Carapace flat; front straight with slight median sulcus; anterolateral margin with 3 broad, well divided lobes, sharp epibranchial tooth. Inner supraorbital tooth absent, or weak. Maxilliped 3 merus distal margin strongly convex; not covering nephridiopore operculum when retracted. Antennular fossa inclined at ~33° angle from horizontal. Gonopod 2 as long as gonopod 1. Maximum cl. 66 mm. Intertidal. Temperate South America (Chile). 1 species (Thoma et al. 2012).

Otmaroxanthus Števčić, 2011 Diagnosis. Carapace moderately convex; front with 4 teeth; anterolateral margin with 3 blunt well separated teeth, blunt epibranchial tooth. Inner supraorbital tooth longer than outer supraorbital tooth. Maxilliped 3 merus distal margin straight; not covering nephridiopore operculum when retracted. Antennular fossa inclined at ~50° angle from horizontal. Gonopod 2 one-third length of gonopod 1. Maximum cl. 31 mm. Shelf (60–140 m). Tropical Eastern Pacific (Panama). 1 species (Garth 1940; Thoma et al. 2012).

Peloeus Eydoux & Souleyet, 1842 Diagnosis. Carapace strongly convex; front with 4 blunt teeth; anterolateral margin with 3 broad, well divided lobes, sharp epibranchial tooth. Inner supraorbital tooth as long as outer supraorbital tooth. Maxilliped 3 merus longer than wide, not covering

nephridiopore operculum when retracted. Antennular fossa inclined at ~25° angle from horizontal. Gonopod 2 longer than gonopod 1. Maximum cl. 60 mm. Subtidal, shelf (15–40 m). Eastern Indo-Pacific (Peru). 1 species (Thoma et al. 2012).

Platyxanthus A. Milne-Edwards, 1863 Diagnosis. Carapace moderately convex; front with 4 teeth; anterolateral margin with 8 blunt well separated teeth, sharp epibranchial tooth. Inner supraorbital tooth as long as outer supraorbital tooth. Maxilliped 3 merus distal margin strongly convex; covering nephridiopore operculum when retracted. Antennular fossa inclined at ~38° angle from horizontal. Gonopod 2 longer than gonopod 1. Maximum cl. 60 mm. Intertidal. Tropical Eastern Pacific, Temperate South America (Ecuador–Chile). 1 species (Thoma et al. 2012). References Garth JS (1940) Some new species of brachyuran crabs from Mexico and the Central and South American mainland. Allan Hancock Pacific Expeditions 5, 53–95, pls 11–26. Rathbun MJ (1911) The stalk-eyed Crustacea of Peru and the adjacent coast. Proceedings of the United States National Museum 38, 531–620, pls 36–56. Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Thoma BP, Ng PKL, Felder DL (2012) Review of the family Platyxanthidae Guinot, 1977 (Crustacea, Decapoda, Brachyura, Eriphioidea), with

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the description of a new genus and a key to genera and species. Zootaxa 3498, 1–23. doi:10.11646/zootaxa.3498.1.1

Goneplacoidea MacLeay, 1838 The superfamily Goneplacoidea is taxonomically challenging. For most of their history, the goneplacoids have comprised a series of potentially disparate groups assembled largely for taxonomic convenience (Castro 2007; Ng et  al. 2008). Not surprisingly, numerous, sometimes conflicting schemes have been proposed (e.g. Castro 2007; Guinot 1969a, 1969b, 1969c; Karasawa and Kato 2003; Karasawa and Schweitzer 2006; Števčić 2005). Here, we follow the 11 family classification of Castro et al. (2010) and Davie et al. (2015), realising that this arrangement is a work in progress and significant changes are likely in future revisions. No known synapomorphies unite the goneplacoids, and they are probably polyphyletic as currently conceived (Castro et al. 2010; Tsang et  al. 2014). As a result, goneplacoids are currently recognised more by what they are not, than by what they are. They are not portunoids, lacking the swimming legs or a transverse ridge on the male pleonite 3; they are not pilumnoids, lacking the sigmoid gonopod 1 and minute gonopod 2; they are not xanthoids, usually lacking fused male pleonites 3–5; nor are they eriphiioids (themselves difficult to characterise), typically having a wider pleon and less carapace ornamentation. After excluding the aforementioned groups, most of what remains among conventionally ‘crablike’ heterotrematous crabs are goneplacoids.

Although no single feature reliably distinguishes Goneplacoidea, the following general observations may aid in their recognition among samples: • Most goneplacoids occur in relatively deep water at shelf to upper slope depths on open substrates, and so often have slender walking legs and elongated dactyli with soft setae on the flexor margin, rather than the more robust legs and dactyli with small spinules, common in xanthoids and eriphioids that usually live in tight crevices. • Most crabs with long-stalked eyes in intertidal and shallow subtidal habitats are ocypodoids, but crabs with similarly stalked eyes from greater depths are usually goneplacoids, particularly Goneplacidae, Euryplacidae and Sotoplacidae • Goneplacoids are usually naked or sparsely setose, rarely densely covered in setae as in Acidopsidae. Diagnosis. Carapace transversely ovate to hexagonal or squarish, as wide as long to wider than long. Antennule folding transversely or obliquely. Endostomial ridges present. Buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular. Cheliped palm ovate in cross-section. Pereopods 2–5 inserted laterally on body, similar, as walking legs, dactyli slender, distally pointed. Male pleonites 1–6 and telson free, or male pleonites 3–4, 3–5 or 4–5 fused, immovable, even if some or all sutures visible. Gonopod 1 stout to slender, typically distally tapering, weakly curved to sinuous, without complex lobes or folds. Gonopod 2 straight.

Key to families of Goneplacidae 1. – 2. – 3. – 4. – 5. –

Carapace and pereopods covered with thick, coarsely setose ridges and convoluted rugosites (Fig. 14.54)���������������������������������������������������������������������������������������������������������������������������������������������������������������Vultocinidae Carapace and pereopods smooth or uneven, granular or at most with few low ridges; carapace surface glabrous or evenly setose������������������������������������������������������������������������������������������������������������������������������������������������������������2 Male genital papilla largely concealed by supplementary plate (calcified structure formed by invagination of surface of thoracic sternite 8) (Fig. 14.42n–p)������������������������������������������������������������ Chasmocarcinidae Male thoracic sternite 8 without supplementary plate; when pleon deflexed, male genital papilla either exposed, lying in open groove between thoracic sternites 7 and 8, or not visible externally, lying in closed gutter formed by appressed margins of sternites 7 and 8�����������������������������������������������������������������������������������������3 Male telson lateral margins distinctly concave (Fig. 14.53c). Male genital papilla not visible externally, lying in closed gutter formed by appressed margins of sternites 7 and 8 (Fig. 14.53d)���������������������������������������� Sotoplacidae Male telson lateral margins straight to convex. Male genital papilla exposed, lying in open groove between thoracic sternites 7 and 8�����������������������������������������������������������������������������������������������������������������������������������������������������������4 Gonopod 2 two-thirds length of gonopod 1 or less (Figs 14.45q–t, 14.48l, n, 14.52g, h)�������������������������������������������������������5 Gonopod 2 as long as or longer than gonopod 1 (Figs 14.41x, y, 14.44c, d, 14.48i–k, m, o–s)����������������������������������������������9 Cheliped merus with stridulatory ridge (rasp) on inner distal margin; ischium and merus fused (Fig. 14.48e)�����������������������������������������������������������������������������������������������������������������������������Goneplacidae (Bathyplax only) Cheliped merus without stridulatory ridge on inner distal margin; ischium and merus articulating������������������������������6

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6.

Gonopod 2 two-thirds length of gonopod 1 (Fig. 14.49e, f). Vulva with 2 or more rounded lobes on margin (Fig. 14.49d)������������������������������������������������������������������������������������������������������������������������������������������������������������ Litocheiridae – Gonopod 2 half gonopod 1 length or less (e.g. Fig. 14.45q–t). Vulva with simple margin or single vulvar cover����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������7 7. Carapace anterolateral margin unarmed; widest posteriorly, posterolateral margins divergent or subparallel. Pereopods 2–5 merus with extensor and flexor margins armed with row of small spines (Fig. 14.52a, b)��������������������������������������������������������������������������������������������������������������������������������������������������������Scalopidiidae – Carapace with 1 or more anterolateral spines; widest near or anterior to midlength, posterolateral margins converging (Figs 14.45a–n, 14.47i, o). Pereopods 2–5 merus at most with spines on extensor margin; flexor margin always unarmed�������������������������������������������������������������������������������������������������������������������������������������������������8 8. Eyes 1.4 times or longer than width of front (Fig. 14.47i, o). Gonopod 2 0.4–0.5 length of gonopod 1 (Fig. 14.48l, n)����������������������������������������������������������������������������������������Goneplacidae (Microgoneplax, Paragoneplax only) – Eyes shorter than or equal to width of front (Fig. 14.45a–n). Gonopod 2 one-third length of gonopod 1 or less (Fig. 14.45p–s)������������������������������������������������������������������������������������������������������������������������������������������������� Euryplacidae 9. Maxilliped 3 merus distolateral margin produced to acute, triangular lobe (Fig. 14.44b)�������������������������������� Conleyidae – Maxilliped 3 merus distolateral margin rounded, square or obtusely angular (Figs 14.41s, t, 14.50n, m, 14.51g)������������� 10 10. Carapace surface usually densely setose and coarsely granular (Fig. 14.41a, b). Pleonite 1 wider than pleonite 3 only if surface smooth and glabrous (Fig. 14.41m, n)���������������������������������������������������������������������Acidopsidae – Carapace surface glabrous and smooth to finely granular, at most sparsely setose (Fig. 14.46, 14.47b–s, 14.50a–e). Pleonite 3 as wide as or wider than pleonite 1 (Figs 14.46, 14.50i, 14.47b–s)������������������������������������������������11 11. Male thoracic sternum with ‘press-button’ of pleonal-locking mechanism on anterior half of sternite 5, near suture 4/5 (Fig. 14.48b)����������������������������������������������������������������������������������������������������������������Goneplacidae (in part) – Male thoracic sternum with ‘press-button’ of pleonal-locking mechanism on posterior half of sternite 5, near midlength or further posteriad (Figs 14.50h, 14.51g)�������������������������������������������������������������������������������������������������11 12. Male pleonites 3–4 or 3–5 fused but demarcation between fused somites clearly indicated by complete transverse groove (Fig. 14.50i)��������������������������������������������������������������������������������������������������������������������������Mathildellidae – Male pleonites 1–6 freely articulating or 3–4 fused (Fig. 14.51e, f) (if pleonites 3–4 fused, demarcation indicated only medially and at lateral margins (Fig. 14.51d)�����������������������������������������������������������������������Progeryonidae References Castro P (2007) A reappraisal of the family Goneplacidae MacLeay, 1838 (Crustacea, Decapoda, Brachyura) and revision of the subfamily Goneplacinae, with the description of 10 new genera and 18 new species. Zootaxa 29, 609–774. Castro P, Guinot D, Ng PKL (2010) A new family for Sotoplax robertsi Guinot, 1984, with a diagnosis and key to the Goneplacoidea Macleay, 1838 (Crustacea: Decapoda: Brachyura). Zootaxa 2356, 36–56. doi:10.11646/zootaxa.2356.1.2 Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. Guinot D (1969a) Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures. VII. Les Goneplacidae. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 41, 241–265, pl. 1. Guinot D (1969b) Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures. VII. Les Goneplacidae (suite et fin). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 41, 688–724, pls 1–5. Guinot D (1969c) Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures. VII. Les Goneplacidae (suite). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 41, 507–528.

Karasawa H, Kato H (2003) The phylogeny, systematics and fossil record of the Goneplacidae MacLeay (Crustacea, Decapoda, Brachyura) revisited. Contributions to Zoology 72, 147–152. doi:10.1163/18759866-0720203016 Karasawa H, Schweitzer CE (2006) A new classification of the Xanthoidea sensu lato (Crustacea: Decapoda: Brachyura) based on phylogenetic analysis and traditional systematics and evaluation of all fossil Xanthoidea sensu lato. Contributions to Zoology 75, 23–73. doi:10.1163/18759866-0750102002 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Tsang LM, Schubart CD, Ahyong ST, Lai JCY, Au EYC, et al. (2014) Evolutionary history of true crabs (Crustacea: Decapoda: Brachyura) and the origin of freshwater crabs. Molecular Biology and Evolution 31, 1173–1187. doi:10.1093/molbev/msu068

Acidopsidae Števčić, 2005 Figure 14.41, Plate 43m–o Acidopsids are typically small crabs primarily from reefs of the Indo-West Pacific where they range from the intertidal to

512

Marine Decapod Crustacea

Fig. 14.41.  Acidopsidae. Carapace: a, Raoulia limosa Ng, 1987. Habitus: b, Thecaplax capillosa Ng and Rahayu, 2018. Antennular basal article (A), antennal articles 1–3: c, Acidops; d, Raoulia. Male, right half of pleonites 1–4, thoracic sternites 7, 8, coxae 4, 5, genital papilla (shaded): e, Acidops; f, Raoulia; g, Thecaplax; h, Typhlocarcinodes. Female, thoracic sternites 4–6, paired vulvae: i, Acidops; j, Raoulia. Male, thoracic sternites 4, 5, 6: k, Caecopilumnus; l, Raoulia. Male pleon: m, Caecopilumnus; n, Raoulia. Right front, orbit, eyestalk, anterior view: o, Acidops; p, Crinitocinus; q, Raoulia; r, Typhlocarcinodes. Maxilliped 3: s, Acidops; t, Raoulia. Chela fingers: u, Acidops; v, Crinitocinus; w, Parapilumnus. Gonopods 1, 2: x, Raoulia; y, Thecaplax.

shallow shelf depth. Many acidopsids are coarsely setose and superficially resemble pilumnids, with which they were previously been classified. Only one acidopsid genus is found in the tropical Atlantic. The taxonomy of the family was reviewed by Ng and Rahayu (2014) who recognised two subfamilies. Diagnosis. Carapace surface smooth, granular or at most with few low ridges, setose; anterolateral margin unarmed or at most with low, coarsely granular convex lobes; without prominent teeth. Male thoracic sternite 8 without

supplementary plate, male genital papilla exposed, lying in open groove between thoracic sternites 7 and 8 (Fig. 14.41e– h). Male pleon triangular; pleonites 3–5 fused although demarcations partially to fully visible (Fig. 14.41m, n). Eyestalk short, with conspicuous cornea. Maxilliped 3 merus anterolateral margin obtusely angular, or rounded. Pereopods 2–5 flexor and extensor margins fringed with setae. Female vulva with simple margin or single vulvar cover (Fig. 14.41i, j). Gonopod 2 longer than or as long as gonopod 1 (Fig. 14.41x, y).

Key to subfamilies and genera of Acidopsidae 1.

Maxilliped 3 merus distolateral margin angular (Fig. 14.41s). Antennal article 1 quadrate (Fig. 14.41c). Genital papilla coxal, exiting coxa 5 directly into sterno-pleonal cavity, protected by pleonite 3 when

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2. – 3. – 4. – 5. – 6.



513

pleon closed. Male thoracic sternite 8 visible as small plate when pleon closed (Fig. 14.41e). Vulvae close to midline of thoracic sternite 6 (Fig. 14.41i). Male sternal ‘press-button’ on posterior half of sternite 4����������������������������������������������������������������������������������������������������������������������������������������������Acidopsinae … 2 Maxilliped 3 merus distolateral margin convex (Fig. 14.41t). Antennal article 1 wider than long (Fig. 14.41d). Genital papilla coxo-sternal, exiting coxa 5 directly under or on groove between thoracic sternites 7, 8, partly exposed, unprotected by sternites or pleon. Male thoracic sternite 8 visible as large subrectangular plate when pleon closed (Fig. 14.41f–h). Vulvae distant from midline of thoracic sternite 6 (Fig. 14.41j). Male sternal ‘press-button’ on anterior half of sternite 4������������������������������������������������� Raouliinae … 4 Cheliped fingers cutting edges blade-like (Fig. 14.41w)��������������������������������������������������������������������������������������Parapilumnus Cheliped fingers cutting edges dentate (Fig. 14.41u, v)��������������������������������������������������������������������������������������������������������������3 Eye as wide as peduncle, with prominent cornea (Fig. 14.41p). Major cheliped dactylus with prominent proximal cutting tooth (Fig. 14.41v)���������������������������������������������������������������������������������������������������������������������Crinitocinus Eye tapering, cornea reduced (Fig. 14.41o). Major cheliped dactylus with large and small teeth (Fig. 14.41u)����������������������������������������������������������������������������������������������������������������������������������������������������������������������Acidops Eye mobile. Epistome partially sunken (Fig. 14.41r). Male genital papilla protected by mobile plate in groove between thoracic sternites 7, 8 (Fig. 14.41h)���������������������������������������������������������������������������������� Typhlocarcinodes Eye fused to orbit. Epistome prominently sunken (Fig. 14.41q). Male genital papilla at least partly exposed in groove between thoracic sternites 7, 8 (Fig. 14.41f, g)�������������������������������������������������������������������������������������������������������5 Male thoracic sternite 4 about as wide as long (Fig. 14.20k). Pleonite 1 subrectangular, lateral margins straight or gently convex (Fig. 14.41m). Male genital papilla partially exposed on long groove between thoracic sternites 7, 8�����������������������������������������������������������������������������������������������������������������������������������������Caecopilumnus Male thoracic sternite 4 wider than long (Fig. 14.41l). Pleonite 1 with lateral part tapering to acute triangle (Fig. 14.41n). Male genital papilla completely or partially exposed on long groove between thoracic sternites 7, 8 (Fig. 14.41f, g)��������������������������������������������������������������������������������������������������������������������������������������������������������6 Carapace anterolateral margin entire, without teeth, spines or lobes (Fig. 14.41a). Male genital papilla exposed over most of distal length between thoracic sternites 7, 8 (Fig. 14.41f). Gonopod 1 with stout basal half, distal half sharply tapering to narrow tip with scattered subdistal spinules (Fig. 14.41x)�������������������������������������������������������������������������������������������������������������������������������������������������������Raoulia Carapace anterolateral margin with granulate lobes (Fig. 14.41b). Male genital papilla completely exposed in groove between thoracic sternites 7, 8 (Fig. 14.41g). Gonopod 1 gradually tapering towards tip, with numerous subdistal spinules (Fig. 14.41y)�������������������������������������������������������������������������������������������������������������� Thecaplax

Subfamily Acidopsinae Števčić, 2005

Acidops Stimpson, 1871

Diagnosis. Male thoracic sternite 8 visible as small plate when pleon closed (Fig. 14.41e). Vulvae close to midline of thoracic sternite 6 Maxilliped 3 merus distolateral margin angular. Antennal article 1 quadrate. Male genital papilla coxal, exiting coxa 5 directly into sterno-pleonal cavity, protected by pleonite 3 when pleon closed.

Diagnosis. Acidopsinae. Eye tapering, cornea reduced. Chelae fingers cutting edges dentate. Major cheliped dactylus with large and small teeth. Maximum cl. 11 mm. Intertidal–shelf (0–135 m; reef, sand). Tropical Atlantic, Tropical Eastern Pacific. 2 species (Ng and Rahayu 2014: rediagnosis, differences between species).

Crinitocinus Ng & Rahayu, 2014

Parapilumnus Kossmann, 1877

Diagnosis. Acidopsinae. Eye as wide as peduncle, with prominent cornea. Chelae fingers cutting edges dentate. Major cheliped dactylus with prominent proximal cutting tooth. Maximum cl. 9 mm (Pl. 43m). Shelf (20–71 m). Western and Central Indo-Pacific. 1 species (Ng and Rahayu 2014).

Diagnosis. Acidopsinae. Eye as wide as peduncle, with prominent cornea. Chelae fingers cutting edges blade-like. Major cheliped dactylus with 1 or 2 proximal blunt teeth. Maximum cl. 13 mm (P l. 43n). Subtidal, shelf (1–30 m). Central Indo-Pacific. 2 species (Ng 2002; Ng and Chen 2004; Ng and Rahayu 2014).

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Marine Decapod Crustacea

Subfamily Raouliinae Števčić, 2005 Diagnosis. Male thoracic sternite 8 visible as large subrectangular plate when pleon closed. Vulvae distant from midline of thoracic sternite 6. Maxilliped 3 merus distolateral margin convex. Antennal article 1 wider than long. Male genital papilla coxo-sternal, exiting coxa 5 directly under or on groove between thoracic sternites 7, 8, partly exposed, unprotected by sternites or pleon. Caecopilumnus Borradaile, 1903 Diagnosis. Raouliinae. Carapace anterolateral margin curved, granular, setose, sometimes with low lobes covered with tubercles. Epistome prominently sunken. Male thoracic sternite 4 about as wide as long. Pleonite 1 subrectangular, lateral margins straight or gently convex. Eye fused to orbit. Male genital papilla partially exposed on long groove between thoracic sternites 7, 8. Gonopod 1 stout, with distal spinules. Maximum cl. 11 mm. Intertidal–shelf (0–95 m; reef). Temperate Northern Pacific, Western and Central Indo-Pacific. 3 species (Ng and Rahayu 2014: rediagnosis, description of 3 species).

Raoulia Ng, 1987 Diagnosis. Raouliinae. Carpace anterolateral margin subcristate, entire, without teeth, spines or lobes. Epistome prominently sunken. Male thoracic sternite 4 wider than long. Pleonite 1 with lateral part tapering to acute triangle. Eye fused to orbit. Male genital papilla exposed over most of distal length between thoracic sternites 7, 8. Gonopod 1 with stout basal half, distal half sharply tapering to narrow tip with scattered subdistal spinules. Maximum cl. 9 mm (Pl. 43o). Intertidal–shelf (0–62 m; reef). Indo-West Pacific. 5 species (Ng and Rahayu 2014: rediagnosis, descriptions of 5 species).

Thecaplax Ng & Rahayu, 2014 Diagnosis. Raouliinae. Carpace anterolateral margin with granulate lobes. Epistome prominently sunken. Male thoracic sternite 4 wider than long. Pleonite 1 with lateral part tapering to acute triangle. Eye fused to orbit. Male genital papilla completely exposed in groove between thoracic sternites 7, 8. Gonopod 1 gradually tapering towards tip, with numerous subdistal spinules. Maximum cl. 12 mm. Intertidal. Central Indo-Pacific (Lombok). 1 species (Ng and Rahayu 2014).

Typhlocarcinodes Alcock, 1900 Diagnosis. Raouliinae. Carpace anterolateral margin curved, granular, setose, with 2 low teeth. Epistome partially sunken. Male thoracic sternite 4 wider than long. Pleonite 1 with lateral part tapering to acute triangle. Eye mobile. Male genital papilla protected by mobile plate in groove between thoracic sternites 7, 8. Gonopod 1 stout, with distal spinules. Maximum cl. 7 mm. Subtidal, shelf (12–100 m). Tropical Atlantic (W Africa). 1 species (Ng and Rahayu 2014: rediagnosis).

References Ng PKL (2002) The Indo-Pacific Pilumnidae XVI. On the identity of Pilumnus cristimanus A. Milne Edwards, 1873, and the status of Parapilumnus Kossmann, 1877 (Crustacea: Decapoda: Brachyura), with description of a new species from rubble beds in Guam. Micronesica 34, 209–226. Ng PKL, Chen H-L (2004) On the male of Parapilumnus cristimanus (A. Milne Edwards, 1873), and its presence in the South China Sea (Crustacea: Decapoda: Brachyura: Goneplacidae). Malayan Nature Journal 56, 403–407. Ng PKL, Rahayu DL (2014) Revision of the family Acidopsidae Števčić, 2005, and the systematic position of Typhlocarcinodes Alcock, 1900, Caecopilumnus Borradaile, 1902, and Raoulia Ng, 1987, with descriptions of two new genera and five new species (Crustacea: Brachyura: Goneplacoidea). Zootaxa 3773, 1–63. doi:10.11646/zootaxa.3773.1.1

Chasmocarcinidae Serène, 1964 Figures 14.42, 14.43, Plate 44a–c The Chasmocarcinidae are a group of small compact crabs arrayed in three subfamilies. The genera of Chasmocarcininae and Megaesthesiinae are marine with representatives living from subtidal to deep-sea depths. Members of Trogloplacinae inhabit coastal freshwater and estuarine rivers, but one species, Trogloplax jolliveti Guinot, 1986, is from an inland cave system at 650 m above sea level. At least one freshwater species of Australocarcinus has direct development, the hatched megalopae carried by the female (Davie and Guinot 1996). Ng and Castro (2016) reviewed and diagnosed all genera and species. Their key to subfamilies and genera has been adapted here. The family differs from all other goneplacoids in that the male genital papilla is partially concealed by a calcified structure (termed a supplementary plate) formed by an invagination of the anterior surface of thoracic sternite 8, which lies against thoracic sternite 7. The supplementary plate is otherwise known only in members of the Palicoidea as an independent derivation (Guinot et al. 2013). Diagnosis. Carapace usually widest posteriorly, usually without anterolateral spine; surface smooth or uneven, granular, glabrous or sparsely setose. Male thoracic sternum with ‘press-button’ on anterior half of sternite 5, near suture 4/5 (Fig. 14.42n). Male thoracic sternite 8 with double supplementary coxosternal plate formed by invagination and anterior extension of sternite, largely concealing genital papilla (Fig. 14.42n, o, p). Male pleon triangular; pleonites 3–5 fused although demarcations partially visible. Eyestalk variously developed, usually with conspicuous cornea. Maxilliped 3 merus anterolateral margin obtusely angular. Pereopods 2–5 sparsely setose; merus usually unarmed (rarely with spinous flexor margin). Female vulva with simple margin or single vulvar cover. Gonopod 2 longer than gonopod 1, or slightly shorter than gonopod 1, or as long as gonopod 1 (Fig. 14.43).

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Key to subfamilies and genera of Chasmocarcinidae 1. – 2. – 3. – 4. –

Antennular peduncle articles cylindrical, swollen. Eyestalk long, thin (Fig. 14.42d)����������������������Megaesthesiinae … 3 Antennular peduncle with articles not enlarged (Fig. 14.42a–c, e). Eyestalk compact (thin in cavernicolous species only)���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Antennular peduncle basal article rectangular; flagellum partially folded into fossa (Fig. 14.42e). Vulva large, with thick margins (Fig. 14.42m). Freshwater, estuaries, caves��������������������������������������������� Trogloplacinae … 4 Antennular peduncle basal article round or quadrate; flagellum not folded into fossa (Fig. 14.42a–c). Vulva small, with thin margins. Marine���������������������������������������������������������������������������������������������������Chasmocarcininae … 5 Gonopod 2 longer than gonopod 1; basal, distal articles equally long (Fig. 14.43m, n)���������������������������������Megaesthesius Gonopod 2 about two-thirds length of gonopod 1; basal article short, distal article spatuliform, with short elongation (Fig. 14.43a, b)��������������������������������������������������������������������������������������������������������������������������������������Alainthesius Cornea reduced, non-pigmented (Fig. 14.42e). Walking legs long, slender, subcylindrical, almost glabrous; pereopod 5 ~3 times carapace length. Inhabitant of terrestrial caves���������������������������������������������������������������Trogloplax Cornea pigmented. Walking legs robust, compressed; dactylus and propodus fringed with setae; pereopod 5 shorter than twice carapace length. Inhabitant of rivers, estuaries�����������������������������������������������������Australocarcinus

Fig. 14.42.  Chasmocarcinidae. Right orbit, eyestalk, antennular and antennal peduncles: a, Amboplax; b, Chasmocarcinus; c, Notopelta; d, Megaesthesius; e, Trogloplax. Carapace: f, Chasmocarcinus; g, Deltopelta; h, Megaesthesius. Carapace, front: i, j, Angustopelta; k, Hephthopelta; l, Statommatia. Female sternum with vulvae: m, Australocarcinus. Male sternum: n, Trogloplax (genital papillae shaded). Male thoracic sternites 6–8, pleonites 1–3: o, Camatopsis; p, Microtopsis. Maxilliped 3: q, Chasmocarcinops; r, Hephthopelta. Minor chela: s, Camatopsis; t, Chinommatia. Major chela, ischium merus: u, Angustopelta; v, Statommatia; w, Notopelta. Pereopod 5, propodus, dactylus: x, Chasmocarcinus.

516

Marine Decapod Crustacea

Fig. 14.43.  Chasmocarcinidae. Gonopods 1, 2: a, b, Alainthesius; c, d, Angustopelta; e, f, Camatopsis; g, h, Chasmocarcinus; i, j, Chinommatia; k, l, Hephthopelta; m, n, Megaesthesius; o, Microtopsis (gonopod 1 only); p, q, Statommatia; r, s, Tenagopelta.

5. – 6. – 7.

Eyestalks mobile, sometimes tightly fitting orbit (Fig. 14.42a–c)���������������������������������������������������������������������������������������������6 Eyestalks sessile (Fig. 14.42i–l)�����������������������������������������������������������������������������������������������������������������������������������������������������13 Carapace subtriangular (Fig. 14.42g). Western Atlantic��������������������������������������������������������������������������������������������Deltopelta Carapace subtrapezoidal or subquadrate (Fig. 14.42f)���������������������������������������������������������������������������������������������������������������7 Eyestalks with reniform cornea (Fig. 14.42c). Male major cheliped ischium prolonged on distal margin (Fig. 14.42w)������������������������������������������������������������������������������������������������������������������������������������������������������������������Notopelta – Eyestalks with round cornea. Male major cheliped ischium not prolonged on distal margin��������������������������������������������8 8. Plate between thoracic sternites 7 and 8 extending over lateral half of exposed part of male thoracic sternum, not reaching sterno-pleonal cavity (Fig. 14.42p). Gonopod 1 strongly twisted (Fig. 14.43o)������ Microtopsis – Plate between thoracic sternites 7 and 8 reaching across most of exposed part of male thoracic sternum, reaching sterno-pleonal cavity (Fig. 14.42o). Gonopod 1 straight or slightly twisted�����������������������������������������������������9 9. Suborbital ridge below orbital margin (Fig. 14.42a). Western Atlantic������������������������������������������������������������������ Amboplax – Without suborbital ridge below orbital margin (Fig. 14.42b, c)����������������������������������������������������������������������������������������������10 10. Minor chela fixed finger usually with 1 or 2 sharp teeth on distal portion of cutting margin, submarginal surfaces with comb-like setae (Fig. 14.42s). Gonopod 2 three-quarters length of gonopod 1, with long basal article, short distal article (Fig. 14.43e, f)��������������������������������������������������������������������������������������������������� Camatopsis – Minor chela fixed finger with many small, subequal teeth or 1 blunt tooth, submarginal surfaces without comb-like setae (Fig. 14.42t). Gonopod 2 half or less length or much longer than gonopod 1, basal article short or very long, distal article short or whip-like�������������������������������������������������������������������������������������������������������������11 11. Carapace globose, convex when viewed frontally (e.g. Fig. 14.42k). Gonopod 2 longer than gonopod 1 (Fig. 14.43r, s)������������������������������������������������������������������������������������������������������������������������������������������������������������Tenagopelta – Carapace flat when viewed frontally (e.g. Fig. 14.42j, l). Gonopod 2 as long or shorter than gonopod 1����������������������� 12 12. Pterygostomian region bulging or convex (Fig. 14.42f). Male pereopod 5 sometimes with subdistal spur (Fig. 14.42x). Gonopod 2 three-quarters or as long as gonopod 1 (Fig. 14.43g, h). Americas��������������Chasmocarcinus – Pterygostomian region not bulging. Male pereopod 5 without subdistal spur. Gonopod 2 half as long as gonopod 1 (Fig. 14.43i, j)������������������������������������������������������������������������������������������������������������������������������������� Chinommatia 13. Maxilliped 3 merus elongate, narrower than ischium (Fig. 14.42q)�����������������������������������������������������������Chasmocarcinops – Maxilliped 3 merus ovate, as wide as ischium (Fig. 14.42r)����������������������������������������������������������������������������������������������������14 14. Carapace globose, convex when viewed frontally (Fig. 14.42k). Cheliped merus lower surface with 3 large teeth on outer margin. Gonopod 2 longer than gonopod 1 (Fig. 14.43k, l)������������������������������������������������� Hephthopelta

14 – Brachyura – crabs

– 15. –

517

Carapace flat when viewed frontally (Fig. 14.42l). Cheliped merus lower surface with 4 large teeth or row of tubercles on outer margin. Gonopod 2 less than half as long as gonopod 1 (Fig. 14.43c, d, p, q)��������������������������������15 Cheliped merus lower inner margin with distal tooth (Fig. 14.42u. Carapace anterolateral margin usually with tooth or tubercle (Fig. 14.42i) or not (Fig. 14.42j). Gonopod 1 with constriction between stout basal part and narrower shorter distal part, curved outwards (Fig. 14.43c)�����������������������������������������������������������Angustopelta Cheliped merus lower inner margin unarmed (Fig. 14.42v). Carapace anterolateral margin unarmed (Fig. 14.42l). Gonopod 1 tapering evenly from basal part to longer slender tapering distal part (Fig. 14.43p)������������������������������������������������������������������������������������������������������������������������������������������������������������ Statommatia

Subfamily Chasmocarcininae Serène, 1964 The subfamily contains the great majority of genera and species. It is entirely marine. Diagnosis. Antennular peduncle basal article round or quadrate; flagellum not folded into fossa. Buccal cavity quadrate or rectangular. Eyestalks compact. Implicit generic attributes of Chasmocarcininae. Carapace subquadrate or subtrapezoidal; flat when viewed frontally; anterolateral margin without tooth or tubercle; without suborbital ridge below orbital margin. Plate between thoracic sternites 7 and 8 reaching across most of exposed part of male thoracic sternum, reaching sterno-pleonal cavity. Eyestalks mobile, sometimes tightly fitting orbit; cornea reduced, pigmented. Maxilliped 3 merus ovate, as wide as ischium. Male major cheliped ischium not prolonged on distal margin. Cheliped merus lower surface inner margin unarmed. Pereopods 2, 3 meri with teeth or tubercles. Gonopod 1 straight. Amboplax Ng & Castro, 2016 Diagnosis. Chasmocarcininae. Carapace globose, convex when viewed frontally; with suborbital ridge below orbital margin. Cheliped merus lower surface without teeth. Minor chela fixed finger usually with 1 or 2 sharp teeth on distal portion of cutting margin, submarginal surfaces with comb-like setae. Gonopod 2 longer than gonopod 1. Maximum cl. 5 mm. Subtidal, shelf (15–34 m). Tropical Atlantic (Brazil). 1 species (Ng and Castro 2016).

Angustopelta Ng & Castro, 2016 Diagnosis. Chasmocarcininae. Carapace anterolateral margin with tooth or tubercle, or without tooth or tubercle. Eyestalks sessile, subcylindrical. Cheliped merus lower surface with 4 large teeth; inner margin with distal tooth. Gonopod 1 with constriction between stout basal part and narrower shorter distal part, curved outwards. Gonopod 2 half or less as long as gonopod 1. Maximum cl. 19 mm (Pl. 44a). Slope (266–1775 m). Central Indo-Pacific. 4 species (Ng and Castro 2016: key to species).

Camatopsis Alcock & Anderson, 1899 Diagnosis. Chasmocarcininae. Cheliped merus lower surface without teeth. Minor chela fixed finger usually with 1 or 2

sharp teeth on distal portion of cutting margin, submarginal surfaces with comb-like setae. Gonopod 1 distal segment slightly curved. Gonopod 2 three-quarters length of gonopod 1, with long basal article, short distal article. Maximum cl. 10.5 mm (Pl. 44b). Subtidal–slope (13–695 m). Temperate Northern Pacific, IndoWest Pacific. 6 species (Ng and Castro 2016: key to species).

Chasmocarcinops Alcock, 1900 Diagnosis. Chasmocarcininae. Eyestalks sessile, subcylindrical. Maxilliped 3 merus elongate, narrower than ischium. Cheliped merus lower surface with teeth. Gonopod 2 about one-third length of gonopod 1, slender, distal part long, slightly curved. Maximum cl. 9 mm (Pl. 44c). Subtidal, shelf (9–49 m). Western and Central Indo-Pacific. 1 species (Ng and Castro 2016).

Chasmocarcinus Rathbun, 1898 Diagnosis. Chasmocarcininae. Cheliped merus lower surface without teeth. Minor chela fixed finger with many small, subequal teeth, submarginal surfaces without comb-like setae. Pereopods 2, 3 meri with tubercles, without teeth. Gonopod 2 three-quarters or as long as gonopod 1. Maximum cl. 9.5 mm. Subtidal–slope (9–1967 m). Tropical W Atlantic, Eastern IndoPacific, Temperate South America. 10 species (Ng and Castro 2016: key to species).

Chinommatia Ng & Castro, 2016 Diagnosis. Chasmocarcininae. Carapace pterygostomial region not bulging. Cornea pigmented. Male pereoppod 5 simple, never with subdistal spur. Cheliped merus lower surface with row of sharp tubercles. Gonopod 2 half or less as long as gonopod 1. Maximum cl. 17 mm. Subtidal–slope (10–549 m). Central Indo-Pacific. 5 species (Ng and Castro 2016: key to species).

Deltopelta Ng & Castro, 2016 Diagnosis. Chasmocarcininae. Carapace subtriangular. Cornea pigmented. Cheliped merus lower surface with teeth. Maximum cl. 5.2 mm. Subtidal–slope (17–855 m). Tropical Atlantic (West Indies). 1 species (Ng and Castro 2016).

518

Marine Decapod Crustacea

Hephthopelta Alcock, 1899

Alainthesius Ng & Castro, 2016

Diagnosis. Chasmocarcininae. Carapace globose, convex when viewed frontally; anterolateral margin with tooth or tubercle, or without tooth or tubercle. Eyestalks sessile, subcylindrical. Cheliped merus lower surface with 3 large teeth on outer margin. Gonopod 2 longer than gonopod 1. Maximum cl. 10 mm. Slope (203–1022 m). Western and Central Indo-Pacific. 2 species (Ng and Castro 2016: key to species).

Diagnosis. Megaesthesiinae. Gonopod 2 longer than gonopod 1; basal, distal articles long. Maximum cl. 5.6 mm. Shelf, slope (72–509 m). Western and Central Indo-Pacific. 2 species (Ng and Castro 2016: key to species).

Microtopsis Komai, Ng & Yamada, 2012 Diagnosis. Chasmocarcininae. Plate between thoracic sternites 7 and 8 extending over lateral half of exposed part of male thoracic sternum, not reaching sterno-pleonal cavity. Cheliped merus lower surface with row of small tubercles. Gonopod 1 short, twisted. Gonopod 2 almost as long as gonopod 1. Maximum cl. 3.6 mm. Subtidal, shelf (15–90 m). Central Indo-Pacific. 2 species (Ng and Castro 2016: key to species).

Notopelta Ng & Castro, 2016 Diagnosis. Chasmocarcininae. Carapace globose, convex when viewed frontally. Cornea flat, reniform. Male major cheliped ischium prolonged on distal margin. Cheliped merus lower surface with teeth. Gonopod 2 three-quarters length of gonopod 1, with long basal article, short distal article. Maximum cl. 6 mm. Subtidal, shelf (17–75 m). Central Indo-Pacific. 1 species (Ng and Castro 2016).

Statommatia Ng & Castro, 2016 Diagnosis. Chasmocarcininae. Carapace slightly convex when viewed frontally. Eyestalks sessile, subcylindrical. Cheliped merus lower surface with teeth, or with row of small tubercles. Gonopod 1 tapering evenly from basal part to longer slender tapering distal part. Gonopod 2 half or less as long as gonopod 1. Maximum cl. 10 mm. Shelf, slope (80–866 m). Western Indo-Pacific, Central IndoPacific. 5 species (Ng and Castro 2016: key to species).

Tenagopelta Ng & Castro, 2016 Diagnosis. Chasmocarcininae. Carapace globose, convex when viewed frontally. Cheliped merus lower surface with 3 large teeth on outer margin. Gonopod 2 longer than gonopod 1. Maximum cl. 15 mm. Shelf, slope (100–970 m). Central Indo-Pacific. 3 species (Ng and Castro 2016: key to species).

Subfamily Megaesthesiinae Števčić 2005 The subfamily contains only six species in two genera, all marine. Diagnosis. Antennular peduncle articles cylindrical, swollen. Buccal cavity arched, sunken. Vulva large, with thick margins. Eyestalks long, immobile, cornea long.

Megaesthesius Rathbun, 1909 Diagnosis. Megaesthesiinae. Gonopod 2 about two-thirds length of gonopod 1; basal article short, distal article spatuliform, with short elongation. Maximum cl. 3.5 mm. Subtidal, shelf (4–250 m). Temperate Northern W Pacific, Central Indo-Pacific. 4 species (Davie 2013; Ng and Castro 2016: key to species).

Subfamily Trogloplacinae Guinot, 1986 The subfamily contains five species in two genera. One genus is confined to fresh water and may appear in upper estuarine environments (Davie and Guinot 1996). The other is from caves high in the karst mountains of Papua New Guinea (Guinot 1986). Diagnosis. Antennular peduncle basal article rectangular; flagellum partially folded into fossa. Buccal cavity quadrate or rectangular. Vulva small, with thin margins. Eyestalks compact. Australocarcinus Davie, 1988 Diagnosis. Trogloplacinae. Cornea spherical, pigmented. Walking legs robust, compressed; dactylus and propodus fringed with setae; pereopod 5 shorter than twice carapace length. Maximum cl. 11 mm. Upper estuary, freshwater rivers (N Qld, Australia; New Caledonia; Palau). 4 species (Davie and Guinot 1996; Ng and Castro 2016: key to species).

Trogloplax Guinot, 1986 Diagnosis. Trogloplacinae. Cornea reduced, non-pigmented. Walking legs long, slender, subcylindrical, almost glabrous; pereopod 5 about triple carapace length. Maximum cl. 20 mm. Inside cave, 650 m asl (Papua New Guinea). 1 species (Guinot 1986; Ng and Castro 2016). References Davie PJF (2013) A new species of Megaesthesius Rathbun, 1909 (Crustacea: Brachyura: Chasmocarcinidae) from deep water off Western Australia. Zootaxa 3647, 470–478. doi:10.11646/zootaxa.3647.3.4 Davie PJF, Guinot D (1996) Two new freshwater crabs in Australocarcinus Davie, with remarks on Trogoplacinae Guinot and Goneplacidae Macleay (Crustacea: Decapoda: Brachyura). Memoirs of the Queensland Museum 39, 277–287. Guinot D (1986) Description d’un crabe cavernicole aveugle de NouvelleBretagne (Papouasie Nouvelle-Guinée), Trogloplax joliveti gen. nov. sp.

14 – Brachyura – crabs

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margins unarmed, sparsely setose. Female vulva with simple margin or single vulvar cover. Gonopod 2 as long as gonopod 1 (Fig. 14.44c, d). Conleyus Ng & Ng, 2003 Shelf (30–70 m; buried 1 m deep in coral rubble). Central IndoPacific (Guam). 1 species. Maximum cl. 12 mm. Reference Ng PKL, Ng NK (2003) Conleyus defodio, a new genus and new species of carcinoplacine crab (Crustacea: Brachyura: Goneplacidae) from deep rubble beds in Guam. Micronesica 35, 431–439.

Euryplacidae Stimpson, 1871

Fig. 14.44.  Conleyidae. Conleyus defodio Ng & Ng, 2003. a, right carapace; b, maxilliped 3; c, d, gonopods 1, 2. e, male sternum. nov., et établissement d’une sous-famille nouvelle, Trogloplacinae subfam. nov. Comptes Rendus de l’Académie, Paris, Série 3 303, 307–312. Guinot D, Tavares M, Castro P (2013) Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa. Zootaxa 3665, 1–414. doi:10.11646/zootaxa.3665.1.1 Ng PKL, Castro P (2016) Revision of the family Chasmocarcinidae Serène, 1964 (Crustacea, Brachyura, Goneplacoidea). Zootaxa 4209, 1–182. doi:10.11646/zootaxa.4209.1.1

Conleyidae Števčić, 2005 Figure 14.44 The family is confined to a single species, Conleyus defodio Ng & Ng, 2003, almost blind, found buried deep in coral rubble (Ng and Ng 2003). Diagnosis. Carapace and pereopods granular, uneven, few low granular ridges, sparsely setose; 3 short, granular anterolateral teeth (Fig. 14.44a). Male thoracic sternum with ‘pressbutton’ on posterior half of sternite 5, near suture 5/6. Male thoracic sternite 8 without supplementary plate, male genital papilla exposed, lying in open groove between thoracic sternites 7 and 8 (Fig.  14.44e). Male pleon broadly triangular; pleonites 1–6 freely articulating; telson semicircular. Eyestalk reduced, in short orbit; first anterolateral tooth small. Maxilliped 3 merus anterolateral margin produced to acute, triangular lobe (Fig. 14.44b). Pereopods 2–5 extensor and flexor

Figure 14.45, Plate 44d–f Euryplacids are goneplacoid crabs from subtidal to upper slope depths. Most are wider than long, with a smooth carapace and few anterolateral teeth, and are outwardly most similar to members of Goneplacidae. Castro and Ng (2010b) reviewed the taxonomy of all genera and species of Euryplacidae in detail. Their key focussed on the form of the male pleon, said to be either triangular or T-shaped. This dichotomy, used in their key, is difficult to interpret and the shapes of the pleon vary, as they stated, ‘almost as a continuum’ (e.g. Fig. 14.45u–w). However, the genera can be separated on the shape of the carapace and this feature is relied on in this key. Diagnosis. Carapace widest near or anterior to midlength, posterolateral margins converging; with 1 or more anterolateral spines; surface smooth finely granular, glabrous or sparsely setose. Male thoracic sternum with ‘press-button’ on anterior half of sternite 5, near suture 4/5. Male thoracic sternite 8 without supplementary plate, male genital papilla exposed, lying in open groove between thoracic sternites 7 and 8 (Fig.  14.45p). Male pleon reaching anteriorly onto thoracic sternite 4 beyond level of cheliped coxa articular condyle pleonites 1–6 freely articulating; telson lateral margins straight to convex (Fig. 14.45u–w). Eyestalk no longer than width of front. Maxilliped 3 merus anterolateral margin obtusely angular. Pereopods 2–5 extensor and flexor margins unarmed, sparsely setose. Female vulva with simple margin or single vulvar cover. Gonopod 2 0.3 or more length of gonopod 1 (Fig. 14.45q–t). Implied generic attributes. Carapace clearly wider than long, smooth; front not projecting. Cornea spherical. Gonopod 1 with acute apex.

Key to genera of Euryplacidae 1. –

Carapace anterolateral margin with 1 sharp, isolated, oblique spine��������������������������������������������������������������������������������������2 Carapace anterolateral margin with 1–3 teeth, never solitary or spine-like���������������������������������������������������������������������������4

520

2. – 3. – 4. – 5. – 6. –

Marine Decapod Crustacea

Carapace exorbital angle produced as sharp tooth or spine; orbit longer than front, supraorbital margin convex (Fig. 14.45c)������������������������������������������������������������������������������������������������������������������������������������������������������� Frevillea Carapace exorbital angle obsolete or blunt tooth; orbit shorter than front, supraorbital margin straight or concave������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Carapace anterolateral margins oblique, with 1 long (one-eighth carapace width), sharp laterally-directed spine. Cornea spherical (Fig. 14.45h)����������������������������������������������������������������������������������������������������������������������Nancyplax Carapace anterolateral margins subparallel, with 1 short (one-twentieth carapace width), sharp oblique spine. Cornea reniform (Fig. 14.45n)�������������������������������������������������������������������������������������������������������������������������Villoplax Carapace anterolateral margin with 1 small tooth close to square exorbital angle; orbit as long or longer than front, usually with defining dorsal ridge (Fig. 14.45d)������������������������������������������������������������������������������Henicoplax Carapace anterolateral margin with 2 or more teeth posterior to exorbital angle; orbit longer or shorter than front, without defining dorsal ridge�������������������������������������������������������������������������������������������������������������������������������5 Carapace anterolateral margin with 2 obsolete teeth; almost circular, slightly wider than long, surface granular (Fig. 14.45k)�������������������������������������������������������������������������������������������������������������������������������������������������Systroplax Carapace anterolateral margin with 2 or more well defined teeth; subtrapezoidal, posterolateral and posterior margins more or less defined, slightly or clearly wider than long, surface smooth����������������������������������������6 Carapace anterolateral margin convex, with small notch separating 2 lobes, 1 sharp tooth posteriorly; surface with large red dorsal circle (Fig. 14.45j). Gonopod 1 with arrowhead-shaped apex (Fig. 14.45s)����������������������������������������������������������������������������������������������������������������������������������������������������������� Psopheticoides Carapace anterolateral margin convex or almost straight, with 2 or 3 teeth posterior to exorbital angle;surface without large red dorsal circle. Gonopod 1 with sharp apex (Fig. 14.45q)������������������������������������������������7

Fig. 14.45.  Euryplacidae. Carapace: a, Eucrate; b, Euryplax; c, Frevillea; d, Henicoplax; e, Heteroplax; f, Machaerus; g, Platyozius; h, Nancyplax; i, Nectopanope; j, Psopheticoides; k, Systroplax; l, Trissoplax; m, Trizocarcinus; n, Villoplax; o, Xenocrate. p, male thoracic sternites 4–8, pleonite 1 (genital papillae shaded), Eucrate. Gonopods 1, 2 (with detail of apices): q, r, Eucrate; s, t, Psopheticoides. Male pleon: u, Eucrate; v, Nancyplax; w, Trizocarcinus.

14 – Brachyura – crabs

521

7. – 8.

Carapace with postorbital ridge curving parallel to anterior front and orbits (Fig. 14.45g)��������������������������������Platyozius Carapace without postorbital ridge�����������������������������������������������������������������������������������������������������������������������������������������������8 Carapace little wider than long; anterolateral margin almost straight, oblique, obsolete exorbital angle and 2 sharp teeth equidistant. Front shelf-like, projecting (Fig. 14.45o)������������������������������������������������������������ Xenocrate – Carapace clearly wider than long; anterolateral margins convex or almost parallel, with 2 or 3 teeth, exorbital angle tooth-like; front not projecting����������������������������������������������������������������������������������������������������������������������9 9. Carapace anterolateral margin about as long as posterolateral margin, with sharp exorbital tooth followed by 2 well spaced teeth (Fig. 14.45m)���������������������������������������������������������������������������������������������������Trizocarcinus – Carapace anterolateral margin shorter than posterolateral margin, with obsolete or triangular exorbital tooth followed by 2 or 3 contiguous teeth�����������������������������������������������������������������������������������������������������������������������������10 10. Carapace anterolateral margin oblique, convex, greatest width more than 1.3 times distance between exorbital teeth����������������������������������������������������������������������������������������������������������������������������������������������������������������������������11 – Carapace anterolateral margins subparallel, greatest width little more than distance between exorbital teeth���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������13 11. Orbit almost as long as front. Eyestalk nearly half front, much longer than cornea (Fig. 14.45l)�����������������������Trissoplax – Orbit shorter than front. Eyestalk much shorter than front, slightly longer than cornea (Fig. 14.45a)�������������������������� 12 12. Carapace anterolateral margin with 2 teeth posterior to exorbital tooth, first truncate, second sharp (Fig. 14.145i)������������������������������������������������������������������������������������������������������������������������������������������������������������ Nectopanope – Carapace anterolateral margin with 3 teeth posterior to exorbital tooth, first obtuse, blunt, second slightly prominent to acute, third small, blunt or obsolete (Fig. 14.45a)������������������������������������������������������������������������������ Eucrate 13. Carapace anterolateral margin with 2 teeth, second sharper, exorbital angle scarcely protecting cornea laterally; front as wide as half carapace greatest width (Fig. 14.45b)������������������������������������������������������������������� Euryplax – Carapace anterolateral margin with 3 teeth, second largest, exorbital angle lateral to cornea; front as wide as one-third carapace greatest width�������������������������������������������������������������������������������������������������������������������������������������14 14. Carapace anterolateral margin with first tooth minute. Cornea much shorter than peduncle (Fig. 14.45e)����������� Heteroplax – Carapace anterolateral margin with 3 large teeth, second blunt. Cornea slightly shorter than peduncle (Fig. 14.45f)����������������������������������������������������������������������������������������������������������������������������������������������������������������Machaerus Eucrate De Haan, 1835 Diagnosis. Carapace little wider than long; anterolateral margin convex, exorbital angle obsolete, first tooth obtuse, blunt, second tooth slightly prominent to acute, third small, blunt. Orbit shorter than front. Eyestalk much shorter than front, slightly longer than cornea. Maximum cl. 38 mm. Subtidal–slope (1–500 m). Western and Central Indo-Pacific, Temperate Australasia (S Qld, Australia). 8 species (Castro and Ng 2010b: key to species). Eucrate crenata (De Haan, 1835) has been introduced to the Mediterranean (Galil 1997).

Euryplax Stimpson, 1859 Diagnosis. Carapace anterolateral margin with 2 teeth, second sharper, exorbital angle scarcely protecting cornea laterally. Cornea slightly shorter than peduncle. Maximum cl. 17 mm. Subtidal, shelf (1–90 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Castro and Ng 2010b).

Frevillea A. Milne-Edwards, 1880 Diagnosis. Carapace little wider than long; orbit longer than front, supraorbital margin convex; anterolateral margin parallel,

with 1 short (one-twentieth carapace width), sharp oblique spine; exorbital angle produced as sharp tooth or spine. Cornea much shorter than peduncle. Gonopod 1 with acute apex. Maximum cl. 19 mm. Subtidal–slope (1–476 m). Tropical W Atlantic. 3 species (Abele and Kim 1986; Castro and Ng 2010b).

Henicoplax Castro & Ng, 2010 Diagnosis. Orbit as long or longer than front, usually with defining dorsal ridge. Carapace anterolateral margin with 1 small tooth close to square exorbital angle. Cornea much shorter than peduncle. Maximum cl. 6 mm. Subtidal, shelf (1–37 m). Western and Central Indo-Pacific. 5 species (Castro and Ng 2010b: table of species).

Heteroplax Stimpson, 1858 Diagnosis. Carapace anterolateral margin with 3 teeth, first minute, second largest, exorbital angle lateral to cornea. Cornea much shorter than peduncle. Maximum cl. 8 mm. Subtidal, shelf (18–50 m). Central Indo-Pacific (Japan, Hong Kong). 1 species (Castro and Ng 2010b).

522

Marine Decapod Crustacea

Machaerus Leach, 1818 Diagnosis. Carapace anterolateral margin with 3 teeth, second blunt, largest, exorbital angle lateral to cornea. Cornea slightly shorter than peduncle. Maximum cl. 23 mm. Subtidal–slope (4–400 m). Tropical E Atlantic. 2 species (Castro and Ng 2010b).

Nancyplax Lemaitre, García-Gómez, von Sternberg & Campos, 2001 Diagnosis. Carapace little wider than long; orbit shorter than front, supraorbital margin concave. Anterolateral margin parallel, with 1 short (one-twentieth carapace width), sharp oblique spine. Cornea about as long as peduncle. Maximum cl. 12 mm. Shelf (55–155 m). Tropical Atlantic (Caribbean). 1 species (Castro and Ng 2010b; Lemaitre et al. 2001).

Nectopanope Wood-Mason & Alcock, 1891 Diagnosis. Carapace anterolateral margin convex, with 1 truncate tooth, 1 sharp tooth. Cornea subreniform, flattened; about as long as peduncle. Maximum cl. 15 mm. Shelf (200 m). Western Indo-Pacific. 1 species (Ng et al. 2019: rediagnosis, figures).

Platyozius Borradaile, 1902 Diagnosis. Carapace little wider than long; postorbital ridge curving parallel to anterior front and orbits; anterolateral margin with 3 short, square teeth. Cornea about as long as peduncle, or slightly shorter than peduncle. Maximum cl. 8.5 mm (Pl. 44d). Intertidal–shelf (0–200 m). Indo-West Pacific. 1 species (Castro 2020; Castro and Ng 2010b).

Psopheticoides Sakai, 1969 Diagnosis. Carapace clearly wider than long, with large red dorsal circle; anterolateral margin convex, with small notch separating 2 lobes, 1 sharp tooth posteriorly. Cornea reniform; longer than peduncle. Gonopod 1 with arrowhead-shaped apex. Maximum cl. 22 mm. Shelf, slope (50–322 m). Temperate Northern W Pacific, Central Indo-Pacific. 1 species (Castro and Ng 2010b).

Systroplax Castro & Ng, 2010 Diagnosis. Carapace almost circular, slightly wider than long, surface granular; anterolateral margin with 2 obsolete teeth. Cornea about as long as peduncle. Maximum cl. 32 mm. Shelf–slope (165–282 m). Central Indo-Pacific. 1 species (Castro and Ng 2010a, 2010b).

Trissoplax Castro & Ng, 2010 Diagnosis. Carapace anterolateral margin straight, exorbital angle triangular, first tooth angular, second tooth most prominent,

third small, blunt. Orbit almost as long as front. Eyestalk nearly half frontal width, much longer than cornea. Maximum cl. 20 mm (Pl. 44e). Subtidal, shelf (1–40 m). Western and Central Indo-Pacific, Temperate Southern Africa. 2 species (Castro and Ng 2010b).

Trizocarcinus Rathbun, 1914 Diagnosis. Carapace anterolateral margin about as long as posterolateral margin, with 2 well spaced teeth, second sharper, exorbital tooth sharp. Cornea about as long as peduncle. Maximum cl. 21 mm. Shelf, slope (36–462 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Alves-Júnior et al. 2020; Castro and Ng 2010b).

Villoplax Castro & Ng, 2010 Diagnosis. Orbit shorter than front, supraorbital margin concave. Carapace anterolateral margin oblique, with 1 long (one-eighth carapace width), sharp laterally-directed spine; exorbital angle produced as sharp tooth or spine. Cornea reniform, about as long as peduncle. Maximum cl. 20 mm. Subtidal, shelf (1–144 m). Tropical Eastern Pacific. 1 species (Castro and Ng 2010b).

Xenocrate Ng & Castro, 2007 Diagnosis. Carapace little wider than long; front shelf-like, projecting; anterolateral margin almost straight, oblique, obsolete exorbital angle and 2 sharp teeth equidistant. Cornea about as long as peduncle. Maximum cl. 40 mm (Pl. 44f). Shelf, slope (80–300 m). Central Indo-Pacific. 1 species (Castro and Ng 2010b; Ng and Castro 2007). References Abele LG, Kim W (1986) An illustrated guide to the marine decapods crustaceans of Florida. State of Florida Department of Environmental Regulation Technical Series 8(1), 1–760. Alves-Júnior FdA, Ng PKL, Castro P, Mantelatto FL, Souza-Filho JF (2020) Extension of geographical range and first record of Trizocarcinus Rathbun, 1914 (Brachyura: Euryplacidae) from the Western Tropical South Atlantic. Nauplius 28, e2020022. doi:10.1590/2358-2936e2020022 Castro P (2020) Brachyuran crabs (Crustacea: Brachyura) of eleven families of Dorippoidea, Goneplacoidea, Homoloidea, Palicoidea, Pilumnoidea, and Trapezioidea from Papua New Guinea. In: Corbari L, Chan T-Y and Ahyong ST (Eds) Deep-sea crustaceans from Papua New Guinea. Tropical Deep-Sea Benthos vol. 31. Mémoires du Muséum National d’Histoire Naturelle, Paris 213, 141–205. Castro P, Ng PKL (2010a) A new genus and species of goneplacid crab (Decapoda, Brachyura, Goneplacidae) from the Western Pacific. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 51–60. doi:10.1163/ej.9789004170865.i-366.40 Castro P, Ng PKL (2010b) Revision of the family Euryplacidae Stimpson, 1871 (Crustacea: Decapoda: Brachyura: Goneplacoidea). Zootaxa 2375, 1–130. doi:10.11646/zootaxa.2375.1.1

14 – Brachyura – crabs

Galil BS (1997) Two Lessepsian migrant decapods new to the coast of Israel. Crustaceana 70, 111–114. doi:10.1163/156854097X00393 Lemaitre R, García-Gómez J, Sternberg RV, Campos NH (2001) A new genus and a new species of crab of the family Goneplacidae MacCleay, 1838 (Crustacea: Decapoda: Brachyura) from the tropical western Atlantic. Proceedings of the Biological Society of Washington 114, 951–963. Ng PKL, Castro P (2007) On a new genus and species of euryplacid crab (Crustacea: Decapoda: Brachyura: Goneplacoidea) from the Philippines. Zootaxa 1549, 43–53. doi:10.11646/zootaxa.1549.1.3 Ng PKL, Priyaja P, Kumar AB, Devi SS (2019) A collection of crabs (Crustacea, Brachyura) from the southwestern coast of India, with a discussion of the systematic position of Nectopanope Wood-Mason in Wood-Mason & Alcock, 1891 (Euryplacidae). ZooKeys 818, 1–24. doi:10.3897/zookeys.818.32108

Goneplacidae MacLeay, 1838 Figures 14.46–14.48, Plate 44g–i Goneplacid crabs occur on soft, open substrates from the shore to slope depths but are most common in deeper shelf waters. Most are small burrowing species with long eyestalks and a trapezoid carapace, but larger species are heavy-bodied with short-eyestalks, resembling progeryonids and mathildellids. Castro’s (2007) key to genera (of Goneplacinae) relies in part on the relative size of the eyestalks, front and carapace width, and general descriptors of carapace shape. For the uninitiated, some of these characters can be difficult to use where differences are subtle. The difference between ‘reniform’ and ‘spherical’ cornea are also not especially clear to a novice trying to identify members of this difficult family. Our key has been significantly reorganised largely using the information in Castro’s (2007) major revision, and includes three genera erected since 2007. Castro (2007) excluded the monotypic Bathyplax A. Milne-Edwards, 1880 (Fig. 14.47a), from Goneplacidae. The systematic position of the only

523

species has generated considerable debate (Castro 2007; Guinot 1969; Tavares 1996), including assignment to its own subfamily (Števčić 2005). Although Bathyplax differs from most other goneplacids in having a short gonopod 2 and several unusual features, its carapace is virtually indistinguishable from some species of Carcinoplax and, at present, is best accommodated in Goneplacidae. A poorly known monotypic genus previously assigned to the Panopeidae, Homoioplax Rathbun, 1914 (type species, Pseudorhombilia haswelli Miers, 1884), from the Arafura Sea and vicinity, was recently determined to be a based on juveniles of Carcinoplax (Ng et al. 2022). Diagnosis. Carapace and pereopods smooth or uneven, granular, or at most with few low ridges; surfaces glabrous or sparsely setose. Male thoracic sternum with ‘press-button’ on anterior half of sternite 5, near suture 4/5. Male thoracic sternite 8 without supplementary plate (Fig. 14.48b), male genital papilla exposed, lying in open groove between thoracic sternites 7 and 8 (Fig. 14.48a). Male pleon broadly triangular or narrow, linguiform; pleonites 1–6 freely articulating or (rarely) with pleonites 3–5 fused. Eyestalk variously developed, with conspicuous cornea. Maxilliped 3 merus anterolateral margin obtusely angular. Pereopods 2–5 extensor and flexor margins unarmed, sparsely setose. Female vulva with simple margin or single vulvar cover. Gonopod 2 greater than or equal to 0.4 length of gonopod 1, usually longer than gonopod 1. Implicit generic attributes. Carapace, pereopods, eye peduncles without setae. Antennules separated by septum. Cornea prominent. Male pleonites 3–5 free; telson wider than long. Right cheliped dactylus cutting edge with even dentition. Pereopods 2–5 meri extensor margin each without distal tooth. Pereopod 5 dactylus slender.

Key to genera of Goneplacidae See notes below

1. – 2. – 3. – 4.

Carapace anterolateral margin without teeth1 behind exorbital angle; exorbital angle obsolete, tooth-like or dominating lateral margin (Fig. 14.47f, g, l, i, m–o, r)�����������������������������������������������������������������������������������������������������������2 Carapace anterolateral margin with 1 or 2 teeth2 behind exorbital angle; exorbital angle obsolete or tooth-like, never dominating lateral margin��������������������������������������������������������������������������������������������������������������������������9 Carapace squarish-trapezoid; orbits transverse; exorbital angle obsolete or tooth-like�������������������������������������������������������3 Carapace much wider than long; orbits oblique, sloping posteriorly; exorbital angle dominating lateral margin�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������5 Carapace quadrate, parallel-sided over most of length; anterolateral margin convex, directed anterolaterally. Cornea narrower than eyestalk (Fig. 14.47m)��������������������������������������������������������������������������������Notonyx Carapace trapezoidal, tapering posteriorly; anterolateral margin convex or straight, directed laterally. Cornea wider than eyestalk�������������������������������������������������������������������������������������������������������������������������������������������������������4 Carapace with convex anterolateral margin; exorbital angle not produced. Eyestalk as long as front width (Fig. 14.47f)����������������������������������������������������������������������������������������������������������������������������������������������������������������� Guinoplax

524

Marine Decapod Crustacea

Fig. 14.46.  Goneplacidae. a, Entricoplax vestita (De Haan, 1835); b, Psopheticus stridulans Wood-Mason, 1892.

Fig. 14.47.  Goneplacidae. Carapace: a, Bathyplax; b, Carcinoplax; c, Exopheticus; d, Goneplacoides; e, Goneplax; f, Guinoplax; g, Hadroplax; h, Menoplax; i, Microgoneplax; j, Neogoneplax costata Castro, 2007; k, Neogoneplax serratipes Castro, 2007; l, Neommatocarcinus; m, Notonyx; n, Ommatocarcinus; o, Paragoneplax; p, Pedroplax; q, Pycnoplax; r, Singhaplax; s, Thyraplax. Male pleon: t, Carcinoplax; u, Entricoplax; v, Neommatocarcinus; w, Microgoneplax; x, Singhaplax.

14 – Brachyura – crabs

525

Fig. 14.48.  Goneplacidae. a, male, end of right thoracic sternites 7, 8, coxa 5, (gonopore shaded), Psopheticus. b, male sternum (sternites 4–8 numbered, telson dashed), Psopheticus. Female sternum, vulvae: c, Carcinoplax; d, Pycnoplax. e, pterygostomium, left cheliped, Bathyplax. f, major chela, Guinoplax. Pereopod 5: g, Paragoneplax; h, Goneplacoides. Gonopods 1, 2: i, Carcinoplax; j, Goneplacoides; k, Menoplax; l, Microgoneplax; m, Neogoneplax; n, Paragoneplax; o, Pedroplax (gonopod 1); p, Psopheticus; q, Pycnoplax; r, Singhaplax; s, Thyraplax.

– 5. – 6. – 7. – 8. – 9. –

Carapace widest at triangular exorbital angle, produced to spine. Eyestalk longer than front width (Fig. 14.47g)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������5 Eye peduncle with distal iridescence. Gonopod 1 distally twisted. Gonopod 2 about half as long as gonopod 1 (Fig. 14.48n)���������������������������������������������������������������������������������������������������������������������������������������������������������Paragoneplax Eye peduncle without distal iridescence. Gonopod 1 weakly curved, without twist. Gonopod 2 slightly longer than gonopod 1 teeth������������������������������������������������������������������������������������������������������������������������������������Hadroplax Eyestalk more than 1.5 times front width, overlapping exorbital tooth (Fig. 14.47l, n)��������������������������������������������������������7 Eyestalk shorter than or no more than 1.2 times front width (Fig. 14.47i, o, r)���������������������������������������������������������������������8 Male pleonites free. Antennules separated by median septum�������������������������������������������������������������������� Ommatocarcinus Male pleonites 3–5 fused (Fig. 14.47v). Antennules not separated by septum���������������������������������������Neommatocarcinus Gonopod 2 much shorter than gonopod 1, with expanded apex (Fig. 14.48l). Male telson subtriangular (Fig. 14.47w)���������������������������������������������������������������������������������������������������������������������������������������������������������Microgoneplax Gonopod 2 as long or longer than gonopod 1 (Fig. 14.48r). Male telson subtriangular or semicircular (Fig. 14.47x)���������������������������������������������������������������������������������������������������������������������������������������������������������������� Singhaplax Carapace anterolateral margin with 1 tooth (Figs 14.46b, 14.47c–e, k, p)�����������������������������������������������������������������������������10 Carapace anterolateral margin with 2 teeth (Figs 14.46a, 14.47b, h, q, s; first or both obsolete if carapace oval)�������������������������������������������������������������������������������������������������������������������������������������������������������������������������15

526

Marine Decapod Crustacea

10.

Carapace little longer than wide, appearing squarish; front half as wide as carapace; anterolateral margin with blunt tooth next to exorbital angle. Eye very short, subglobular (Fig. 14.47p). Gonopod 1 with 2 subdistal spines (Fig. 14.48o)�������������������������������������������������������������������������������������������������������������������������������������Pedroplax – Carapace noticeably wider than long; front about one-third carapace width; anterolateral margin with sharp tooth next to or set back from exorbital angle. Eye elongated, much longer than wide. Gonopod 1 simple������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������11 11. Carapace strongly tapering over posterior three-quarters, with exorbital and anterolateral teeth similar, triangular (Fig. 14.47d). Pereopod 5 dactylus flat, wide (Fig. 14.48h). Gonopod 2 with flagellum apex curled (Fig. 14.48j)����������������������������������������������������������������������������������������������������������������������������������������������� Goneplacoides – Carapace almost rectangular or circular, with exorbital and anterolateral teeth dissimilar. Pereopod 5 dactylus slender. Gonopod 2 with curved flagellum (Fig. 14.48m)���������������������������������������������������������������������������������� 12 12. Gonopod 1 triangular, ~4 times as long as wide (Fig. 14.48p). Pereopods 3–5 meri extensor margins with row of sharp teeth (Fig. 14.46b)������������������������������������������������������������������������������������������������������������������������������ Psopheticus – Gonopod 1 slender, ~6 times as long as wide (Fig. 14.48m). Pereopods 3–5 meri extensor margins without or with 1 distal tooth����������������������������������������������������������������������������������������������������������������������������������������������������������������13 13. Carapace usually with small anterolateral tooth set close to exorbital spine (Fig. 14.47j, k)3��������������������������������� Neogoneplax – Carapace with anterolateral tooth set close to midlength�������������������������������������������������������������������������������������������������������14 14. Carapace ~1.3 times as wide as long, anterolateral and posterolateral margins of similar lengths, obtusely angled (Fig. 14.47c)�������������������������������������������������������������������������������������������������������������������������������������������������� Exopheticus – Carapace ~1.5 times as wide as long, anterolateral margin shorter than posterolateral margins each almost straight, scarcely angled (Fig. 14.47e)������������������������������������������������������������������������������������������������������������������������Goneplax 15. Carapace, pereopods, eye peduncles setose (Fig. 14.46a). Pleonite 1 wider than pleonite 2 (Fig. 14.47u)�����������Entricoplax – Carapace, pereopods, eye peduncles without or with few setae. Pleonite 1 about as wide as pleonite 2 (Fig. 14.47t)���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������16 16. Carapace anterolateral margin with 2 dissimilar teeth, first right-angled, second tooth either much smaller or long, acute (Fig. 14.47h). Gonopod stout, tapering over distal fifth, with distal denticles, truncate (Fig. 14.48k)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������17 – Carapace anterolateral margin with 2 similar teeth or anterior tooth obsolete or absent (Fig. 14.47b, q, s). Gonopod 1 tapering at least over distal half, apex acute, oblique, rarely truncate (Fig. 14.48i, q, s)���������������������������18 17. Carapace exorbital angle an almost right-angled tooth; second anterolateral tooth slender, acute, prominently longer than first tooth (Fig. 14.47h). Gonopod 2 straight (Fig. 14.48k)��������������������������������������� Menoplax – Carapace exorbital angle low, not produced; second anterolateral tooth shorter, smaller than first tooth. Gonopod 2 strongly recurved distally���������������������������������������������������������������������������������������������������������������������Kingelus 18. Carapace slightly wider posterior to larger anterolateral tooth; front scarcely projecting anterior to exorbital angles, eyes hidden laterally (Fig. 14.47s)����������������������������������������������������������������������������������������������� Thyraplax – Carapace widest at second anterolateral tooth; front projecting well anterior to exorbital angles, eyes exposed laterally (Fig. 14.47b, q)���������������������������������������������������������������������������������������������������������������������������������������������19 19. Gonopod 1 longer than gonopod 2. Cheliped merus with stridulatory ridge on inner distal margin (Fig. 14.48e)������������������������������������������������������������������������������������������������������������������������������������������������������������������Bathyplax – Gonopod 2 longer than gonopod 1. Cheliped merus without stridulatory ridge on inner distal margin���������������������� 20 20. Gonopod 1 thin, evenly tapering (Fig. 14.48i). Vulvae half width of corresponding sternite 6, without cover (Fig. 14.48c)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Carcinoplax – Gonopod 1 with wide basal half (Fig. 14.48q). Vulvae less than one-quarter width of corresponding sternite 6, with cover (Fig. 14.48d)�������������������������������������������������������������������������������������������������������������������������� Pycnoplax 1

Ommatocarcinus fibriophthalmus Yokoya, 1933 has a small anterolateral tooth and is notable for eyestalks that are twice the carapace width. Anterolateral teeth may be obsolete in large specimens of Carcinoplax. 3 Neogoneplax costata Castro, 2007 differs from the other two species in the shapes of the carapace, male pleon and gonopods. Its carapace resembles those of Paragoneplax and Singhaplax. 2

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Bathyplax A. Milne-Edwards, 1880 Diagnosis. Carapace widest at second anterolateral tooth; front projecting well anterior to exorbital angles, eyes exposed laterally; anterolateral margin usually with 2 sharp teeth posterior to low exorbital angle. Eyestalk 0.2–0.3 times front width. Cheliped merus with stridulatory ridge on inner distal margin. Gonopod 1 thin, evenly tapering, apex acute. Gonopod 2 about half length of gonopod 1, distal part shorter than basal part. Vulvae fifth width of sternum, with cover. Maximum cl. 26 mm. Shelf, slope (55–1106 m). Tropical W Atlantic. 1 species (Poupin and Corbari 2016; Tavares 1996).

Carcinoplax H. Milne Edwards, 1852 Diagnosis. Carapace widest at second anterolateral tooth; front projecting well anterior to exorbital angles, eyes exposed laterally; anterolateral margin usually with 2 blunt teeth posterior to exorbital tooth (sometimes obsolete in large specimens). Eyestalk 0.2–0.3 times front width. Cheliped merus without stridulatory ridge on inner distal margin. Gonopod 1 thin, evenly tapering, apex acute. Gonopod 2 as long as or longer than gonopod 1, distal part shorter than basal part. Vulvae half width of corresponding sternite 6, without cover. Maximum cl. 54 mm (Pl. 44g). Shelf, slope (17–984 m). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Southern Africa. 25 species (Guinot and Castro 2007: rediagnosis, key to species; Ng and Castro 2020).

Entricoplax Castro, 2007 Diagnosis. Carapace, pereopods, eye peduncles setose; carapace widest behind second anterolateral tooth; front scarcely projecting anterior to exorbital angles, eyes hidden laterally; anterolateral margin with 2 minute teeth posterior to exorbital tooth; exorbital angle obsolete. Eyestalk 0.5 times front width. Gonopod 1 thin, evenly tapering, apex acute. Gonopod 2 longer than gonopod 1, distal part longer than basal part. Maximum cl. 23 mm. Shelf (15–110 m). Temperate Northern W Pacific, Central Indo-Pacific. 1 species (Castro 2007; Shen 1932: figure).

Exopheticus Castro, 2007 Diagnosis. Carapace ~1.3 times as wide as long, anterolateral and posterolateral margins of similar lengths, obtusely angled; anterolateral margin with 1 sharp tooth at midpoint; exorbital angle tooth-like. Eyestalk elongate, 0.5 times front width. Gonopod 1 slender, ~6 times as long as wide. Gonopod 2 about as long as gonopod 1, distal part shorter than basal part. Maximum cl. 30 mm. Shelf, slope (110–418 m). Temperate Northern W Pacific, Central Indo-Pacific. 2 species (Castro 2007: key to species).

Goneplacoides Castro, 2007 Diagnosis. Carapace widest at anterolateral tooth; anterolateral margin with 1 tooth close to and similar to exorbital tooth; exorbital angle tooth-like. Eyestalk 0.8 times front width. Pereopods 2–5 meri extensor margin each with distal tooth. Pereopod 5 dactylus

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flat, wide. Gonopod 1 bent distally. Gonopod 2 with flagellum apex curled. Maximum cl. 16 mm. Shelf, slope (156–660 m). Central Indo-Pacific. 1 species (Castro 2007; Komatsu and Takeda 2003: species description).

Goneplax Leach, 1814 Diagnosis. Carapace ~1.5 times as wide as long, anterolateral margin shorter than posterolateral margins each almost straight, scarcely angled; anterolateral margin with 1 sharp tooth at midpoint; exorbital angle spine-like. Eyestalk 0.5–1.2 times front width. Pereopods 2–5 meri extensor margin each with distal tooth. Gonopod 1 slender, ~6 times as long as wide. Gonopod 2 about as long as gonopod 1, distal part shorter than basal part. Maximum cl. 24 mm. Intertidal–slope (0–700 m). Temperate Northern and Tropical Atlantic (including Mediterranean), Temperate Southern Africa. 4 species (Castro 2007; Guinot and Castro 2007).

Guinoplax Castro & Ng, 2010 Diagnosis. Carapace widest just posterior to exorbital angle, anterolateral margin convex; front projecting well anterior to exorbital angles, eyes exposed laterally; anterolateral margin without tooth; exorbital angle obsolete. Eyestalk 0.8 times front width. Right cheliped dactylus cutting edge with large basal tooth. Gonopod 1 triangular, ~3 times as long as wide. Gonopod 2 almost as long as gonopod 1, distal part as long as basal part. Maximum cl. 5.5 mm. Slope (430–460 m). Central Indo-Pacific. 1 species (Castro and Ng 2010).

Hadroplax Castro, 2007 Diagnosis. Carapace quadrate, parallel-sided over most of length. Anterolateral margin convex, directed anterolaterally; anterolateral margin without tooth; exorbital angle tooth-like. Eyestalk 0.8 times front width. Pereopods 2–4 meri extensor margin each with distal tooth, 5 unarmed. Gonopod 1 thin, evenly tapering, apex acute. Gonopod 2 longer than gonopod 1, distal part longer than basal part. Maximum cl. 7.5 mm. Shelf (27–147 m). Temperate Northern W Pacific, Central Indo-Pacific. 2 species (Castro 2007).

Kingelus Ng, 2022 Diagnosis. Carapace widest at second anterolateral tooth; front projecting well anterior to exorbital angles, eyes exposed laterally; exorbital angle low, not produced; anterolateral margin with triangular, almost right-angled tooth and much smaller tooth posterior to exorbital tooth. Eyestalk about 0.2 times front width. Gonopod 1 stout, distinctly tapering over distal one-fifth, with small flap and base of taper, with distal denticles, truncate. Gonopod 2 longer than gonopod 1, strongly recurved distally. Maximum cl. 4 mm. Shelf. Central Indo-Pacific (Indonesia). 1 species (Ng 2022: description, figures).

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Marine Decapod Crustacea

Menoplax Castro, 2007 Diagnosis. Carapace widest at second anterolateral tooth; front projecting well anterior to exorbital angles, eyes exposed laterally; exorbital angle tooth-like, almost right-angled; anterolateral margin with 1 right-angled tooth and long spine posterior to exorbital tooth. Eyestalk 0.2 times front width. Gonopod 1 stout, tapering over distal one-fifth, with distal denticles, truncate. Gonopod 2 longer than gonopod 1, distal part longer than basal part. Maximum cl. 13 mm. Shelf, slope (65–1300 m). Western and Central Indo-Pacific. 1 species (Castro 2007; Guinot 1989: figures).

Microgoneplax Castro, 2007 Diagnosis. Carapace widest at exorbital tooth; anterolateral margin without tooth; exorbital angle dominating lateral margin. Eyestalk 0.7–1.0 times front width. Male telson subtriangular. Gonopod 1 denticulate over distal half. Gonopod 2 about half as long as gonopod 1, with expanded apex. Maximum cl. 5 mm. Shelf, slope (100–300 m). Temperate Northern W Pacific, Central Indo-Pacific. 7 species (Castro 2007: key to 5 species)

Neogoneplax Castro, 2007 Diagnosis. Carapace widest at exorbital tooth; anterolateral margin with 1 sharp small tooth close to exorbital tooth; exorbital angle spine-like. Eyestalk 0.8–1.1 times front width. Gonopod 1 slender, ~6 times as long as wide. Gonopod 2 longer than gonopod 1, distal part longer than basal part. Maximum cl. 10 mm. Shelf, slope (50–512 m). Temperate Northern W Pacific, Central and Eastern Indo-Pacific. 3 species (Castro 2007: key to species).

Neommatocarcinus Takeda & Miyake, 1969 policeman crab Diagnosis. Carapace widest at exorbital tooth; anterolateral margin without tooth; exorbital angle dominating lateral margin. Antennules not separated by septum. Eyestalk 2 times front width. Male pleonites 3–5 fused. Pereopods 2–5 meri extensor margin each with distal tooth. Gonopod 1 sinuous, tapering. Gonopod 2 longer than gonopod 1, distal part shorter than basal part. Maximum cl. 17 mm. Shelf, slope (18–594 m). Temperate Australasia (New Zealand). 1 species (McLay 1988; Takeda and Miyake 1969).

Notonyx A. Milne-Edwards, 1873 Diagnosis. Carapace quadrate, parallel-sided over most of length. Anterolateral margin convex, directed anterolaterally; anterolateral margin without tooth; exorbital angle obsolete. Eyestalk 0.3– 0.4 times front width; cornea narrower than eyestalk. Gonopod 1 stout. Gonopod 2 longer than gonopod 1, distal part longer than basal part. Maximum cl. 10 mm.

Intertidal, shelf (0–82 m). Western and Central Indo-Pacific. 12 species. Castro (2007) diagnosed the genus. New species have been added since (Clark and Ng 2011; Naruse and Takeda 2010; Ng and Clark 2010; Rahayu 2011; Rahayu and Ng 2010).

Ommatocarcinus White, 1851 Diagnosis. Carapace widest at exorbital tooth; anterolateral margin without tooth; exorbital angle dominating lateral margin. Eyestalk 3 times front width (or more). Pereopods 2–5 meri extensor margin each with distal tooth. Gonopod 1 sinuous, tapering. Gonopod 2 longer than gonopod 1, distal part shorter than basal part. Maximum cl. 30 mm. Subtidal–slope (5–333 m). Temperate and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 4 species (Castro 2007: key to species).

Paragoneplax Castro, 2007 Diagnosis. Carapace widest at exorbital tooth; anterolateral margin without tooth; exorbital angle dominating lateral margin. Eyestalk 0.8–0.9 times front width. Pereopods 2–5 meri extensor margin each with distal tooth. Gonopod 1 tapering, distally twisted. Gonopod 2 half as long as gonopod 1. Maximum cl. 13 mm. Shelf (53–280 m). Central Indo-Pacific. 2 species (Castro 2007; Jiang and Liu 2011).

Pedroplax Ng & Komai, 2011 Diagnosis. Carapace widest at anterolateral tooth; anterolateral margin with 1 tooth close to and similar to exorbital tooth; exorbital angle obsolete. Eyestalk 0.5 times front width. Gonopod 1 with 2 subdistal spines. Gonopod 2 longer than gonopod 1, distal part with bifurcate apex. Maximum cl. 5.6 mm. Shelf (25–59 m). Central Indo-Pacific (Japan, Coral Sea). 2 species (Davie 2012; Ng and Komai 2011).

Psopheticus Wood-Mason, 1892 squeaker crab Diagnosis. Carapace widest at anterolateral tooth; anterolateral margin with 1 sharp tooth posterior to exorbital tooth; exorbital angle spine-like. Eyestalk 0.5 times front width. Pereopods 2 merus extensor margin with 1 distal tooth, pereopods 3–5 with row of teeth. Gonopod 1 triangular, ~4 times as long as wide. Gonopod 2 about as long as gonopod 1, distal part shorter than basal part. Maximum cl. 25 mm (Pl. 44h). Shelf, slope (190–900 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 4 species (Castro 2007: key to species; Guinot 1990: species descriptions). Alcock (1902) called P. stridulans Wood-Mason, 1892, the ‘squeaker crab’ because of its ‘dismal sound, like the squeaking of a pencil on a slate … made by rubbing the spine on the ‘arm’ of this crab against a roughened knob on the adjacent orbit’.

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Pycnoplax Castro, 2007 Diagnosis. Carapace anterolateral margin with 2 sharp teeth posterior to exorbital tooth, widest at second anterolateral tooth; front projecting well anterior to exorbital angles, exorbital angle obsolete; eyes exposed laterally. Eyestalk 0.2 times front width. Cheliped merus without stridulatory ridge on inner distal margin. Gonopod 1 stout, with wide basal half, apex acute. Gonopod 2 longer than gonopod 1, distal part longer than basal part. Vulvae less than one-quarter width of corresponding sternite 6, with cover. Maximum cl. 25 mm (Pl. 44i). Shelf, slope (46–1045 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia. 5 species (Castro 2007: key to species).

Singhaplax Serène & Soh, 1976 Diagnosis. Carapace widest at exorbital tooth; anterolateral margin without tooth; exorbital angle dominating lateral margin. Eyestalk 0.8–1.2 times front width. Male telson subtriangular or semicircular. Gonopod 1 sinuous, tapering. Gonopod 2 longer than or as long as gonopod 1, distal part longer than basal part. Maximum cl. 4.5 mm. Shelf, slope (80–440 m). Central Indo-Pacific. 10 species (Castro 2007; Naruse and Castro 2010; Takeda and Komatsu 2010).

Thyraplax Castro, 2007 Diagnosis. Carapace widest behind second anterolateral tooth; front scarcely projecting anterior to exorbital angles, eyes hidden laterally; anterolateral margin with first tooth obsolete or absent, second tooth small, sharp; exorbital angle obsolete. Eyestalk 0.2–0.3 times front width. Gonopod 1 sinuous, with wide basal third, apex oblique or truncate. Gonopod 2 longer than gonopod 1, distal part shorter than basal part. Maximum cl. 16 mm. Shelf, slope (250–800 m). Central and Eastern Indo-Pacific. 5 species (Castro 2007: key to species). References Alcock A (1902) A naturalist in Indian seas, or four years with the Royal Indian Marine Survey Ship “Investigator”. John Murray, London. Castro P (2007) A reappraisal of the family Goneplacidae MacLeay, 1838 (Crustacea, Decapoda, Brachyura) and revision of the subfamily Goneplacinae, with the description of 10 new genera and 18 new species. Zootaxa 29, 609–774. Castro P, Ng PKL (2010) A new genus and species of goneplacid crab (Decapoda, Brachyura, Goneplacidae) from the Western Pacific. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 51–60. doi:10.1163/ej.9789004170865.i-366.40 Clark PF, Ng PKL (2011) A new species of Notonyx A. Milne-Edwards, 1873 (Crustacea: Decapoda: Brachyura: Goneplacidae) from the Philippines. Zootaxa 2982, 27–32. Davie PJF (2012) A new species of Pedroplax (Decapoda, Brachyura, Goneplacidae) from the Coral Sea, and a new record of Pedroplax megalops from north-western Australia. In: Komatsu H, Okuno J, Fukuoka K (eds) Studies on Eumalacostraca: a homage to Masatsune Takeda. Crustaceana Monographs 17, 105–115. doi:10.1163/9789004202894_009

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Guinot D (1969) Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures. VII. Les Goneplacidae (suite et fin). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 41, 688–724, pls 1–5. Guinot D (1989) Les genre Carcinoplax H. Milne Edwards, 1852 (Crustacea, Brachyura: Goneplacidae). In: Forest J (Ed.), Résultats des Campagnes MUSORSTOM, vol. 5. Mémoires du Muséum National d’Histoire Naturelle, Paris 144, 265–345. Guinot D (1990) Crustacea Decapoda: le genre Psopheticus Wood-Mason, 1892 (Goneplacidae). In: Crosnier A (Ed.), Résultats des Campagnes Musorstom, vol. 6. Mémoires du Muséum National d’Histoire Naturelle, Paris 145, 331–367. Guinot D, Castro P (2007) A new species of Goneplax Leach, 1814 (Crustacea, Decapoda, Brachyura, Goneplacidae) from the south Atlantic and the western limits of the Indo-West Pacific region, long confused with G. rhomboides (Linnaeus, 1758). Zootaxa 1577, 17–31. doi:10.11646/​ zootaxa.1577.1.3 Jiang W, Liu R (2011) On a new species of Paragoneplax Castro (Crustacea: Decapoda: Goneplacidae), P. chenae sp. nov. from the South China Sea. Chinese Journal of Oceanology and Limnology 29, 213–216. doi:10.1007/s00343-011-9072-3 Komatsu H, Takeda M (2003) Two new species of the genus Goneplax (Decapoda, Brachyura, Goneplacidae) from East Asia. Crustaceana 76, 1243–1256. doi:10.1163/156854003773123474 McLay CL (1988) Brachyura and crab-like Anomura of New Zealand [Crabs of New Zealand]. Leigh Laboratory Bulletin 22, 1–463. Naruse T, Castro P (2010) A new species of Microgoneplax Castro, 2007 (Decapoda, Brachyura, Goneplacidae) from Ambon, Indonesia. Crustaceana Monographs 11, 215–220. Naruse T, Takeda M (2010) Notonyx angulatus, a new species of goneplacid crab (Crustacea, Decapoda, Brachyura) from Amami-Oshima, the Ryukyu Islands. Bulletin of the National Museum of Nature and Science. Series A, Zoology 36, 71–74. Ng PKL (2022) The goneplacid crabs (Decapoda, Brachyura) of Raoul Serène and Kasim Moosa, including a new species and genus from Indonesia. Crustaceana 95, 229–240. doi:10.1163/15685403-bja10182 Ng PKL, Castro P (2020) A revision of Carcinoplax abyssicola (Miers, 1885) and seven related species of Carcinoplax H. Milne Edwards, 1852, with the description of two new species and an updated key to the genus (Crustacea, Decapoda, Brachyura, Goneplacidae). Zoosystema 42, 239–284. doi:10.5252/zoosystema2020v42a17 Ng PKL, Clark PF (2010) Two new species of Notonyx A. Milne-Edwards, 1873 (Crustacea: Decapoda: Brachyura: Goneplacidae) from the Philippines and Fiji. Zootaxa 2509, 30–38. doi:10.11646/zootaxa.2509.1.2 Ng PKL, Clark PCF, Ahyong ST (2022) The identity of Homoioplax haswelli (Miers, 1884) (Crustacea: Decapoda: Brachyura: Goneplacidae). Zoological Studies 61, 6. doi:10.6620/ZS.2022.61-06 Ng PKL, Komai T (2011) A new genus for Psopheticus megalops Takeda, 1989 (Crustacea: Brachyura: Goneplacidae). Zootaxa 2934, 61–67. doi:10.11646/​zootaxa.2934.1.6 Poupin J, Corbari L (2016) A preliminary assessment of the deep-sea Decapoda collected during the KARUBENTHOS 2015 Expedition to Guadeloupe Island. Zootaxa 4190, 1–107. doi:10.11646/ zootaxa.4190.1.1 Rahayu DL (2011) A new species of Notonyx A. Milne-Edwards 1873 (Crustacea: Decapoda: Brachyura: Goneplacidae) from Maluku, Indonesia. Zootaxa 2982, 33–39. Rahayu DL, Ng PKL (2010) Notonyx guinotae, a new species of goneplacid crab (Brachyura, Goneplacidae) from Lombok Island, Indonesia. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 269–278.

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Rathbun MJ (1914) A new genus and some new species of crabs of the family Goneplacidae. Proceedings of the United States National Museum 48, 137–154. doi:10.5479/si.00963801.2067.137 Shen CJ (1932) The brachyuran Crustacea of North China. Zoologia Sinica. Series A. Invertebrates of China 9, 1–320, pls 1–10. Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Takeda M, Komatsu H (2010) Two new species of the family Goneplacidae (Decapoda, Brachyura) from the West Pacific. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 319–328. Takeda M, Miyake S (1969) A small collection of crabs from New Zealand. Ohmu 2, 157–193, pls 1–3. Tavares MS (1996) Sur la validité de Bathyplax typhlus oculiferus Miers, 1886 (Decapoda, Brachyura). Crustaceana 69, 413–423. doi:10.1163/​ 156854096X01005

Litocheiridae Kinahan, 1856 Figure 14.49 Litocheirids are small crabs from shallow mudflats of southern Australia (Poore 2004). The family was diagnosed by Števčić (2005). Litocheirids closely resemble many goneplacids but are distinguished by features of the male and female reproductive structures. The gonopod 1 is stout, twisted and grapsoid-like (Fig. 14.49f; Guinot et al. 2013; Ng et al. 2008) and females have rounded lobes on inner margins of the vulva. Diagnosis. Carapace smooth, glabrous; anterolateral tooth small. Male thoracic sternum with ‘press-button’ on posterior one-third of sternite 5. Male thoracic sternite 8 without supplementary plate, male genital papilla exposed, lying in open groove between thoracic sternites 7 and 8 (Fig.  14.49c). Male pleon broadly triangular; pleonites 1–6 freely articulating. Eyestalk large, with conspicuous cornea. Maxilliped 3 merus anterolateral margin obtusely angular. Pereopods 2–5 extensor and flexor margins unarmed,

sparsely setose. Female vulva with 2 or more rounded lobes on margin (Fig.  14.49d). Gonopod 2 two-thirds length of gonopod 1 (Fig. 14.49e, f). Georgeoplax Türkay, 1983 Diagnosis. Carapace about as wide as long; front single-edged; anterolateral margin curved, defined by slight knob (Fig. 14.49g). Pereopods 2–5 without setae. Maximum cl. 12 mm. Subtidal (10–20 m). Temperate SW Australia. 1 species (Poore 2004; Türkay 1983).

Litocheira Kinahan, 1856 Diagnosis. Carapace wider than long; front double-edged; anterolateral margin straight, defined by sharp spine (Fig.  14.49a, b). Pereopods 2–5 setose. Maximum cl. 11 mm. Intertidal, subtidal (0–15 m). Temperate Australia. 1 species (Poore 2004). Platyozius perpusillus Nobili, 1906, from French Polynesia is doubtfully included in this genus (Ng et al. 2008). References Guinot D, Tavares M, Castro P (2013) Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa. Zootaxa 3665, 1–414. doi:10.11646/zootaxa.3665.1.1 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Türkay M (1983) Georgeoplax, new genus for Litocheira glabra Baker, 1906 (Crustacea: Decapoda: Brachyura). Australian Museum Memoir 18, 101–105. doi:10.3853/j.0067-1967.18.1984.375

Fig. 14.49.  Litocheiridae. Litocheira bispinosa Kinahan, 1856: a, habitus; b, front; c, male right thoracic sternites 7, 8, coxa 5 (numbered), genital papilla (gp); d, female thoracic sternum (sternites numbered; v, vulva); e, f, gonopods 1, 2. g, Georgeoplax glabra (Baker, 1906).

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Mathildellidae Karasawa & Kato, 2003 Figure 14.50, Plate 44j Mathildellids are deepwater crabs from lower shelf and slope depths. Most inhabit the Indo-West Pacific but one genus is represented in the Central and South Atlantic. The genera were redefined by Guinot and Richer de Forges (1981a, b) and the family was erected by Karasawa and Kato (2003). Mathildellids, particularly Mathildella and Neopilumnoplax, could be mistaken for progeryonids but are separable by differences in male pleonal somite fusion. Some juvenile geryonids, such as Chaceon and Geryon, which also occur in deep water, may be confounded with mathildellids, but have long, slender anterolateral carapace spines at body sizes that overlap mathildellids.

531

Diagnosis. Carapace anterolateral margins with 3 or more prominent acute teeth; surface smooth or uneven, granular or at most with few low ridges, glabrous or sparsely setose. Male thoracic sternum with ‘press-button’ on posterior half of sternite 5, near suture 5/6. Male thoracic sternite 8 without supplementary plate, male genital papilla exposed, lying in open groove between thoracic sternites 7 and 8 (Fig. 14.50h). Male pleon broadly triangular; pleonites 3–4 or 3–5 fused but demarcation between fused somites clearly indicated by complete transverse groove (Fig. 14.50i). Eyestalk short, with conspicuous cornea. Maxilliped 3 merus anterolateral margin obtusely angular (Fig. 14.50m, n). Pereopods 2–5 extensor and flexor margins unarmed or spinous, usually sparsely setose. Female vulva with simple margin or single vulvar cover. Gonopod 2 longer than gonopod 1 (Fig. 14.50l), or as long as gonopod 1 (Fig. 14.50k; Intesius].

Fig. 14.50.  Mathildellidae. a, Beuroisia major (Sakai, 1978). Carapace: b, Intesius; c, Mathildella; d, Neopilumnoplax; e, Platypilumnus jamesoni Richer de Forges, 1996. Right epistomial region: f, Mathildella; g, Neopilumnoplax. h, sternum, coxa 5, gonopore (shaded), pleon (dashed), Beuroisia. i, male pleon, Intesius. j, pereopod 4, Platypilumnus. Gonopods 1, 2: k, Intesius; l, Mathildella. Maxilliped 3: m, Intesius; n, Neopilumnoplax.

532

Marine Decapod Crustacea

Key to genera of Mathildellidae 1. – 2. – 3. – 4. –

Carapace anterolateral margin with exorbital tooth followed by and 1 or 2 anterolateral teeth forming a serrate plate, followed by 2 broad well separated teeth, last tooth sharp (Fig. 14.50a)�������������������������������������� Beuroisia Carapace anterolateral margin with exorbital tooth almost indistinguishable from first anterolateral tooth or with numerous spines (Fig. 14.50b–e)��������������������������������������������������������������������������������������������������������������������������������2 Pereopods 2–5 meri–propodi extensor margins spinose, meri flexor margins spinose (Fig. 14.50j)������������Platypilumnus Pereopods 2–5 meri smooth, at most with scarce granulation�������������������������������������������������������������������������������������������������3 Carapace sparsely setose. Cheliped carpus inner margin with 1 prominent spine. Gonopod 1 longer than gonopod 2 (Fig. 14.50k)�������������������������������������������������������������������������������������������������������������������������������������������������� Intesius Carapace glabrous. Cheliped carpus inner margin with 2 prominent spines. Gonopod 1 shorter than gonopod 2 (Fig. 14.50l)���������������������������������������������������������������������������������������������������������������������������������������������������������������4 With 1 endostomial ridge on each side (Fig. 14.50f)����������������������������������������������������������������������������������������������� Mathildella With 2 endostomial ridges on each side, inner ridge, shorter, smooth or granular (Fig. 14.50g)�������������� Neopilumnoplax Beuroisia Guinot & Richer de Forges, 1981

Neopilumnoplax Serène, 1969

Diagnosis. Carapace smooth; front bilobed, smooth; anterolateral margin with exorbital tooth and 1 or 2 anterolateral teeth forming a serrate plate, followed by 2 broad well separated teeth, last tooth sharp; with 1 endostomial ridge on each side, not reaching anterior margin of buccal frame. Cheliped carpus inner margin unarmed or with 1 or 2 acute tubercles. Pereopods 2–5 meri smooth, at most with scarce granulation. Maximum cl. 61 mm. Slope (200–635 m). Temperate Northern W Pacific, Indo-West Pacific. 3 species (Guinot and Richer de Forges 1981a: figures; 1981b: all species compared; Sakai 1983).

Diagnosis. Carapace with low epibranchial and other ridges; front straight, smooth; anterolateral margin with shallow concavity between exorbital tooth and fused first anterolateral tooth, second-fourth teeth sharp; with 2 endostomial ridges on each side, inner shorter. Cheliped carpus inner margin with 2 spines. Pereopods 2–5 meri smooth, at most with scarce granulation. Gonopod 1 shorter than gonopod 2. Maximum cl. 16 mm. Shelf, slope (137–1053 m). Tropical W Atlantic, Western IndoPacific, Temperate Australasia, Temperate South America, Temperate Southern Africa. 7 species (Ahyong and Ng 2016; De Matos-Pita and Ramil 2016: key to species).

Intesius Guinot & Richer de Forges, 1981 Diagnosis. Carapace almost smooth or finely granulate, sparsely setose; front bilobed, denticulate; anterolateral margin denticulate, with small sharp exorbital tooth followed by 3 short pointed teeth; with 1 endostomial ridge on each side, not reaching anterior margin of buccal frame. Cheliped carpus inner margin with 1 spine. Pereopods 2–5 meri smooth, at most with scarce granulation. Gonopod 1 longer than gonopod 2. Maximum cl. 40 mm. Slope (220–535 m). Western and Central Indo-Pacific, Temperate Australasia. 5 species (Ahyong 2008; Padate et al. 2021: all species compared).

Mathildella Guinot & Richer de Forges, 1981 Diagnosis. Carapace smooth; front straight or gently curved, smooth; anterolateral margin with shallow concavity between exorbital tooth and fused first anterolateral tooth, second-fourth teeth sharp; with 1 endostomial ridge on each side, reaching anterior margin of buccal frame. Cheliped carpus inner margin with 2 spines. Pereopods 2–5 meri smooth, at most with scarce granulation. Gonopod 1 shorter than gonopod 2. Maximum cl. 34 mm (Pl. 44j). Shelf, slope (180–787 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia. 6 species (Ahyong and Ng 2016; Guinot and Richer de Forges 1981a: figures; 1981b: all species compared).

Platypilumnus Alcock, 1894 Diagnosis. Carapace smooth; front usually bilobed, denticulate; anterolateral margin with c. 15 irregular unequal sharp spines; with 1 short endostomial ridge on each side. Cheliped carpus inner margin with 2 spines. Pereopods 2–5 meri–propodi extensor margins spinose, meri flexor margins spinose. Maximum cl. 40 mm. Shelf, slope (178–950 m). Western and Central Indo-Pacific. 4 species (Richer de Forges 1996: key to species). References Ahyong ST (2008) Deepwater crabs from seamounts and chemosynthetic habitats off eastern New Zealand (Crustacea: Decapoda: Brachyura). Zootaxa 1708, 1–72. doi:10.11646/zootaxa.1708.1.1 Ahyong ST, Ng PKL (2016) The species of Mathildella Guinot and Richer de Forges, 1981 and Neopilumnoplax Serène in Guinot, 1969 (Decapoda: Brachyura: Mathildellidae). Journal of Crustacean Biology 36, 538–552. doi:10.1163/1937240X-00002446 De Matos-Pita SS, Ramil F (2016) New species of Neopilumnoplax Serène in Guinot, 1969 (Decapoda, Brachyura, Mathildellidae) from Northwest Africa with a key to the genus. Marine Biodiversity 46, 253–260. doi:10.1007/s12526-015-0361-5 Guinot D, Richer de Forges B (1981a) Crabes de profondeur, nouveaux ou rares, de l’Indo-Pacifique (Crustacea, Decapoda, Brachyura) (première partie). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série 24, 1113–1153, pls 1–7. [Dated 1980, published 1981]

14 – Brachyura – crabs

Guinot D, Richer de Forges B (1981b) Crabes de profondeur, nouveaux ou rares, de I’ Indo-Pacifique (Crustacea, Decapoda, Brachyura) (deuxième partie) Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 3, 227–260. Karasawa H, Kato H (2003) The family Goneplacidae (Crustacea: Decapoda: Brachyura): systematics, phylogeny, and fossil records. Paleontological Research 7, 129–151. doi:10.2517/prpsj.7.129 Padate VP, Cubelio SS, Takeda M (2021) Rare deep-water crabs (Crustacea: Decapoda) from Indian waters, with description of one new species. Nauplius 29, e2021034. doi:10.1590/2358-2936e2021034 Richer de Forges B (1996) The genus Platypilumnus Alcock and description of P. jamiesoni n. sp. from New Caledonia (Crustacea, Decapoda, Brachyura). Records of the Australian Museum 48, 1–6. doi:10.3853/​ j.0067-1975.48.1996.278 Sakai T (1983) Description of new genera and species of Japanese crabs together with systematically and biogeographically interesting species. (I). Researches on Crustacea 12, 1–44, pls 1–8.

Progeryonidae Števčić, 2005 Figure 14.51, Plate 44k Progeryonids are a small group of deepwater crabs that resemble mathildellids and goneplacids, such as Carcinoplax.

533

The family was erected by Števčić (2005) and rediagnosed by Castro and Ng (2008). Diagnosis. Carapace anterolateral margins with 1 or more teeth, sometimes prominent acute; surface smooth or uneven, granular or at most with few low ridges, glabrous or sparsely setose. Male thoracic sternum with ‘press-button’ on posterior half of sternite 5, near midlength or more posteriad near suture 5/6. Male thoracic sternite 8 without supplementary plate, male genital papilla exposed, lying in open groove between thoracic sternites 7 and 8 (Fig.  14.51g). Male pleon broadly triangular; pleonites 1–6 freely articulating (Fig. 14.51e, f) or 3–4 fused and suture indicated medially and laterally (Fig. 14.51d). Eyestalk short, with conspicuous cornea. Maxilliped 3 merus anterolateral margin obtusely angular (Fig.  14.51h). Pereopods 2–5 smooth or granular; extensor and flexor margins unarmed, sparsely setose. Female vulva with simple margin or single vulvar cover. Gonopod 2 longer than gonopod 1 (Fig. 14.51i, j).

Key to genera of Progeryonidae 1. – 2. –

Carapace subcircular, strongly convex (Fig. 14.51c)����������������������������������������������������������������������������������������������Rhadinoplax Carapace wider than long, flat (Fig. 14.51a, b)�����������������������������������������������������������������������������������������������������������������������������2 Carapace anterolateral margin with 4 teeth behind exorbital tooth, last 2 spiniform (Fig. 14.51a). Male pleonites 3 and 4 fused (Fig. 14.51d)���������������������������������������������������������������������������������������������������������������������� Paragalene Carapace anterolateral margin convex, with 1 tooth at widest point (Fig. 14.51b). Male pleonites 3 and 4 free (Fig. 14.51e, f)�������������������������������������������������������������������������������������������������������������������������������������������������������Progeryon

Fig. 14.51.  Progeryonidae. Carapace: a, Paragalene danieleae Tavares & de Melo, 2010; b, Progeryon mus Ng & Guinot, 1999; c, Rhadinoplax microphthalmus (Guinot & Richer de Forges, 1981). Male pleonites 3–6, telson: d, Paragalene; e, Progeryon; f, Rhadinoplax. g, male thoracic sternum, Rhadinoplax. Maxilliped 3: h, Progeryon. Gonopods 1, 2: i, j, Rhadinoplax.

534

Marine Decapod Crustacea

Paragalene Kossmann, 1878

Scalopidiidae Števčić, 2005

Diagnosis. Carapace ~1.5 times as wide as long, with conspicuous transverse gastric and branchial ridges; front with 2 acute submedian teeth; anterolateral margin with 4 teeth, last 2 spiniform. Male pleonites 3, 4 fused, with suture visible medially and laterally. Maximum cl. 51 mm. Slope (20–250 m; in caves and on hard bottoms with algal growth and shells). Temperate Northern and Tropical Atlantic, including Mediterranean. 2 species (Castro and Ng 2008: discussion; Tavares and de Melo 2010: second species).

Figure 14.52, Plate 44i The family, erected for a single genus by Števčić (2005), was further justified by Ng and Castro (2013) who remarked on the tubular, partially calcified genital papilla lying in a channel on the sternum. The family is notable for spines along both the flexor and extensor margins of pereopods 2–5. Its species are superficially similar to rhizopine pilumnids. Diagnosis. Carapace widest posteriorly, posterolateral margins divergent or subparallel; anterolateral margin unarmed; surface smooth or uneven, granular or at most with few low ridges; sparsely setose. Male thoracic sternum with ‘press-­ button’ on posterior half of sternite 5, near suture 5/6. Male thoracic sternite 8 without supplementary plate, male genital papilla exposed, lying in open groove between thoracic sternites 7 and 8 (Fig. 14.52e, f). Male pleonite 3 to telson distinctly T-shaped; pleonites 1–6 freely articulating; telson subtriangular, lateral margins straight to convex. Eyestalk short, with small cornea. Maxilliped 3 merus anterolateral margin obtusely angular. Pereopods 2–5 merus extensor and flexor margins armed with row of small spines (Fig.  14.52a, b). Female vulva with simple margin or single vulvar cover. Gonopod 2 about one-third length of gonopod 1 (Fig. 14.52g, h).

Progeryon Bouvier, 1922 Diagnosis. Carapace trapezoidal, smooth; front lamellar, with slight median notch; anterolateral margin convex, with 1 tooth at widest point. Male pleonites 3, 4 free. Maximum cl. 47 mm. Slope–bathyal (350–2165 m). Temperate Northern Atlantic, Western Indo-Pacific, Central Indo-Pacific, Eastern Indo-Pacific. 5 species (Bouvier 1922; Guinot and Richer de Forges 1981; Ng and Guinot 1999: key to 4 Indo-West Pacific species).

Rhadinoplax Castro & Ng, 2008 Diagnosis. Carapace subcircular, strongly convex, smooth; front lamellar, with slight median notch; anterolateral margin curved, without teeth or with 2 obsolete teeth. Male pleonites 3, 4 free. Maximum cl. 53 mm (Pl. 44k). Slope (239–580 m). Central Indo-Pacific. 1 species (Castro and Ng 2008). References Bouvier E-L (1922) Observations complémentaires sur les Crustacés décapodes (Abstraction faite les Carides) provenant des Campagnes de S. A. S. le Prince de Monaco. Résultats des Campagnes Scientifiques Accompliés sur son Yacht par Albert 1er, Prince Souverain de Monaco 62, 1–106. Castro P, Ng PKL (2008) Rhadinoplax, a new genus of Progeryonidae Števčić, 2005, for Carcinoplax microphthalmus Guinot & Richer de Forges, 1981, and a redescription of Paragalene longicrura (Nardo, 1868) (Crustacea: Decapoda: Brachyura: Goneplacoidea). Zootaxa 1777, 53–68. doi:10.11646/zootaxa.1777.1.2 Guinot D, Richer de Forges B (1981) Crabes de profondeur, nouveaux ou rares, de l’Indo-Pacifique (Crustacea, Decapoda, Brachyura) (première partie). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série 24, 1113–1153, pls 1–7. Ng PKL, Guinot D (1999) On a new species of deep-water crab of the genus Progeryon (Decapoda, Brachyura, Geryonidae) from Hawaii. Crustaceana 72, 685–692. doi:10.1163/156854099503726 Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Tavares M, de Melo GAS (2010) First species of Paragalene Kossmann, 1878 (Decapoda, Brachyura, Progeryonidae) from the Western Atlantic. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 335–344. doi:10.1163/ej.9789004170865.i-366.203

Caenopedia Ng & Castro, 2013 Diagnosis. Eyestalk mobile (Fig. 14.52a, c). Male pleonite 3 wider than pleonite 1 (Fig. 14.52e). Maximum cl. 11 mm. Shelf (42–52 m). Western Indo-Pacific (Madagascar). 1 species (Ng and Castro 2013).

Scalopidia Stimpson, 1858 Diagnosis. Eyestalk fused to orbit (Fig.  14.52b, d). Male pleonite 3 about as wide as pleonite 1 (Fig. 14.52f). Maximum cl. 19 mm (Pl. 44i). Subtidal (4–20 m). Western and Central Indo-Pacific. 3 species (Ng and Castro 2013). References Ng PKL, Castro P (2013) On the genus Scalopidia Stimpson, 1858 (Crustacea: Brachyura: Goneplacoidea: Scalopidiidae), with the description of one new genus and three new species. Zootaxa 3731, 58–76. doi:10.11646/​ zootaxa.3731.1.2 Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159.

Sotoplacidae Castro, Guinot & Ng, 2010 Figure 14.53 The family is confined to Sotoplax robertsi Guinot, 1984 from the tropical Americas (Castro et al. 2010). The single

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535

Fig. 14.52.  Scalopidiidae. a, Caenopedia ocularia Ng & Castro, 2013; b, Scalopidia spinosipes Stimpson 1858. Left orbit c, Caenopedia; d, Scalopidia. Male right thoracic somites 7, 8, pleonites 1–5, genital papilla (shaded): e, Caenopedia; f, Scalopidia. Gonopods 1, 2: g, h, Scalopidia.

species can be distinguished from all other goneplacoids by the long male telson with concave margins that taper to a very narrow tip. Diagnosis. Carapace trapezoid, widest across anterior; 1 anterolateral tooth behind exorbital tooth; surface

glabrous (Fig. 14.53a). Male thoracic sternum with ‘pressbutton’ on anterior half of sternite 5, near suture 4/5. Male thoracic sternite 8 without supplementary plate, male genital papilla exposed, lying in open groove between thoracic sternites 7 and 8 (Fig. 14.53b, d). Male pleon triangular; pleonites 1–6 freely articulating; telson length about twice width, lateral margins distinctly concave (Fig.  14.53c). Eyestalk large, with conspicuous cornea. Maxilliped 3 merus anterolateral margin obtusely angular. Pereopods 2–5 extensor and flexor margins unarmed, sparsely setose. Female vulva with simple margin or single vulvar cover. Gonopod 2 about quarter length of gonopod 1. Sotoplax Guinot, 1984 Maximum cl. 6.7 mm. Shelf (33–54 m). Tropical Atlantic, Tropical Eastern Pacific (Mexico, Brazil). 1 species (Castro et al. 2010; Guinot 1984). References

Fig. 14.53.  Sotoplacidae. Sotoplax robertsi Guinot, 1984: a, Carapace; b, male sternum; c, pleon; d, lateral part of thoracic sternites 7, 8, pleonites 1–3 (dashed), genital opening (shaded); e, f, gonopods 1, 2.

Castro P, Guinot D, Ng PKL (2010) A new family for Sotoplax robertsi Guinot, 1984, with a diagnosis and key to the Goneplacoidea Macleay, 1838 (Crustacea: Decapoda: Brachyura). Zootaxa 2356, 36–56. doi:10.11646/zootaxa.2356.1.2 Guinot D (1984) Découverte d’un nouveau genre de crabe dans le Golfe du Mexique, Sotoplax robertsi gen. nov., sp. nov. (Crustacea Decapoda Brachyura). Anales del Instituto de Ciencias Marinas y Limnología. Universidad Nacional Autónoma de México 11, 91–98.

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References Ng PKL, Manuel-Santos MR (2007) Establishment of the Vultocinidae, a new family for an unusual new genus and new species of Indo-West Pacific crab (Crustacea: Dccapoda: Brachyura: Goneplacoidea), with comments on the taxonomy of the Goneplacidae. Zootaxa 1558, 39–68. doi:10.11646/zootaxa.1558.1.3 Ng PKL, Richer de Forges B (2009) Vultocinus anfractus Ng & ManuelSantos, 2007 (Decapoda, Brachyura, Vultocinidae): a new record for New Caledonia, with notes on female characters. Crustaceana 82, 627–634. doi:10.1163/156854009X407731

Hexapodoidea Miers, 1886

Fig. 14.54.  Vultocinidae. a, Vultocinus anfractus Ng & ManuelSantos, 2007; b, telson; c, d, gonopods 1, 2.

Vultocinidae Ng & Manuel-Santos, 2007 Figure 14.54, Plate 44m The family was erected for a single species, Vultocinus anfractus Ng & Manuel-Santos, 2007, differing from all other goneplacoid families in its coarsely setose and rugose carapace (Ng and Manuel-Santos 2007; Ng and Richer de Forges 2009). The convoluted pattern of carapace and pereopod setation superficially resembles that of the pilumnoid genus Vellumnus, which of course otherwise differs by superfamily distinctions. Diagnosis. Carapace and pereopods covered with thick, coarsely setose ridges and convoluted rugosites (Fig. 14.54a). Male thoracic sternum with ‘press-button’ on posterior half of sternite 5, near suture 5/6. Male thoracic sternite 8 without supplementary plate, male genital papilla exposed, lying in open groove between thoracic sternites 7 and 8. Male pleon narrow; pleonites 3–4 with concave lateral margins, fused but demarcation between fused somites clearly indicated by complete transverse groove (Fig. 14.54b). Eyestalk short, with conspicuous cornea. Maxilliped 3 merus anterolateral margin obtusely angular. Pereopods 2–5 extensor and flexor margins unarmed, sparsely setose. Female vulva semicircular, with raised rim, without vulvar cover. Gonopod 2 as long as gonopod 1 (Fig. 14.54c, d). Vultocinus Ng & Manuel-Santos, 2007 Maximum cl. 30 mm (Pl. 44m). Shelf, 32–200 m. Central IndoPacific. 1 species.

Hexapodoids, comprising one family, are instantly recognisable as the only brachyuran crabs with three pairs of walking legs (pereopods 2–4). Although reduction of pereopod 5 (or 4 and 5) has occurred on other brachyuran families, in none is pereopod 5 completely lost in adults. The wide subcylindrical habitus of hexapodids facilitates their occupation of narrow mud burrows. Three works are essential for the identification of hexapodids. Manning and Holthuis (1981) reviewed species from West Africa and included a key to 11 genera. Some of their misinterpretations were corrected by Guinot (2006) following rediscovery of type material ­ of poorly known species. Rahayu and Ng (2014) provided a key to all genera. Few species have been described since. Diagnosis. Carapace wider than long, regions poorly defined. Antennule folding transversely. Buccal cavity approximately quadrate. Maxilliped 3 merus triangular, tapering distally, rarely subquadrate–hexagonal. Pereopods 2–4 only visible; pereopod 5 absent in adults. Male pleon usually with some pleonites fused. Gonopod 1 stout to slender.

Hexapodidae Miers, 1886 Figures 14.55, 14.56, Plate 44n, o Implicit generic attributes. Carapace regions indistinct; front without basal constriction. Male thoracic sternite 4 smooth; thoracic sternal groove absent or without transverse extension. Telson lateral margins convex. Eye movable, small, cornea scarcely or not wider than peduncle. Maxilliped 3 broad, ischium as wide or longer than merus length, completely covering buccal cavity; propodus slender; exopod with flagellum. Pereopods 2–4 short, merus of longest pair shorter or subequal to length to carapace; dactyli not enlarged or flattened dorsoventrally. Pereopodal meri without longitudinal groove. Gonopod 1 not twisted into a tight spiral.

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537

Key to genera of Hexapodidae 1. – 2. – 3. – 4. – 5. – 6. – 7. –

Eye hammer-shaped, cornea large, much wider than peduncle (Fig. 14.55a). Carapace front with distinct basal constriction, T-shaped (Fig. 14.55a)����������������������������������������������������������������������������������������������������������������Hexaplax Eye small, cornea scarcely or not wider than peduncle (Fig. 14.55b–e). Carapace front without basal constriction (Fig. 14.55b–e)�������������������������������������������������������������������������������������������������������������������������������������������������������2 Pereopods 2–4 long, merus of longest pair much longer than carapace (Fig. 14.55b). Male thoracic sternal groove long, narrow, transverse, extending laterally from sternoabdominal cavity to beyond base of maxilliped 3 (Fig. 14.55g). Eyes fixed������������������������������������������������������������������������������������������������������������� Lambdophallus Pereopods 2–4 short, merus of longest pair shorter or subequal to length to carapace (Fig. 14.55b–e). Male thoracic sternal groove absent, short or without transverse extension from sternoabdominal cavity to base of maxilliped 3 (Fig. 14.55f, h). Eyes movable������������������������������������������������������������������������������������������������������������3 Maxilliped 3 propodus strongly dilated distally (Fig. 14.56D)�������������������������������������������������������������������������������������������������4 Maxilliped 3 propodus slender (Fig. 14.56A–C)��������������������������������������������������������������������������������������������������������������������������6 Gonopod 1 short, twisted into a tight spiral (Fig. 14.56v)������������������������������������������������������������������������������������������Spiroplax Gonopod 1 not twisted into tight spiral���������������������������������������������������������������������������������������������������������������������������������������5 Male pleonite 6 long; telson lateral margins concave (Fig. 14.56l). Gonopod 1 slender, sinuous, directed anteriorly (Fig. 14.56w)�������������������������������������������������������������������������������������������������������������������������������������Thaumastoplax Male pleonite 6 short; telson lateral margins convex (Fig. 14.56a). Gonopod 1 curved into figure-8 shape (Fig. 14.56o)������������������������������������������������������������������������������������������������������������������������������������������������������������ Hexalaughlia Male pleonites free, all tapering; pleonite 6 not constricted (Fig. 14.56h)����������������������������������������������������������������������Stevea Male pleonites 3–5 fused; pleonite 6 usually constricted�����������������������������������������������������������������������������������������������������������7 Maxilliped 3 exopod without flagellum. Pereopods 2–4 dactyli enlarged, triangular, flattened dorsoventrally (Fig. 14.55e). Male pleonite 6 with convex lateral margins (Fig. 14.56i)�������������������������Pseudohexapus Maxilliped 3 exopod with flagellum (Fig. 14.56B). Pereopods 2–4 dactyli not enlarged or flattened dorsoventrally. Male pleonite 6 constricted proximally�������������������������������������������������������������������������������������������������������8

Fig. 14.55.  Hexapodidae. a, Hexaplax megalops Doflein, 1904; b, Lambdophallus sexpes Alcock, 1900; c, Latohexapus granosus Huang, Hsueh & Ng, 2002; d, Mariaplax daviei Rahayu & Ng, 2014; e, Pseudohexapus platydactylus Monod, 1956. Thoracic sternites, pleon: f, Hexapus. Thoracic sternite 4: g, Lambdophallus. Anterior thoracic sternum, pleon of male: h, Latohexapus. Pereopod 4: i, Mariaplax .

538

Marine Decapod Crustacea

Male pleonite 6 much longer than of pleonites 3–5, constricted subproximally (Fig. 14.56f)������������������������������ Paeduma Male pleonite 6 as long as or shorter than pleonites 3–5�����������������������������������������������������������������������������������������������������������9 Male pleonite 6 wider than long (Fig. 14.56j, o). Gonopod 1 bent sharply laterally at distal third (Fig. 14.56v)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������10 – Male pleonite 6 as wide or longer than wide������������������������������������������������������������������������������������������������������������������������������11 10. Telson trilobed, with distinct lateral angular projection (Fig. 14.56n)��������������������������������������������������������������������� Tritoplax – Telson semicircular (Fig. 14.56i)����������������������������������������������������������������������������������������������������������������������������������� Rayapinus 11. Carapace regions distinct (Fig. 14.55c). Thoracic sternite 4 granular, with transverse groove across entire anterior margin (Fig. 14.55h)������������������������������������������������������������������������������������������������������������������������������� Latohexapus – Carapace regions indistinct. Thoracic sternite 4 smooth (Fig. 14.55f)��������������������������������������������������������������������������������� 12 12. Maxilliped 3 slender, ischium width ~0.7 merus length (Fig. 14.56B, C)�����������������������������������������������������������������������������13 – Maxilliped 3 broad, ischium as wide as or longer than merus length (Fig. 14.56A)�����������������������������������������������������������14 13. Male pleonite 6, telson narrower than pleonites 3–5 (Fig. 14.56g). Gonopod 1 slender, sinuous, tapering abruptly to filiform apex, with recurved spines near midlength (Fig. 14.56t)�����������������������������Parahexapus – Male pleonite 6, telson almost as wide as pleonites 3–5 (Fig. 14.56c). Gonopod 1 stout, bent 60° at midpoint, directed anterolaterally, distal part tapered (Fig. 14.56q)��������������������������������������������������������������������������������������� Hexapus 14. Maxilliped 3 ischium expanded mesially (Fig. 14.56A). Gonopod 1 gently curved (Fig. 14.56p)����������������������Hexapinus – Maxilliped 3 ischium not expanded mesially. Gonopod 1 curved or not�����������������������������������������������������������������������������15 15. Pereopodal meri (at least pereopod 3) with longitudinal groove (Fig. 14.55i). Gonopod 1 curved or twisted, sometimes S-shaped (Fig. 14.56s)����������������������������������������������������������������������������������������������������������������������������Mariaplax – Pereopodal meri without longitudinal groove. Gonopod 1 straight, tip covered by dense setae, as wide as proximal part (Fig. 14.56x)�������������������������������������������������������������������������������������������������������������������������������������� Theoxapus 8. – 9.

Fig. 14.56.  Hexapodidae. Male pleon: a, Hexalaughlia; b, Hexapinus; c, Hexapus; d, Lambdophallus (with tip of gonopod 1); e, Latohexapus; f, Paeduma; g, Parahexapus; h, Pseudohexapus; i, Rayapinus; j, Spiroplax; k Stevea; l, Thaumastoplax; m, Theoxapus; n, Tritoplax. Gonopod 1: o, Hexalaughlia; p, Hexapinus; q, Hexapus; r, Latohexapus; s, Mariaplax; t, Parahexapus; u, Rayapinus; v, Spiroplax; w, Thaumastoplax; x, Theoxapus. Maxilliped 3: A, Hexapinus; B, Hexapus; C, Parahexapus; D, Spiroplax.

14 – Brachyura – crabs

539

Hexalaughlia Guinot, 2006

Latohexapus Huang, Hsueh & Ng, 2002

Diagnosis. Male thoracic sternal groove absent. Male pleonites 3–5 fused, narrower distally, pleonite 6, as long as 3–5, wider distally. Maxilliped 3 slender, ischium width ~0.7 merus length, not covering buccal cavity; propodus lower margin expanded. Gonopod 1 curved into figure-8 shape. Maximum cl. 12 mm. Subtidal, shelf (18–65 m). Temperate Northern Pacific, Central Indo-Pacific. 2 species (Guinot 2006).

Diagnosis. Carapace regions distinct, separated by grooves, granulated; male thoracic sternite 4 granular, with transverse groove across entire anterior margin. Male pleonites 3–5 fused, pleonite 6 wider than long, constricted basally. Maxilliped 3 broad, ischium as wide or longer than merus length, completely covering buccal cavity. Gonopod 1 slender, directed anterolaterally, distal quarter not concealed under pleon. Maximum cl. 8 mm. Subtidal. Central Indo-Pacific (Taiwan). 1 species (Huang et al. 2002; Rahayu and Ng 2014).

Hexapinus Manning & Holthuis, 1981 Diagnosis. Male pleonites 3–5 fused, pleonite 6 wider than long, constricted basally. Maxilliped 3 ischium expanded mesially, completely covering buccal cavity. Gonopod 1 gently curved, distal fifth exposed. Maximum cl. 18 mm. Subtidal, shelf (5–27 m). Temperate Northern W Pacific, Central Indo-Pacific. 5 species (Naruse 2020; Rahayu and Ng 2014).

Hexaplax Doflein, 1904 Diagnosis. Carapace front with distinct basal constriction, T-shaped. Male thoracic sternal groove absent. Male pleonites 3–5 fused (sutures sometimes visible), pleonite 6 shorter than 3–5, constricted basally; telson subtriangular. Eye hammershaped, cornea large, much broader than peduncle. Maxilliped 3 not covering buccal cavity. Pereopods 2–4 long, slender, merus of longest pair longer than carapace. Gonopod 1 straight to sinuous, apex concealed under pleon. Maximum cl. 13 mm (Pl. 44n). Slope (280–766 m). Central Indo-Pacific. 3 species (Rahayu and Ng 2014).

Hexapus De Haan, 1835 Diagnosis. Male pleonites 3–5 fused, pleonite 6 wider than long, constricted basally; telson subtriangular. Maxilliped 3 slender, ischium width ~0.7 merus length, not covering buccal cavity. Gonopod 1 stout, bent 60° at midpoint, directed anterolaterally, distal part gradually tapered, exposed. Maximum cl. 5.5 mm. Estuarine, intertidal, subtidal (0–14 m). Western and Central Indo-Pacific. 3 species (Rahayu and Ng 2014).

Lambdophallus Alcock, 1900 Diagnosis. Male thoracic sternal groove long, narrow, transverse, extending laterally from sternoabdominal cavity to beyond base of maxilliped 3. Male pleonites free, 3–5 discoid, pleonite 6 wider distally. Eye fixed, small, cornea scarcely or not wider than peduncle. Maxilliped 3 broad, ischium as wide or longer than merus length, completely covering buccal cavity. Pereopods 2–4 long, slender, merus of longest pair longer than carapace. Gonopod 1 L-shaped. Maximum cl. 4.5 mm. Subtidal, shelf (5–120 m). Western Indo-Pacific (Bay of Bengal). 1 species (Alcock and McArdle 1902: figures).

Mariaplax Rahayu & Ng, 2014 Diagnosis. Male pleonites 3–5 fused, pleonite 6 wider than long, constricted basally. Maxilliped 3 broad, ischium as wide or longer than merus length, completely covering buccal cavity. Pereopod 3 (sometimes others) merus with longitudinal groove. Gonopod 1 curved or twisted, sometimes S-shaped, apex exposed. Maximum cl. 8 mm (Pl. 44o). Intertidal–slope (0–256 m). Temperate Northern W Pacific, Central Indo-Pacific, Temperate Australasia. 18 species (­R ahayu and Ng 2014; Rahayu and Widyastuti 2018: key to species).

Paeduma Rathbun, 1897 Diagnosis. Male pleonites 3–5 fused, pleonite 6 linear, with subproximal constriction. Maxilliped 3 broad, ischium as wide or longer than merus length, completely covering buccal cavity. Maximum cl. 15.3 mm. Habitat not known. Tropical Eastern Pacific. 1 species (Guinot 2006).

Parahexapus Balss, 1922 Diagnosis. Male pleonites 3–5 fused, much narrower distally, pleonite 6 not constricted. Maxilliped 3 slender, ischium width ~0.7 merus length, not covering buccal cavity. Gonopod 1 slender, sinuous, tapering abruptly to filiform apex, with recurved spines near midlength. Maximum cl. 5.6 mm. Subtidal, shelf (5–27 m). Tropical E Atlantic. 1 species (Manning and Holthuis 1981).

Pseudohexapus Monod, 1956 Diagnosis. Male thoracic sternal groove absent. Male pleonites 3–5 fused, pleonite 6 half length of 3–5, constricted proximally. Maxilliped 3 broad, ischium as wide or longer than merus length, completely covering buccal cavity; exopod without flagellum. Pereopods 2–4 dactyli enlarged or flattened dorsoventrally. Gonopod 1 stout, apex curved mesially, with terminal spinules and few subterminal setae. Maximum cl. 9 mm. Subtidal (5–15 m). Tropical E Atlantic. 1 species (Manning and Holthuis 1981).

540

Marine Decapod Crustacea

Rayapinus Rahayu & Ng, 2014 Diagnosis. Male pleonites 3–5 fused, pleonite 6 wider than long, constricted basally. Maxilliped 3 broad, ischium as wide or longer than merus length, completely covering buccal cavity. Gonopod 1 stout, bent subdistally, apex not concealed under pleon. Maximum cl. 4.6 mm. Subtidal (1.5–2 m). Central Indo-Pacific (Ryukyu Is, Japan). 1 species (Rahayu and Ng 2014).

Spiroplax Manning & Holthuis, 1981 Diagnosis. Male pleonites 3–5 fused, as wide as long, pleonite 6 short, wider than long; telson subtriangular. Maxilliped 3 slender, ischium width ~0.7 merus length, not covering buccal cavity; propodus strongly dilated distally. Gonopod 1 short, strongly twisted into a tight spiral. Maximum cl. 5.5 mm. Estuarine, intertidal, shallow subtidal (sometimes commensal with the ghost shrimp, Kraussillichirus kraussi (Stebbing, 1920). Temperate Southern Africa. 1 species (Emmerson 2016; Manning and Holthuis 1981).

Stevea Manning & Holthuis, 1981 Diagnosis. Male thoracic sternal groove absent. Male pleonites free, tapering, pleonite 6 tapering, not constricted. Maxilliped 3 broad, ischium as wide or longer than merus length, completely covering buccal cavity. Maximum cl. 9.4 mm. Subtidal (18–24 m). Tropical Eastern Pacific. 1 species (Guinot 2006; Manning and Holthuis 1981).

Thaumastoplax Miers, 1881 Diagnosis. Male pleonites 3–5 fused, pleonite 6 as long as 3–5, not constricted; telson lateral margins concave. Maxilliped 3 broad, ischium as wide or longer than merus length, completely covering buccal cavity; propodus strongly dilated distally. Gonopod 1 slender, sinuous, directed anteriorly. Maximum cl. 11 mm. Estuarine, subtidal, shelf (3–34 m). Tropical E Atlantic. 1 species (Manning and Holthuis 1981).

Theoxapus Rahayu & Ng, 2014 Diagnosis. Male pleonites 3–5 fused, pleonite 6 shorter than 3–5, ovate. Maxilliped 3 broad, ischium as wide or longer than merus length, completely covering buccal cavity. Gonopod 1 straight, tip covered by dense setae, as wide as proximal part. Maximum cl. 5 mm. Subtidal (7–10 m). Tropical E Atlantic. 1 species (Rahayu and Ng 2014).

Tritoplax Manning & Holthuis, 1981 Diagnosis. Male pleonites 3–5 fused, as wide as long, pleonite 6 wider than long, constricted proximally; telson trilobed, with

distinct lateral angular projection. Maxilliped 3 broad, ischium as wide or longer than merus length, completely covering buccal cavity. Gonopod 1 slender, sinuous, bent laterally, apex recurved. Maximum cl. 10 mm. Shelf (30–70 m). Central Indo-Pacific, Temperate Southern Africa. 2 species (Manning and Holthuis 1981). References Alcock A, McArdle SB (1902) Illustrations of the zoology of the Royal Indian Marine Surveying Steamer “Investigator”. Crustacea, Part X. Vol. 10. Trustees of the Indian Museum, Calcutta. Emmerson WD (2016) A guide to, and checklist for, the Decapoda of Namibia, South Africa and Mozambique. Vol. 2. Cambridge Scholars Publishing, Cambridge. Guinot D (2006) Rediscovery of the holotype of Paeduma cylindraceum (Bell, 1859) and description of a new genus of Hexapodidae (Decapoda, Brachyura). Zoosystema 28, 553–571. Huang J-F, Hsueh P-W, Ng PKL (2002) Crabs of the family Hexapodidae (Decapoda: Brachyura) from Taiwan, with description of a new genus and new species. Journal of Crustacean Biology 22, 651–660. doi:10.1163/​ 20021975-99990277 Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Naruse T (2020) A new species of Hexapinus Manning & Holthuis, 1981 (Decapoda: Brachyura: Hexapodidae) from Hatoma Island, Ryukyu Islands, Japan. Crustacean Research 49, 49–55. doi:10.18353/ crustacea.49.0_49 Rahayu DL, Ng PKL (2014) New genera and new species of Hexapodidae (Crustacea, Brachyura) from the Indo-West Pacific and east Atlantic. Raffles Bulletin of Zoology 62, 396–483. Rahayu DL, Widyastuti E (2018) Additions to the Indonesian crab fauna of the genus Mariaplax Rahayu & Ng, 2014, and notes on Hexapus timika Rahayu & Ng, 2014 (Crustacea, Decapoda, Brachyura, Hexapodidae). Zootaxa 4379, 231–246. doi:10.11646/zootaxa.4379.2.5

Hymenosomatoidea MacLeay, 1838 false spider crabs, flat-backed crabs, crown crabs Hymenosomatoid crabs are small, cryptic, delicate crabs with long thing legs, more ‘spider’-like than majoid crabs some of which are commonly called ‘spider crabs’. The name ‘false spider crabs’ is used to distinguish them. The alternative names, ‘flat-backed crabs’ or ‘crown crab’, recognise their unique feature, a ‘hymenosomian rim’, a ridge defining the edge of the dorsal carapace. Hymenosomatoids differ from all other crabs in having the last pleonite fused to the telson. Most of the 136 hymenosomatoid species are marine, from shallow-water environments, but ten species in eight genera are exclusively freshwater, some of those in caves (Chuang and Ng 1994). Three other genera include estuarine species. The largest hymenosomatoid has a carapace of ~26 mm but the family includes some of the smallest crab species. Elamenopsis minima Lucas & Davie, 1982, is ovigerous at a carapace length of 1.7 mm (Lucas and Davie 1982) and

14 – Brachyura – crabs

others are not much bigger. Micas lucasi (Richer de Forges, 1993) is ovigerous at 2.2 mm (Richer de Forges 1993). The systematic position of hymenosomatids has long been controversial. They share many features with heterotreme brachyurans, notably Majoidea and Dorippioidea, and molecular data supports this conclusion. They differ from all heterotremes, however, in having the male gonopore on the thoracic sternum as in thoracotreme crabs, making these two sections difficult to diagnose unambiguously. The thoracotrematous condition of hymenosomatids, however, appears to have been derived differently from that of true thoracotremes, possibly in connection with the peculiar overall endophragmal organisation (Ahyong et al. 2007; Guinot 2011b; Guinot and Richer de Forges 1997; Guinot et al. 2013). The taxonomy of the single family was established by Melrose (1975) concentrating on the fauna of New Zealand and reviewed again by Lucas (1980) with special attention to the fauna of Australia. More than half of the genera now accepted were established since then, four by Poore et  al. (2016). Two subfamilies were recognised by Guinot (2011a) but this is an interim subdivision of the family. Odiomariinae was established for three genera possessing small uropods but uropods are also seen in two species of Hymenosomatinae, Hymenosoma hodgkini Lucas, 1980 and Elamenopsis guinotae Poore, 2010. The family includes a group of genera, exemplified by Neorhynchoplax, in which the female broods its embryos in a posteroventral thoracic concavity closed by the female pleon without

541

attachments to the pleopods (Hsueh 2018; Lucas 1980; Ng and Chuang 1996). The rostrum has been described in some species as ‘trifid’ or ‘trilobed’. Poore et  al. (2016) interpreted this situation as having a simple rostrum accompanied by a pair of projections at the anterior margins of the supraorbital eave, referred to here as a ‘pseudorostral’ teeth or lobes. The carapace width and length are those of the hymenosomian rim, excluding the rostrum. Diagnosis. Carapace slightly wider than long to about as long as wide (excluding rostrum), pyriform, triangular, to circular, surface flat, soft, pliable, poorly calcified. Antenna folding longitudinally alongside antennules. Buccal cavity approximately quadrate. Maxilliped 3 ischium subequal to or larger than merus. Pereopods 2–5 long, slender, pereopod 5 about twice carapace length or longer. Male and female pleon with telson fused to pleonite 6, others variously fused; female pleon often domed, forming brood chamber. Uropods absent, or rarely present. Gonopod 1 much longer than gonopod 2.

Hymenosomatidae MacLeay, 1838 Figures 14.57–14.60, Plate 45a–f Implicit generic attributes. Carapace with lateral body wall extended beyond hymenosomian rim. Epistome present anterior to buccal cavern. Pleonite 1 posterior margin smooth. Eyes free, pigmented. Male uropod absent. Female incubating embryos attached to pleopods.

Key to subfamilies and genera of Hymenosomatidae 1. – 2. – 3. – 4. – 5. – 6. – 7. –

Male uropods intercalated laterally at base of pleotelson (Fig. 14.59a, g)����������������������������������������������� Odiomarinae … 2 Male uropods absent (Fig. 14.59b–f, h) (rare exceptions)���������������������������������������������������������������� Hymenosomatinae … 4 Carapace almost circular. Rostrum anterior margin concave in dorsal view (Fig. 14.58i)���������������������������������� Richerius* Carapace wider than long. Rostrum ending in triangular or rounded apex (Fig. 14.57d, f)������������������������������������������������3 Gonopod 1 tapering, with 2 distal processes (Fig. 14.60p). Maxilliped 3 merus broadly ovoid (Fig. 14.59q)��������������������������������������������������������������������������������������������������������������������������������������������������������������� Odiomaris* Gonopod 1 tapering abruptly distally, with 1 or more short distal lobes (Fig. 14.60m). Maxilliped 3 merus distolaterally square (Fig. 14.59l)���������������������������������������������������������������������������������������������������������������������������� Amarinus* Epistome absent (Fig. 14.58r)������������������������������������������������������������������������������������������������������������������������������������Hymenosoma Epistome present (Fig. 14.58o–p, s–v)�������������������������������������������������������������������������������������������������������������������������������������������5 Maxillipeds 3 separated by wide gape (Fig. 14.58p)��������������������������������������������������������������������������������������������������������������������6 Maxillipeds 3 almost entirely covering buccal cavern (Fig. 14.58s, t)������������������������������������������������������������������������������������17 Maxilliped 3 narrow, with palp overlapping ischium, reaching close to posterior margin of buccal cavern (Fig. 14.59n, o)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������7 Maxilliped 3 with short palp barely overlapping merus (Figs 14.58p, 14.59p)�����������������������������������������������������������������������9 Male pleotelson semicircular (Fig. 14.59b). Eyestalks fused to orbit, without pigment (Fig. 14.57f, g)���������������������������������������������������������������������������������������������������������������������������������������������������������Cancrocaeca* Male pleotelson trilobed (Fig. 14.59d, e). Eyestalks free, pigmented (Fig. 14.57q)�����������������������������������������������������������������8

542

Marine Decapod Crustacea

Fig. 14.57.  Hymenosomatidae. a, Halicarcinus ovatus Stimpson, 1858; b, Micas filholi (De Man, 1888). Carapace: c, Aletheiana; d, Amarinus; e, Apechocinus; f, g, Cancrocaeca (with front); h, Crustaenia; i, Culexisoma; j, Elamena castanea Naruse, Mendoza & Ng, 2008; k, Elamena globoasa Chuang & Ng, 1991; l, Elamenopsis; m, n, Guaplax (with front); o, Halicarcinides; p, Halimena; q, Hymenicoides; r, Hymenosoma; s, Insulaplax.

8. – 9. – 10. – 11. –

Male pleotelson with lateral lobes auriculate (Fig. 14.59d). Gonopod 1 about twice as wide as long, with distal outer angle strongly swollen (Fig. 14.60n)��������������������������������������������������������������������������������������������� Hymenicoides Male pleotelson with lateral lobes smaller than medial lobe (Fig. 14.59e). Gonopod 1 ~3 times as wide as long, with distal outer angle not swollen (Fig. 14.60o)�������������������������������������������������������������������������������������� Limnopilos* Rostrum elongate, more than 0.7 times carapace length in male, 0.3 times in female; with longitudinal supraorbital eave at base (Figs 14.57i, 14.58p). Female without internal brood cavity, embryos attached to pleopods����������������������������������������������������������������������������������������������������������������������������������������������������������������Culexisoma Rostrum at most 0.2 times carapace length; supraorbital eave absent or evident as small pseudorostral tooth (Fig. 14.57e, h, l). Female with internal brood cavity������������������������������������������������������������������������������������������������10 Marine or estuarine species (2 freshwater species of Neorhynchoplax)���������������������������������������������������������������������������������11 Living in fresh water in streams or caves�����������������������������������������������������������������������������������������������������������������������������������13 Rostrum broad, short. Carapace laterally oval (Fig. 14.57l). Pereopods 2–5 dactyli without subterminal teeth (Fig. 14.60f, l)��������������������������������������������������������������������������������������������������������������������������������������������������Elamenopsis Rostrum triangular, with pair of pseudorostral teeth/lobes. Carapace longitudinally oval (Figs 14.57h, 14.58e). Pereopods 2–5 dactyli with at least 1 subterminal tooth (Fig. 14.60j)��������������������������������������������������������������� 12

14 – Brachyura – crabs

543

Fig. 14.58.  Hymenosomatidae. Carapace: a, Lucascinus; b, c, Nasutoplax (with rostrum, eye, lateral view) d, Neohymenicus; e, Neorhynchoplax; f, Odiomaris; g, h, Rhynchoplax (with rostrum, eye, lateral view) i, Richerius; j, Samarplax; k, Stimpsoplax; l, Sulaplax; m, Teramnonotus; n, Trigonoplax. Antennae, eyes, epistome, buccal cavern, maxillipeds 3, sternum: o, Amarinus; p, Culexisoma; q, Elamena; r, Hymenosoma; s, Lucascinus; t, Rhynchoplax; u, Teramnonotus; v, Trigonoplax.

12. Pleonite 1 with pair of posterior lobes. Male pleonites 2–pleotelson fused, with remnants of suture between pleonite 5 and telson (Fig. 14.59c). Carapace with lateral tooth on carapace hymenosomian rim (Fig. 14.57h)���������������������������������������������������������������������������������������������������������������������������������������������������������������� Crustaenia – Pleonite 1 without pair of posterior lobes. Male pleonites 3 and 4 fused, 3–5 or 4 and 5 fused (Fig. 14.59f). Carapace with lateral tooth, if present, on branchiostegite�����������������������������������������������Neorhynchoplax 13. Eyes fused to orbit, unpigmented (Fig. 14.57m, n). Living in freshwater in caves���������������������������������������������������������������14 – Eyes free, pigmented (Fig. 14.57c, s). Living in freshwater streams����������������������������������������������������������������������������������������16 14. Rostrum triangular, with pair of similar supraorbital teeth (Fig. 14.57m, n)����������������������������������������������������������Guaplax* – Rostrum vestigial or absent (Fig. 14.58j, l)���������������������������������������������������������������������������������������������������������������������������������15 15. Carapace rim longer than wide, rounded-octagonal. Rostrum absent (Fig. 14.58j)������������������������������������������ Samarplax* – Carapace rim wider than long. Rostrum vestigial (Fig. 14.58l)�������������������������������������������������������������������������������� Sulaplax* 16. Carapace with postocular and lateral teeth. Rostrum minute (Fig. 14.57c). Pereopods 2–5 dactyli with at least 1 subdistal tooth on flexor margin (Fig. 14.60a)����������������������������������������������������������������������������������������Aletheiana* – Carapace without postocular and lateral teeth. Rostrum absent (Fig. 14.57s). Pereopods 2–5 dactyli without teeth on flexor margin (Fig. 14.60f)������������������������������������������������������������������������������������������������������� Insulaplax*

544

Marine Decapod Crustacea

Fig. 14.59.  Hymenosomatidae. Male pleon: a, Amarinus; b, Cancrocaeca; c, Crustaenia; d, Hymenicoides; e, Limnopilos; f, Neorhynchoplax; g, Odiomaris; h, Rhynchoplax. Major chela: i, Lucascinus; j, Rhynchoplax; k, Stimpsoplax. Maxilliped 3: l, Amarinus; m, Apechocinus; n, Cancrocaeca; o, Limnopilos; p, Neohymenicus; q, Odiomaris; r, Richerius.

17. – 18. – 19. – 20. – 21. – 22. – 23. – 24. – 25. –

Carapace dorsal surface including rostrum surrounded by continuous hymenosomian rim (Figs 14.57j, k, 14.58m, n), or at least carapace dorsal surface continuous with rostrum����������������������������������������������18 Carapace with rostrum demarcated from dorsal surface by hymenosomian rim (Figs 14.57a, b, e, o, p, 14.58a, b, d, k)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������21 Carapace approximately triangular. Female thoracic sternite 8 mostly exposed dorsolaterally by carapace (Fig. 14.58n). Inhalant water openings fused laterally for more than half its length (Fig. 14.58v)������������� Trigonoplax Carapace polygonal (Fig. 14.57j) or ovate-pyriform (Figs 14.57k, 14.58m). Female thoracic sternite 8 covered dorsolaterally, or nearly so, by carapace (female not known for Chartocinus). Inhalant water openings not fused (Fig. 14.58u)���������������������������������������������������������������������������������������������������������������������������������������������19 Eyestalk mobile. Subhepatic region smooth (Fig. 14.58q)������������������������������������������������������������������������������������������������������ 20 Eyestalk fused to carapace. Subhepatic region with 2 broad tubercles (Fig. 14.58u)������������������������������������� Teramnonotus Rostrum with ventral keel (Fig. 14.57j, k). Pereopod 2–5 dactyli usually with 1 or more teeth on flexor margin, occasionally unarmed����������������������������������������������������������������������������������������������������������������������������������� Elamena Rostrum without ventral keel. Pereopod 2–5 dactyli unarmed���������������������������������������������������������������������������� Chartocinus Carapace rim longer than wide. Rostrum broad, triangular, with rounded tip (Fig. 14.57p). Pereopods 2–5 dactyli curved, with 1 subterminal recurved tooth (Fig. 14.60d)������������������������������������������������������������������������ Halimena Carapace rim as wide or wider than long, Rostrum broad or truncate or with paired pseudorostral teeth. Pereopods 2–5 dactyli with 1 or more teeth on flexor margin (Fig. 14.60b, c, g, h, i)��������������������������������������������������� 22 Rostrum absent, eyes not visible in dorsal view (Fig. 14.57o). Pereopods 2–5 dactyli straight, with subdistal triangular tooth (Fig. 14.60b)�������������������������������������������������������������������������������������������������� Halicarcinides Rostrum present, eyes visible in dorsal view. Pereopods 2–5 dactyli usually with row of teeth along flexor margin (Fig. 14.60e), or 2 subdistal teeth (Fig. 14.60h)�������������������������������������������������������������������������������� 23 Pereopods 2–5 dactyli with 1 or 2 subdistal teeth (Fig. 14.60h, i)����������������������������������������������������������������������������������������� 24 Pereopods 2–5 dactyli with row of teeth along flexor margin (Fig. 14.60e)������������������������������������������������������������������������ 26 Rostrum short, triangular, with similar pair of pseudorostral teeth (Fig. 14.57b)������������������������������������������������������� Micas Rostrum broad, triangular, without pseudorostral teeth (Figs 14.57a, e, 14.58a, d, g, k)�������������������������������������������������� 25 Carapace subcircular (Fig. 14.58d). Maxilliped 3 merus longer than wide (Fig. 14.59p)����������������������������� Neohymenicus Carapace pear-shaped (Fig. 14.57e). Maxilliped 3 merus as wide as long (Fig. 14.59m)�����������������������������������Apechocinus

14 – Brachyura – crabs

545

Fig. 14.60.  Hymenosomatidae. Pereopods 2–5, dactylus: a, Aletheiana; b, Halicarcinides; c, Halicarcinus; d, Halimena; e, Hymenosoma; f, Insulaplax; g, Lucascinus; h, Micas; i, Neohymenicus; j, Neorhynchoplax; k, Teramnonotus; l, Trigonoplax. Gonopod 1; m, Amarinus; n, Hymenicoides; o, Limnopilos; p, Odiomaris; q, Richerius.

26. Rostrum and pseudorostral teeth separate and of similar length (‘trilobed’) (Fig. 14.57a) or together forming broad, approximately triangular plate with 3 short blunt lobes, median the longer������������������� Halicarcinus – Rostrum prominently longer than pseudorostral teeth (Fig. 14.58a–c, g, h, k)������������������������������������������������������������������� 27 27. Rostrum long, laterally compressed, with rounded apex, ventrally expanded, erect in lateral view, 0.35 times carapace length in male, 0.2 times in female. Carapace rim almost circular (Fig. 14.58b, c)����������� Nasutoplax – Rostrum not laterally compressed or ventrally expanded. Carapace rim oval or pyriform (Fig. 14.58a, g, h, k)������������������������������������������������������������������������������������������������������������������������������������������������������������������ 28 28. Carapace rim longer than wide (Fig. 14.58a). Male cheliped with elongate propodus, with short finger and dactylus forming ‘nut-cracker’ (Fig. 14.59i)���������������������������������������������������������������������������������������������������������� Lucascinus – Carapace rim as wide or wider than long (Fig. 14.58g, h, k). Male chela with finger and dactylus about as long as palm (Fig. 14.59j, k)���������������������������������������������������������������������������������������������������������������������������������������������������� 29 29. Rostrum accompanied by pair of short pseudorostral teeth (Fig. 14.58k). Male cheliped fingers with dense setation in gape (Fig. 14.59k)��������������������������������������������������������������������������������������������������������������������������������� Stimpsoplax – Rostrum without pseudorostral teeth (Fig. 14.58g, h). Male cheliped fingers without setation in gape (Fig. 14.59j)��������������������������������������������������������������������������������������������������������������������������������������������������������������Rhynchoplax * genera inhabiting exclusively fresh water and/or estuaries

Subfamily Hymenosomatinae MacLeay, 1838

Apechocinus Ng & Chuang, 1996

Aletheiana Ng & Lukhaup, 2015

Diagnosis. Carapace rim as long as wide (rostrum excluded), narrower anteriorly. Rostrum triangular, as wide as long, quarter carapace length; apex without long setae. Pleonites 1–5, pleotelson free in both sexes. Maxillipeds 3 almost entirely covering buccal cavern; maxilliped 3 merus as wide as long. Pereopods 2–5 dactyli almost straight, with subdistal triangular tooth. Gonopod 1 with swollen base, narrow distal section tapering through sharp proximal ventral curvature and twist, with apex acute sharply bent ventrally. Maximum cl. 2.2 mm. Habitat unknown. Indonesia. 1 species (Ng and Chuang 1996).

Diagnosis. Carapace rim as wide or wider than long (rostrum excluded); with 1 postocular and 1 lateral tooth. Rostrum minute, depressed. Pleonites 1–5, pleotelson free in both sexes. Maxillipeds 3 separated by wide gape. Pereopods 2–5 dactyli curved, with 1 subdistal recurved tooth. Gonopod 1 twisted medially. Female with internal brood cavity incubating free embryos. Maximum cl. 5 mm. Freshwater (amongst vegetation along river banks). Sulawesi, Indonesia. 1 species (Ng and Lukhaup 2015).

546

Marine Decapod Crustacea

Cancrocaeca Ng, 1991

Elamena H. Milne Edwards, 1837

Diagnosis. Carapace rim as long as wide (rostrum excluded), hymenosomian rim almost circular. Rostrum absent, dorsal surface surrounded by continuous hymenosomian rim. Pleonites 1–5, pleotelson free in both sexes; pleotelson semicircular. Maxillipeds 3 separated by wide gape; maxilliped 3 pediform, with long propodus and dactylus, reaching or almost reaching posterior margin of buccal cavern. Pereopods 2–5 dactyli without teeth on flexor margin. Gonopod 1 stout, twisted at base, distal part with 3 processes. Maximum cl. 4.7 mm. Freshwater in caves. Sulawesi, Indonesia. 1 species (Naruse et  al. 2008b: rediagnosis, comparison of 5 cave-dwelling genera; Ng 1991: diagnosis).

Diagnosis. Carapace with lateral body wall not visible beyond hymenosomian rim; rim pear-shaped, or polygonal; subhepatic region smooth. Rostrum triangular or truncate, continuous with carapace surface, with ventral rostral keel. Pleonites 1–5, pleotelson variously fused in both sexes. Maxillipeds 3 almost entirely covering buccal cavern. Pereopods 2–5 dactyli flexor margin unarmed, or with sharp teeth along flexor margin. Gonopod 1 sinuous. Maximum cl. 5 mm (Pl. 45b). Intertidal–shelf (0–116 m). Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 27 species. Elamena has not been reviewed. Two groups of species are recognised, one with a short, truncated rostrum and a polygonal carapace (Fig.  14.57j). The second group has a produced, triangular rostrum and a pyriform carapace (Fig.  14.57k) (Kulabtong et  al. 2013; Naruse et  al. 2008a; Rahayu and Nugroho 2019). Elamena producta Kirk, 1878, is a facultative associate in the mantle cavities of abalones (Haliotis spp.) (McLay 1988).

Chartocinus Ng & Rahayu, 2022 Diagnosis. Carapace longitudinally ovate; hymenosomian rim continuous except at base of rostrum; Rostrum rounded-subtriangular, continuous with carapace surface, without ventral keel. Male sternopleonal cavity shorter than half length of thoracic sternum. Male pleonites 3–5 fused; pleotelson rounded, twice as wide as long. Maxillipeds 3 almost entirely covering buccal cavern. Pereopods 2–5 dactyli without teeth on flexor margin. Gonopod 1 thick, stocky, apically, acute, inner margin strongly sinuous. Maximum cl. 3.1 mm. Estuarine, subtidal. Central Indo-Pacific (Indonesia). 1 species (Ng and Rahayu 2022: description, figures).

Crustaenia Ng & Chuang, 1996 Diagnosis. Carapace rim 1.2 times as long as wide (rostrum excluded); with lateral tooth on hymenosomian rim. Rostrum elliptical, concave, subequal to pseudorostral lobes. Pleonites 3–5 fused in male, 2–5 fused in female; pleonite 1 with pair of posterior lobes. Maxillipeds 3 separated by wide gape. Pereopods 2–5 dactyli curved, with subdistal blunt tooth. Female with internal brood cavity incubating free embryos. Maximum cl. 4 mm. Intertidal, subtidal. Central Indo-Pacific. 1 species (Ng and Chuang 1996).

Culexisoma Poore, Guinot, Komai & Naruse, 2016 Diagnosis. Carapace rim 1.2 times as long as wide (rostrum excluded). Rostrum narrowly triangular, elongate, more than 0.7 times carapace length in male, equilateral, anteriorly upturned, 0.3 times in female; apex with long setae. Pleonites 3–4 fused in male; hemispherical, pleonites 1, 2 fused, 3–5 fused in female. Maxillipeds 3 separated by wide gape. Male cheliped twice as long as carapace length; propodus swollen, fingers gaping slightly, with irregularly toothed cutting edges, dactylus without dominant proximal tooth. Pereopods 2–5 dactyli mostly straight, with row of sharp teeth along proximal three-quarters of flexor margin. Gonopod 1 tapering evenly from base, sinusoidal, with acute apex directed anterodorsally. Maximum cl. 3.5 mm. Subtidal (2–16 m). Central Indo-Pacific. 2 species (Poore et al. 2016).

Elamenopsis A. Milne-Edwards, 1873 Diagnosis. Carapace rim wider than long (rostrum excluded). Rostrum broad, short, deflexed. Pleonites free, or 4–5 fused in male; pleonites 3–5 fused in female. Maxillipeds 3 separated by wide gape. Pereopods 2–5 dactyli without teeth on flexor margin. Gonopod 1 sinuous. Female with internal brood cavity incubating free embryos. Maximum cl. 2.6 mm. Estuarine, marine, intertidal, subtidal (0–15 m). Central IndoPacific, Temperate Australasia. 6 species (Rahayu and Ng 2019: key to species). The ovigerous female of E. minima Lucas & Davie, 1982, at 1.7 mm carapace width, is probably the smallest mature female crab reported (Lucas and Davie 1982). Elamenopsis guinotae Poore, 2010, is anomalous, possessing uropods (Poore 2010).

Guaplax Naruse, Ng & Guinot, 2008 Diagnosis. Carapace rim as wide or wider than long (rostrum excluded). Rostrum triangular with pair of similar pseudorostral teeth. Pleonites 3–5 fused in male. Eyes fused to orbit, unpigmented. Maxillipeds 3 separated by wide gape. Pereopods 2–5 dactyli without teeth on flexor margin. Gonopod 1 straight. Female with internal brood cavity incubating free embryos. Maximum cl. 3.4 mm. Freshwater in caves. Kalimantan, Indonesia. 1 species (Naruse et al. 2008b).

Halicarcinides Lucas, 1980 Diagnosis. Carapace rim as long as wide (rostrum excluded), narrower anteriorly. Rostrum absent, eyes not visible in dorsal view. Pleonites 3–4 fused in both sexes. Maxillipeds 3 almost entirely covering buccal cavern. Pereopods 2–5 dactyli almost straight, with subdistal triangular tooth. Gonopod 1 with swollen base, narrow distal part strongly twisted, sharply curved. Maximum cl. 6 mm (Pl. 45c). Intertidal. Temperate S Australia (rare). 1 species (Lucas 1980; Poore 2004).

14 – Brachyura – crabs

Halicarcinus White, 1846 Diagnosis. Carapace rim as wide or wider than long (rostrum excluded). Rostrum a median lobe similar in males, females, 0.05–0.2 times carapace length, separate from pair of similar pseudorostral teeth, OR together forming quadrate–triangular plate with three short blunt lobes, median the longer; apex with or without setal tuft. Pleonites 1–5, pleotelson free in both sexes. Maxillipeds 3 almost entirely covering buccal cavern. Male cheliped fingers with finely denticulate cutting edges, dactylus with square proximal tooth. Pereopods 2–5 dactyli evenly curved with 1 or 2 rows of short blunt teeth along most of flexor margin, most distal 2 sometimes longer than others. Gonopod 1 with swollen base tapering to finer distal part strongly, evenly curving dorsally through more than 120°, with acute apex directed ventrally, sitting in pocket on lateral ventral face of pleotelson when pleon closed. Maximum cl. 23 mm (Pl. 45c–e). Intertidal–slope (0–494 m). Temperate South America, Temperate Australasia, including subantarctic islands. 7 species (McLay 1988: ecology of New Zealand species; Poore et al. 2016: allocation of several species to other genera). One species, H.  planatus (­Fabricius, 1775) has been found among commercial oysters transported from Chile to Rio de Janeiro, Brazil (Tavares 2003).

Halimena Melrose, 1975 Diagnosis. Carapace rim 1.2 times as long as wide (rostrum excluded). Rostrum with convex lateral margins, tapering to rounded apex, depressed, 0.2 times carapace length; apex without long setae. Pleonites 1–5, pleotelson free in both sexes. Maxillipeds 3 almost entirely covering buccal cavern. Pereopods 2–5 dactyli curved, with 1 subdistal recurved tooth. Maximum cl. 6.2 mm. Subtidal, shelf (0–91 m). Temperate Australasia (New Zealand, endemic, rare). 1 species (Lucas and Hicks 1981; Melrose 1975).

Hymenicoides Kemp, 1917 Diagnosis. Carapace rim as long as wide (rostrum excluded), hymenosomian rim almost circular. Rostrum absent, dorsal surface surrounded by continuous hymenosomian rim. Pleonites 1–5, pleotelson free in both sexes; pleotelson with lateral lobes auriculate, inner surface of lobe rimmed laterally. Maxillipeds 3 separated by wide gape; maxilliped 3 pediform, with long propodus and dactylus, reaching or almost reaching posterior margin of buccal cavern. Pereopods 2–5 dactyli curved, with 1 subdistal recurved tooth. Gonopod 1 strongly bent, with complex apex including semicircular lip and denticulate lobe. Maximum cl. 6 mm. Subtidal. Western Indo-Pacific. 2 species (Naruse and Ng 2007: rediagnosis, key to species).

Hymenosoma Desmarest, 1823 Diagnosis. Carapace rim as wide or wider than long (rostrum excluded), or longer than wide, rounded-octagonal. Rostrum triangular. Epistome short or absent. Pleonites 1–5, pleotelson free in

547

both sexes. Maxillipeds 3 separated by wide gape. Pereopods 2–5 dactyli almost straight, with or without teeth on flexor margin. Gonopod 1 with stout base, distal part with blunt complex apex. Maximum cl. 28 mm. Estuarine, marine, intertidal, subtidal (0–37 m). Temperate Southern Africa, Temperate Australasia. 8 species (Dawson and Griffiths 2012: 5 South African species; Lucas 1980).

Insulaplax Ng, 2019 Diagnosis. Carapace rim wider than long (rostrum excluded); without postocular and lateral teeth. Rostrum minute, depressed. Pleonites 3–5 fused in male. Maxillipeds 3 separated by wide gape. Pereopods 2–5 dactyli without teeth on flexor margin. Gonopod 1 short, stout, medially twisted. Female with internal brood cavity incubating free embryos. Maximum cl. 2.8 mm. Freshwater. Guam. 1 species (Ng 2019).

Limnopilos Chuang & Ng, 1991 Diagnosis. Rostrum absent, dorsal surface surrounded by continuous hymenosomian rim. Pleotelson with lateral lobes smaller than medial lobe, lateral cavity of inner surface thickened distally. Maxillipeds 3 separated by wide gape; maxilliped 3 pediform, with long propodus and dactylus, reaching or almost reaching posterior margin of buccal cavern. Pereopods 2–5 dactyli with sharp teeth along flexor margin. Maximum cl. 6 mm. Freshwater streams. Indonesia, Thailand. 3 species (Chuang and Ng 1991; Naruse and Ng 2007; Naruse et al. 2008b: rediagnosis, key to species).

Lucascinus Poore, Guinot, Komai & Naruse, 2016 Diagnosis. Carapace rim as long as wide (rostrum excluded), narrower anteriorly. Rostrum rod-like, 0.25–0.46 times carapace length, similar in males, females, or triangular; apex with long setae. Pleonites 1–5, pleotelson free in both sexes. Maxillipeds 3 almost entirely covering buccal cavern. Male cheliped 2.5–4 times as long as carapace length; propodus elongate, cylindrical, fingers short, gaping at midpoint (‘nut-cracker’-like). Pereopods 2–5 dactyli mostly straight, with row of sharp spines along flexor margin. Gonopod 1 with swollen base, distal part stout, tapering to apex directed anteriorly, distally setose. Maximum cl. 4.4 mm. Estuarine, marine, intertidal, subtidal (0–3 m). Temperate Northern Pacific, Central Indo-Pacific, Temperate Australasia. 3 species (Poore et al. 2016).

Micas Ng & Richer de Forges, 1996 Diagnosis. Carapace rim as long as wide (rostrum excluded), narrower anteriorly. Rostrum short, triangular, with similar pair of pseudorostral teeth; apex without long setae. Pleonites 3–4 fused in both sexes. Maxillipeds 3 almost entirely covering buccal cavern. Pereopods 2–5 dactyli curved, with 2 subdistal teeth. Gonopod 1 with swollen base, narrow distal section tapering through sharp proximal ventral curvature and twist, with apex acute sharply bent ventrally. Maximum cl. 5.4 mm.

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Intertidal–slope shelf (0–244 m). Central Indo-Pacific, Temperate Australasia. 5 species (Ng and Richer de Forges 1996; Poore et al. 2016: rediagnosis, 5 species).

Nasutoplax Poore, Guinot, Komai & Naruse, 2016 Diagnosis. Carapace rim as long as wide (rostrum excluded), hymenosomian rim almost circular. Rostrum long, laterally compressed median spine with rounded apex, ventrally expanded, erect in lateral view, 0.35 times carapace length in male, 0.2 times in female; lower apex tubercle-like in male, angular in female; apex with long setae, or without long setae. Pleonites 1–5, pleotelson free in male; pleonites 3–5 fused in female. Maxillipeds 3 almost entirely covering buccal cavern. Male cheliped twice as long as carapace length, with swollen propodus with keel on upper margin, fixed finger with basal tooth, oblique distal cutting edge; dactylus with simple proximal tooth, distal cutting edge. Pereopods 2–5 dactyli with row of blunt teeth along most of flexor margin. Gonopod 1 with swollen base, almost straight distal part directed anteriorly, apex truncate, with subterminal blunt projection with spinulose tubercle on anterior face. Maximum cl. 5.5 mm. Subtidal, shelf (2–130 m). Temperate Australia. 1 species (Poore 2004; Poore et al. 2016).

Neohymenicus Lucas, 1980 Diagnosis. Carapace rim as long as wide (rostrum excluded), hymenosomian rim almost circular. Rostrum triangular; apex without long setae. Pleonites 1–5, pleotelson free in both sexes. Maxillipeds 3 almost entirely covering buccal cavern; maxilliped 3 merus longer than wide. Pereopods 2–5 dactyli curved, with subdistal blunt tooth. Gonopod 1 sickle-shaped, with stout base and thinner, strongly-curved distal portion, with a row of long subterminal setae. Maximum cl. 6.5 mm. Intertidal, subtidal (0–10 m). Temperate Australasia (New Zealand), Temperate Northern Pacific (Japan). 2 species (Lucas 1980; McLay 1988: ecology).

Neorhynchoplax Sakai, 1938 Diagnosis. Carapace rim longer than wide, rounded-octagonal; with lateral tooth on branchiostegite. Rostrum triangular with or without pair of similar pseudorostral teeth. Pleonites 3–5 fused in male. Maxillipeds 3 separated by wide gape. Pereopods 2–5 dactyli without teeth on flexor margin, or with sharp teeth along flexor margin. Gonopod 1 distal part curved. Female with internal brood cavity incubating free embryos. Maximum cl. 5 mm. Intertidal, subtidal (0–4 m), freshwater, estuarine, marine. Temperate Northern W Pacific, Western and Central IndoPacific. 33 species (Hsueh 2018: ecology, reproduction, distribution of species; Naruse et  al. 2008b: compared with other cave-dwelling genera; Rahayu et al. 2020: key to 9 Indian species).

Rhynchoplax Stimpson, 1858 Diagnosis. Carapace rim as wide or wider than long (rostrum excluded). Rostrum elongate-subovate in dorsal view, or narrow,

strongly flattened dorsoventrally, third to half as long as carapace in males, less than 0.2 times as long in females, with pair of shorter pseudorostral teeth; apex with long setae, or without long setae. Pleonites 3–4 fused in male; pleonites 3–5 fused, pleotelson fused to pleonite 5 or free in female. Maxillipeds 3 almost entirely covering buccal cavern. Male cheliped twice carapace length at most; fingers not forming ‘nut-cracker’ structure. Pereopods 2–5 dactyli falcate (pereopod 2 not always), each with row of sharp erect recurved teeth along most of flexor margin. Gonopod 1 with swollen base, distal part very slender, not coiled but strongly curved dorsally, terminating in acute apex directed ventrally. Maximum cl. 5 mm. Intertidal, subtidal (0–9 m). Temperate Northern W Pacific, Central Indo-Pacific. 4 species (Komai and Poore 2016: rediagnosis, 2 species; Poore et al. 2016: rediagnosis).

Samarplax Husana, Tan & Kase, 2011 Diagnosis. Carapace rim longer than wide, rounded-octagonal. Rostrum absent, dorsal surface surrounded by continuous hymenosomian rim. Pleonites 3–5 fused in male. Eyes fused to orbit, unpigmented. Maxillipeds 3 separated by wide gape. Pereopods 2–5 dactyli without teeth on flexor margin. Gonopod 1 sinuous. Female with internal brood cavity incubating free embryos. Maximum cl. 5.3 mm. Freshwater in caves. Samar, Philippines. 1 species (Husana et al. 2011).

Stimpsoplax Poore, Guinot, Komai & Naruse, 2016 Diagnosis. Carapace rim as wide or wider than long (rostrum excluded). Rostrum narrowly triangular, ~0.4 of carapace length in male, 0.25 in female, laterally setose, with pair of shorter pseudorostral teeth; apex without long setae. Pleonites 3–4 fused in male; pleonite 3–pleotelson fused in female. Maxillipeds 3 almost entirely covering buccal cavern. Male cheliped 2.5 as long as carapace length, with swollen propodus, fingers with narrow proximal setose gape when closed, cutting edges with row of low, blunt teeth, dactylus with prominent tooth remote from other teeth proximally. Pereopods 2–5 dactyli with row evenly spaced, backwardly directed spines on flexor margin; dactylus of pereopod 2 gently curving, those of pereopods 3–5 more strongly curving. Gonopod 1 with swollen base, narrow distal part strongly twisted, sharply curved ventrally over proximal third, tapering distally to curved apex, distal part directed ventrally, hooking inner side of fused distal pleonal segment. Maximum cl. 3.6 mm. Shelf (25–135 m). Temperate Northern W Pacific, Central Indo-Pacific. 3 species (Poore et al. 2016).

Sulaplax Naruse, Ng & Guinot, 2008 Diagnosis. Carapace rim wider than long (rostrum excluded). Rostrum vestigial, roundly produced downwards. Pleonites 3–5 fused in male. Eyes fused to orbit, unpigmented. Maxillipeds 3 separated by wide gape. Pereopods 2–5 dactyli without teeth on

14 – Brachyura – crabs

flexor margin. Gonopod 1 bent at midpoint. Female with internal brood cavity incubating free embryos. Maximum cl. 5 mm. Freshwater in caves. Sulawesi, Indonesia. 1 species (Naruse et al. 2008b).

Teramnonotus Tavares & Santana, 2015 Diagnosis. Carapace with lateral body wall not visible beyond hymenosomian rim; rim pear-shaped; subhepatic region with 2 broad tubercles. Rostrum triangular, upturned, continuous with carapace surface, ventral rostral keel absent. Pleonites 3–5 fused in female. Pereopods 2–5 dactyli curved, with 2 subdistal teeth. Maximum cl. 4 mm. Subtidal, shelf (8–40 m). Tropical W Atlantic, Central IndoPacific. 3 species (Tavares and Santana 2015).

Trigonoplax H. Milne Edwards, 1853 Diagnosis. Carapace with lateral body wall not visible beyond hymenosomian rim; rim triangular; subhepatic region smooth. Rostrum triangular, upturned, continuous with carapace surface, with ventral rostral keel. Maxillipeds 3 almost entirely covering buccal cavern. Pereopods 2–5 dactyli without teeth on flexor margin. Gonopod 1 distal part curved. Maximum cl. 10 mm. Intertidal–shelf (0–140 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia. 3 species (Lucas 1980; Poore 2004).

Subfamily Odiomarinae Guinot, 2011 Amarinus Lucas, 1980 Diagnosis. Carapace rim wider than long (rostrum excluded). Rostrum spade-like, deflected, ending in triangular tip. Pleonites 1–5, pleotelson free in both sexes. Maxillipeds 3 gaping at ischiummerus junction. Pereopods 2–5 dactyli without teeth on flexor margin. Gonopod 1 tapering abruptly distally, with 1 or more short distal lobes. Male uropod intercalated laterally at base of pleotelson. Maximum cl. 22 mm (Pl. 45a). Estuarine (0–2 m), freshwater in streams up to 1600 m asl. Central Indo-Pacific, Temperate Australasia. 11 species (Guinot 2011a: inclusion in Odiomarinae; Lucas 1980: diagnosis; McLay 1988: ecology of New Zealand species; Naruse et  al. 2008a: new species; Poore 2004: 3 Australian species).

Odiomaris Ng & Richer de Forges, 1996 Diagnosis. Carapace rim wider than long (rostrum excluded). Rostrum spade-like, deflected, ending in triangular tip. Pleonites 1–5, pleotelson free in both sexes. Maxillipeds 3 gaping at ischiummerus junction. Pereopods 2–5 dactyli without teeth on flexor margin. Gonopod 1 tapering, with 2 distal processes. Male uropod intercalated laterally at base of pleotelson. Maximum cl. 6 mm. Estuarine, freshwater. Central Indo-Pacific (New Caledonia). 2 species (Davie and Richer de Forges 1996: second species; Guinot 2011a: rediagnosis; Ng and Richer de Forges 1996: diagnosis).

549

Richerius Guinot & de Mazancourt, 2020 Diagnosis. Carapace rim as long as wide (rostrum excluded), hymenosomian rim almost circular. Rostrum spade-like, deflected, anterior margin concave in dorsal view. Pleonites 1–5, pleotelson free in both sexes. Maxillipeds 3 gaping at ischiummerus junction. Pereopods 2–5 dactyli without teeth on flexor margin. Gonopod 1 curved at base, without corneous process. Male uropod intercalated laterally at base of pleotelson. Maximum cl. 5 mm. Freshwater streams up to 500 m asl. New Caledonia. 1 species (Guinot and de Mazancourt 2020). References Ahyong ST, Lai JCY, Sharkey D, Colgan DJ, Ng PKL (2007) Phylogenetics of the brachyuran crabs (Crustacea: Decapoda): the status of Podotremata based on small subunit nuclear ribosomal RNA. Molecular Phylogenetics and Evolution 45, 576–586. doi:10.1016/j.ympev.2007.03.022 Chuang CTN, Ng PKL (1991) Preliminary descriptions of one new genus and three new species of hymenosomatid crabs from Southeast Asia (Crustacea: Decapoda: Brachyura). Raffles Bulletin of Zoology 39, 363–368. Chuang CTN, Ng PKL (1994) The ecology and biology of Southeast Asian false spider crabs (Crustacea: Decapoda: Brachyura: Hymenosomatidae). In Ecology and Conservation of Southeast Asian Marine and Freshwater Environments including Wetlands. (Eds Sasekumar A, Marshall N, Macintosh DJ) pp. 85–92. Springer, Dordrecht. Davie PJF, Richer de Forges B (1996) Two new species of false spider crabs (Crustacea: Brachyura: Hymenosomatidae) from New Caledonia. Memoirs of the Queensland Museum 39, 257–262. Dawson J, Griffiths CL (2012) Revision of the crown crabs, genus Hymenosoma (Crustacea: Hymenosomatidae), of South Africa. African Natural History 8, 16–29. Guinot D (2011a) Odiomarinae nov. subfam., a new subfamily for two primitive genera of Hymenosomatidae MacLeay, 1838, with preliminary remarks on the family (Crustacea, Decapoda, Brachyura). Zootaxa 2732, 20–32. doi:10.11646/zootaxa.2732.1.2 Guinot D (2011b) The position of the Hymenosomatidae MacLeay, 1838, within the Brachyura (Crustacea, Decapoda). Zootaxa 2890, 40–52. doi:10.11646/zootaxa.2890.1.4 Guinot D, de Mazancourt V (2020) A new freshwater crab of the family Hymenosomatidae MacLeay, 1838 (Crustacea, Decapoda, Brachyura) and an updated review of the hymenosomatid fauna of New Caledonia. European Journal of Taxonomy 671, 1–29. doi:10.5852/ejt.2020.671 Guinot D, Richer de Forges B (1997) Affinités entre les Hymenosomatidae MacLeay, 1838 et les Inachoididae Dana, 1851 (Crustacea, Decapoda, Brachyura). Zoosystema 19, 453–502. Guinot D, Tavares M, Castro P (2013) Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa. Zootaxa 3665, 1–414. doi:10.11646/ zootaxa.3665.1.1 Hsueh PW (2018) A new species of Neorhynchoplax (Crustacea: Decapoda: Brachyura: Hymenosomatidae) from Taiwan. Zootaxa 4461, 350–358. doi:10.11646/zootaxa.4464.3.2 Husana DEM, Tan SH, Kase T (2011) A new genus and species of anchialine Hymenosomatidae (Crustacea, Decapoda, Brachyura) from Samar, Philippines. Zootaxa 3109, 49–59. doi:10.11646/zootaxa.3109.1.3 Komai T, Poore GCB (2016) Resurrection of Rhynchoplax Stimpson, 1858, with the description of two new species from Japan and Australia

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(Crustacea: Decapoda: Brachyura: Hymenosomatidae). Zootaxa 4093, 515–524. doi:10.11646/zootaxa.4093.4.3 Kulabtong S, Kunlapapuk S, Sottiyothin S (2013) A new species of false spider crab of the genus Elamena H. Milne Edwards, 1837 from Upper Gulf of Thailand (Decapoda Hymenosomatidae). Biodiversity Journal 4, 129–134. Lucas JS (1980) Spider crabs of the family Hymenosomatidae (Crustacea; Brachyura) with particular reference to Australian species: systematics and biology. Records of the Australian Museum 33, 148–247. doi:10. 3853/j.0067-1975.33.1980.199 Lucas JS, Davie PJF (1982) Hymenosomatid crabs of Queensland estuaries and tidal mud flats, including descriptions of four new species of Elamenopsis A. Milne Edwards and a new species of Amarinus Lucas. Memoirs of the Queensland Museum 20, 401–419. Lucas JS, Hicks GRF (1981) Further details on the male of the poorly known hymenosomatid crab Halimena aotearoa Melrose. National Museum of New Zealand Records 2, 1–4. McLay CL (1988) Brachyura and crab-like Anomura of New Zealand [Crabs of New Zealand]. Leigh Laboratory Bulletin 22, 1–463. Melrose MJ (1975) The marine fauna of New Zealand: family Hymenosomatidae (Crustacea, Decapoda, Brachyura). New Zealand Oceanographic Institute Memoir 34, 1–123. Naruse T, Ng PKL (2007) On the taxonomy of the genus Hymenicoides Kemp, 1917 (Crustacea: Decapoda: Brachyura: Hymenosomatidae), with resurrection of Limnopilos Chuang & Ng, 1991, and descriptions of two new species. Zootaxa 1621, 17–31. doi:10.11646/ zootaxa.1621.1.2 Naruse T, Mendoza JCE, Ng PKL (2008a) Descriptions of five new species of false spider crabs (Decapoda: Brachyura: Hymenosomatidae) from the Philippines. Marine Biology Research 4, 429–441. doi:10.1080/​ 17451000802232866 Naruse T, Ng PKL, Guinot D (2008b) Two new genera and two new species of troglobitic false spider crabs (Crustacea: Decapoda: Brachyura: Hymenosomatidae) from Indonesia, with notes on Cancrocaeca Ng, 1991. Zootaxa 1739, 21–40. doi:10.11646/zootaxa.1739.1.2 Ng PKL (1991) Cancrocaeca xenomorpha, new genus and species, a blind troglobitic freshwater hymenosomatid (Crustacea: Decapoda: Brachyura) from Sulawesi, Indonesia. Raffles Bulletin of Zoology 39, 59–73. Ng PKL (2019) On the taxonomy of two rare freshwater crabs (Decapoda, Brachyura, Chasmocarcinidae and Hymenosomatidae) from Palau, western Pacific. Crustaceana 92, 979–996. doi:10.1163/​1568540300003917 Ng PKL, Chuang CTN (1996) The Hymenosomatidae (Crustacea: Decapoda: Brachyura) of southeast Asia, with notes on other species. Raffles Bulletin of Zoology Supplement 3, 1–82. Ng PKL, Lukhaup C (2015) Aletheiana tenella, a new genus and new species of freshwater hymenosomatid crab (Crustacea: Decapoda: Brachyura) from Central Sulawesi, Indonesia. Zootaxa 4039, 118–128. doi:10.11646/zootaxa.4039.1.4 Ng PKL, Rahayu DL (2022) A new genus and new species of Hymenosomatidae (Crustacea: Brachyura) from Indonesian Papua. Zootaxa 5092, 480–486. doi:10.11646/zootaxa.5092.4.7 Ng PKL, Richer de Forges B (1996) The Hymenosomatidae (Crustacea: Decapoda: Brachyura) of New Caledonia, with descriptions of two new genera and two new species. Memoirs of the Queensland Museum 39, 263–276. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Poore GCB (2010) Elamenopsis guinotae, a new spider crab from Bass Strait, Australia (Brachyura: Hymenosomatidae). In: Castro P, Davie

PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 261–268. Poore GCB, Guinot D, Komai T, Naruse T (2016) Reappraisal of species attributed to Halicarcinus White, 1846 (Crustacea: Decapoda: Brachyura: Hymenosomatidae) with diagnosis of four new genera and one new species from New Ireland, Papua New Guinea. Zootaxa 4093, 480–514. doi:10.11646/zootaxa.4093.4.2 Rahayu DL, Ng PKL (2019) A new species of Elamenopsis from Papua, Indonesia (Decapoda, Brachyura, Hymenosomatidae). Crustaceana 92, 537–544. doi:10.1163/15685403-00003883 Rahayu DL, Nugroho DA (2019) Three new species of the genus Elamena H. Milne Edwards, 1837 (Crustacea: Decapoda: Hymenosomatidae) from Lombok Island, Indonesia. Raffles Bulletin of Zoology 67, 171–182. Rahayu DL, Jayachandran PR, Jima M, Bijoy Nandan S, Hershey NR, et al. (2020) Notes on Neorhynchoplax alcocki (Kemp 1917) and Neorhynchoplax demeloi (Kemp 1917) from the Kochi Backwater, India (Crustacea: Decapoda: Brachyura: Hymenosomatidae). Thalassas: An International Journal of Marine Sciences 36, 79–84. doi:10.1007/ s41208-019-00176-w Richer de Forges B (1993) Deep sea crabs of the Tasman seamounts (Crustacea: Decapoda: Brachyura). Records of the Australian Museum 45, 11–24. doi:10.3853/j.0067-1975.45.1993.126 Tavares M (2003) On Halicarcinus planatus (Fabricius) (Brachyura, Hymenosomatidae) transported from Chile to Brazil along with the exotic oyster Crassostrea gigas (Thunberg). Nauplius 11, 45–50. Tavares M, Santana W (2015) A new genus and two new species of hymenosomatid crabs (Crustacea: Brachyura: Hymenosomatidae) from the southwestern Atlantic and eastern Australia. Zootaxa 3905, 91–106. doi:10.11646/zootaxa.3905.1.5

Leucosioidea Samouelle, 1819 Leucosioidea include two families of crabs characterised by the inhalent (= afferent) respiratory channel lying alongside the buccal cavern, opening below the orbits, and covered by the exopod of the third maxilliped. Leucosioids are common in shallow coral reef and seagrass environments in tropical to temperate waters. The division of the group into two families on differences in the female pleon is well supported (Guinot and Bouchard 1998; Ng et al. 2008). Diagnosis. Carapace subcircular to ovoid, pyriform or pentagonal, regions often undefined, surface usually smooth, often armed, sometimes with spines or lobes. Opening for inhalent respiratory current below frontal margin or orbits, adjacent to endostome, with distinct canal (inhalent branchial channel) along sides of buccal cavity beneath maxilliped 3 exopod, visible when maxillipeds 3 pushed aside. Antennule folding obliquely. Buccal cavity trapezoid-triangular, margins distinctly convergent anteriorly. Maxilliped 3 merus triangular, tapering distally, rarely subquadrate–hexagonal. Male pleon usually with some pleonites fused. Gonopod 1 usually slender, often twisted, variously ornamented apically.

14 – Brachyura – crabs

551

Key to families of Leucosioidea 1. –

Female with all pleonites freely articulating, not forming brood-chamber with thoracic sternum, egg-mass protruding from sides of pleon when ovigerous������������������������������������������������������������������������������������������������� Iphiculidae Female usually with most pleonites fused, forming brood-chamber with thoracic sternum, egg-mass not visible when ovigerous (Fig. 14.67)������������������������������������������������������������������������������������������������������������������������Leucosiidae

References

Pariphiculus Alcock, 1896

Guinot D, Bouchard J-M (1998) Evolution of the abdominal holding systems of brachyuran crabs (Crustacea, Decapoda, Brachyura). Zoosystema 20, 613–694. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286.

Iphiculidae Alcock, 1896 Figure 14.61, Plate 45g–i Iphiculids are recognisable by the combination of fine, more-or-less pectinate fingers on the chelipeds and the spongiose covering of the carapace and legs. They differ from leucosiids in not having the female pleon fused and dome-shaped. Diagnosis. Female with all pleonites freely articulating, not forming brood-chamber with thoracic sternum, eggmass protruding from sides of pleon when ovigerous. Chela with fine, more-or-less pectinate fingers (Fig. 14.61b–d). Iphiculus Adams & White, 1849 Diagnosis. Carapace wider than long, anterolateral margin with prominent teeth, largest at most lateral extreme (Fig.  14.61b), or with small widely spaced teeth (Fig.  14.61a). Cheliped dactylus opening through arc of ~120°. Maximum cl. 15 mm (Pl. 45g). Subtidal–shelf (11–204 m). Western and Central Indo-Pacific. 2 species (Ihle 1918: figure; Naruse et  al. 2017: figures, distribution).

Diagnosis. Carapace globular or oval, anterolateral margins dentate or spinose (Fig.  14.61c). Cheliped dactylus opening through arc of ~70°. Maximum cl. 32 mm (Pl. 45h, i). Shelf–slope (26–296 m). Temperate Northern W Pacific, Central Indo-Pacific. 5 species (Chen 1989: key to 3 species; Ng and Jeng 2017: fifth species). References Chen H-L (1989) Leucosiidae (Crustacea, Brachyura). In: Forest J (Ed.), Résultats des Campagnes MUSORSTOM, vol. 5. Mémoires du Muséum National d’Histoire Naturelle, Paris 144, 181–264. Ihle JEW (1918) Die Decapoda Brachyura der Siboga-Expedition. III. Oxystomata: Calappidae, Leucosiidae, Raninidae. Siboga-Expéditie 39b2, 159–322. Naruse T, Maenosono T, Uyeno D, Samejima S, Shirawakawa N (2017) Records of six leucosioid crab species from the Ryukyu Archipelago, Japan. Fauna Ryukyuana 36, 1–11. Ng PKL, Jeng MS (2017) Notes on two crabs (Crustacea, Brachyura, Dynomenidae and Iphiculidae) collected from red coral beds in northern Taiwan, including a new species of Pariphiculus Alcock, 1896. ZooKeys 694, 135–156. doi:10.3897/zookeys.694.14871

Leucosiidae Samouelle, 1819 pebble crabs, nut crabs Figures 14.62–14.70, Plates 46–48 Leucosiidae, aptly known as nut crabs or pebble crabs, are a readily recognisable family of small crabs that are common on soft sediments and coral rubble from the intertidal shore

Fig. 14.61.  Iphiculidae. a, Iphiculus convexus Ihle, 1918 (carapace); b, Iphiculus spongiosus Adams & White, 1849; c, Pariphiculus agariciferus Ihle, 1918; d, chela, Pariphiculus.

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Marine Decapod Crustacea

to slope depths. The family is commonly encountered in the Indo-West Pacific where almost 500 species in 60 genera can be found. Fewer genera and species can be found in the Americas. In temperate environments leucosiids inhabit seagrass meadows in shallow waters and less commonly, rocky reefs. On tropical coral reefs some species have a highly eroded and flattened carapace that mimics broken coral (Ng and Komatsu 2016) or fragments of the alga Halimeda (Tan and Richer de Forges 1993). While the systematics of some groups of genera is well understood, their higher relationships are controversial. Early workers (Alcock 1896; Bell 1855; Ihle 1918; Miers 1886) recognised at least four subfamilies, Cryptocneminae, Ebaliinae, Leucosiinae and Philyrinae, but their differentiation is difficult and historically has been extremely fluid (Galil and Ng 2020; Manning and Holthuis 1981). Leucosiinae are perhaps the best delimited by the possession of a thoracic sinus on the lateral margin of the carapace. The remaining subfamilies as used historically intergrade morphologically and are unlikely to represent natural groups (Ng et  al. 2008). Additionally, the generic placement of some species of the largest genera, Ebalia, Nursia, Philyra and Cryptocnemus, requires review. Genera usually assigned to Cryptocneminae have a broad carapace, with lamelliform expansions wrapping around the lateral and posterior margins overhanging most of walking legs, but not all genera that have been historically placed there conform, and Cryptocnemus is similar to members of a group of ebaliine genera (Nursia, Paranursia, Nobiliella) traditionally excluded (Galil and Ng 2020). The cryptocnemines are probably derived from within the ebaliines. Distinguishing Ebaliinae and Philyrinae has been historically problematical, so recent authors have merged the latter with the former (e.g. Davie et  al. 2015; Ng et  al. 2008), a decision followed here. Nevertheless, some of the features previously used to distinguish the philyrines from ebaliines, namely the relative positions of the anterior margin of the endostome and the anterior margin of the inhalant branchial channel, remain useful as key characters. Most genera previously considered as philyrine have the anterior margins of the inhalant branchial channels and endostome transversely aligned, reaching anteriorly to about the same level. In other genera, the anterior margin of the endostome projects beyond the anterior margin of the inhalant branchial channel, including in Nursia and Nursilia which were usually considered to be philyrines (e.g. Chen and Sun 2002; Sakai 1976). In recognising Cryptocneminae and Ebaliinae here, some pragmatic taxonomic changes are introduced. For Cryptocneminae, we emphasise the thin, sharp-edged, expansions of the carapace margins and smooth, polished dorsal surface (apart from the granular median and

epibranchial ridges, when present). As such, we follow Chen and Sun (2002) in treating Paranursia as a cryptocnemine (Fig. 14.62). Likewise, we transfer Leucisca from Cryptocneminae to Ebaliinae because the expanded carapace margins are blunt and somewhat thickened (rather than thin, bladelike), and the dorsal surface is finely granulate (rather than smooth) (Fig. 14.64d, e). Two Australian species originally described in Phlyxia but historically assigned to Ebalia, E. crassipes (Bell, 1855) and E. lambriformis (Bell, 1855), do not correspond to Ebalia because the anterior margins of the inhalant branchial channels and endostome are transversely aligned; we resurrect Phlyxia for their reception, of which P. crassipes Bell, 1855 is the type species. Two other ebaliine genera, Nucia and Heteronucia, supposedly separated by vertical-oblique or horizontal plane of movement of the cheliped fingers (Chen 1989; Chen and Sun 2002), are unsatisfactorily difficult to distinguish on this criterion, given the vagaries of orientation during observation (Tan 1996). We herein focus on a correlated feature that is more easily observed: the relative lengths of the cheliped dactylus and dorsal margin of the propodal palm (dactylus longer in Heteronucia, shorter in Nucia). Accordingly, we transfer Nucia bouvieri Ihle, 1918 to Heteronucia, and Heteronucia elegans Chen & Türkay, 2001, and H. xincunensis Chen & Türkay, 2001, to Nucia. The present subfamilial arrangement is probably artificial, but serves as a primarily pragmatic framework in lieu of phylogenetic analyses through which natural subdivisions can be identified. Some aspects of the diagnosis of leucosiid genera need explanation for the uninitiated. The thoracic sinus of Leucosia and related genera is a groove on the lateral-underside of the carapace beginning in front of the base of the cheliped which serves as a channel for water to the gills; its margins are often granular (Fig. 14.70c–g). The buccal cavern is covered by maxillipeds 3 and the central portion is covered by the maxilliped 3 endopods, of which the anterior portion in front of the mandibles is the endostome. On each side of the endostome, covered by the maxilliped 3 exopod, are the inhalant branchial channels, shallow conduits through which the crab draws water for respiration. The armature and position of the anterior margin of the inhalant branchial channel (whether in line with or positioned behind the level of the endostome) are important key characters (Fig. 14.66a–j). The anterior margin of the inhalant branchial channel is separated from the suborbital margin by a groove or distinct space in most genera (e.g. Randallia), but in some genera, the two structures are appressed (e.g. Arcania, Ixa, Raylilia) (Fig.  14.66n, p), or even indistinguishably merged (e.g. Persephona, Myropsis). Recognising the distinction between the two structures is important because the presence of either two or three teeth/

14 – Brachyura – crabs

lobes on the anterior margin of the inhalant branchial channel is diagnostic, so care must be taken in assessing the number. When the anterior margin of the inhalant branchial channel is separated from the suborbital margin, the lobes/spines on each structure are distinguishable (e.g, Fig.  14.66m, o). However, when the two structures are appressed, the two lobes/spines of the inhalant branchial channel abut the suborbital spine/lobe giving the appearance of three spines/lobes (e.g. Fig.  14.66n, p). Careful examination (from anteroventral view), however, will reveal

553

that the suborbital spine is on a separate plane, behind the two lobes/spines of the inhalant branchial channel. Eleven genera occur only in the Americas, principally in the tropical Western Atlantic and Eastern Pacific. Fiftynine genera occur only in the Indo-West Pacific, Australasia, southern and West Africa. Only Ebalia and Tanaoa occur in both regions. Diagnosis. Female pleonites mostly fused (rarely all free), forming brood-chamber with thoracic sternum, fully enclosing egg-mass.

Key to subfamilies and genera of Leucosiidae 1.

Carapace thoracic sinus present (Fig. 14.70c–h); dorsal surface highly polished (Fig. 14.70a–d)������������������������������������������������������������������������������������������������������������������������������������������������ Leucosiinae … 2 – Carapace thoracic sinus absent; dorsal surface variously sculptured or smooth (Figs 14.62–14.65)����������������������������������8 2. Carapace thoracic sinus anterior margin defined by beaded loop (Fig. 14.70c, e)����������������������������������������������� Euclosiana – Carapace thoracic sinus anterior margin open, without or with few granules (Fig. 14.70d, f, g)����������������������������������������3 3. Male pleonites 3–6 fused, with constriction at suture 5/6 (Fig. 14.70m)�������������������������������������������������������������������� Seulocia – Male pleonites 3–5 fused, anterolaterally inflated; pleonite 6 articulating (Fig. 14.70j–l, n)�����������������������������������������������4 4. Male pleonite 6 without median denticle (Fig. 14.70l)����������������������������������������������������������������������������������������������������� Liusius – Male pleonite 6 with median denticle (Fig. 14.70n)��������������������������������������������������������������������������������������������������������������������5 5. Female maxilliped 3 endopod without longitudinal fringe of setae (Fig. 14.70i). Gonopod 1 straight, uncoiled (Fig. 14.70u)��������������������������������������������������������������������������������������������������������������������������������������������������Urnalana – Female maxilliped 3 endopod with longitudinal fringe of setae (Fig. 14.70h). Gonopod 1 coiled at least once���������������6 6. Gonopod 1 coiled once, apical process encased in setose muff (Fig. 14.70t)������������������������������������������������������������� Soceulia – Gonopod 1 coiled 3 times or more, ultimate coil swollen, setose; apical process cornute���������������������������������������������������7 7. Gonopod 1 coiled 3 times (Fig. 14.70o)��������������������������������������������������������������������������������������������������������������������������� Coleusia – Gonopod 1 tightly-coiled 5 or more times; apical process conical, with swollen setose muff (Fig. 14.70q)�������������������������������������������������������������������������������������������������������������������������������������������������������������������� Leucosia 8. Carapace lateral and posterior margins expanded, with thin sharp edge, forming continuous laminar rim overhanging proximal part of pereopods and pleon; dorsal surface smooth, appearing polished, unornamented or at most with low, granular median and oblique epibranchial ridges (Fig. 14.62)���������������������������������������������������������������������������������������������������������������������������������������������� Cryptocneminae … 9 – Carapace variously globose or angular; lateral margins unexpanded, or if expanded, edge blunt, not thin-sharp, often tubercular or beaded, rarely forming continuous rim overhanging branchial and posterior margins; dorsal surface granular or punctate, often sculptured; median and oblique epibranchial ridges, if present, usually prominent (Figs 14.63–14.68)������������������������������������������������������Ebaliinae … 11 9. Carapace longer than wide (Fig. 14.62c). Cheliped palm with inner lower surface densely setose����������� Onychomorpha – Carapace as long as or wider than long. Cheliped palm with inner lower surface glabrous����������������������������������������������10 10. Gonopod 1 with 3 distal points (Fig. 14.62i). Carapace as long as wide or little wider than long, lateral margin sinuous with concavity behind midlength; front weakly produced, sinuous (Fig. 14.62d, e)������������Paranursia – Gonopod 1 with 1 or 2 distal points or lobes (Fig. 14.62f–h). Carapace wider than long and front weakly produced, strongly produced or as two horn-like lobes, if carapace little wider than long and lateral margin with concavity behind midlength, then front forming two horn-like lobes (Fig. 14.62a, b)������������������������������������������������������������������������������������������������������������������������������������������������������ Cryptocnemus 11. Endostome (covered by maxilliped 3 endopod) reaching beyond anterior margin of inhalant branchial channel (covered by maxilliped 3 exopod) (Fig. 14.66b–f)����������������������������������������������������������������������������������������������� 12 – Endostome reaching level of inhalant branchial channel (covered by maxilliped 3 exopod) (Fig. 14.66a)���������������������� 40

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Marine Decapod Crustacea

12. Cheliped fingers 1.6–1.7 times as long as palm, slender, curved in dorsal view, cutting edges with unequal perpendicular spines (Fig. 14.66x, y)��������������������������������������������������������������������������������������������������������������������������������������13 – Cheliped fingers shorter than to 1.8 (usually less than 1.6) times as long as palm, robust, straight in dorsal view, cutting edges dentate or crenulated, not spinose�������������������������������������������������������������������������������������������������������14 13. Carapace distinctly wider than long (Fig. 14.63k). Male pleonites 3–5 fused, pleonite 6 free. Female pleonites all free (Fig. 14.67a)���������������������������������������������������������������������������������������������������������������������������Coralliocryptus – Carapace about as wide as long (Fig. 14.64r). Male and female pleonites 3–6 fused������������������������������������������������ Nursilia 14. Carapace dorsal surface with 4 parallel longitudinal granular ridges (Fig. 14.65g)�������������������������������������������������� Praosia – Carapace dorsal surface at most with single median longitudinal ridge and diagonal ridges radiating from centre�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������15 15. Orbit mesially open, not closed-off from antennular fossa, basal antennal article movable (Fig. 14.66e)�����������������������16 – Orbit mesially closed-off from antennular fossa by immovable (or very slightly movable) basal antennal article (Fig. 14.66f)��������������������������������������������������������������������������������������������������������������������������������������������������������������������21 16. Carapace rectangular–triangular; branchial region with strong oblique or transverse epibranchial ridge extending posterolaterally, widest posterolaterally�������������������������������������������������������������������������������������������������������������17 – Carapace oval or irregularly polygonal, surface finely granular to prominently tubercular or spinous, epibranchial ridge, if present rarely reaching widest part of carapace�����������������������������������������������������������������������������18 17. Gonopod 1 scarcely distally bulbous with slender apical process (Fig. 14.69e)���������������������������������������������������� Atlantotlos – Gonopod 1 evenly tapering to sharp point (Fig. 14.69k)��������������������������������������������������������������������������������������� Merocryptus 18. Cheliped merus in adults about as long as or shorter than carapace, propodus stout��������������������������������������������������������19 – Cheliped merus in adults exceeding 1.5 times as long as carapace, propodus slender������������������������������������������������������ 20 19. Carapace highest point in midline; surface ranging from finely granular to coarsely tuberculate, lobate or spinous, but without substantial areas of deep, irregular erosions and cavities (Fig. 14.63m)����������������������������� Ebalia – Carapace highest point on branchial regions; surface with substantial areas of deep, irregular or convoluted excavations, often with deep pits and erosions (Fig. 14.64h)��������������������������������������������������������������Lithadia 20. Carapace lateral margin with tooth at midpoint; posterior margin with 2 or 3 teeth (Fig. 14.65f). Male pleonites 3–5 fused�����������������������������������������������������������������������������������������������������������������������������������������Praebebalia – Carapace lateral margin rounded at midpoint; posterior margin straight (Fig. 14.63d). Male pleonites 3–6 fused (Fig. 14.68b)��������������������������������������������������������������������������������������������������������������Ancylodactyla 21. Carapace rectangular–triangular–oval, usually wider than long, dorsal surface usually highly sculptured or eroded, with deep cavities�������������������������������������������������������������������������������������������������������������������������������������������������� 28 – Carapace pentagonal, hexagonal, or subcircular-ovate, wider than long to longer than wide, dorsal surface smooth or variously granulate or tuberculate, not highly sculptured, without deep cavities and erosions������������������ 22 22. Carapace lateral margins expanded laterally and also sometimes posteriorly�������������������������������������������������������������������� 23 – Carapace lateral margins neither expanded laterally nor posteriorly����������������������������������������������������������������������������������� 25 23. Carapace transversely ovate; surface finely and evenly granular; front strongly produced forward beyond orbits (Fig. 14.64e)����������������������������������������������������������������������������������������������������������������������������������������������������������Leucisca – Carapace outline angular, deltoid to polygonal; front at most weakly produced anteriorly beyond orbits�������������������� 24 24. Carapace supraorbital margin with 2 fissures. Male pleonites 3–6 fused (Fig. 14.68k). Female pleonite 1 fully visible (Fig. 14.67g)��������������������������������������������������������������������������������������������������������������������������������������������������Nursia – Carapace supraorbital margin with 1 fissure. Male pleonites 2–6 fused Female pleonite 1 concealed medially by overlap of pleonite 2 (Fig. 14.66f)���������������������������������������������������������������������������������������������������������Nobiliella 25. Pereopods 2–4 smooth (Figs 14.63t, 14.64o, p)������������������������������������������������������������������������������������������������������������������������� 26 – Pereopods 2–4 tuberculate����������������������������������������������������������������������������������������������������������������������������������������������������������� 27 26. Carapace outline subcircular-subhexagonal, slightly wider than long; posterior margin straight, unarmed, lateral margin evenly convex, tuberculate (Fig. 14.64p). Female pleonites 3–6 fused������������������������������������������Nuciops – Carapace longitudinally ovate; posterior margin with 2 or 4 teeth, sometimes with intestinal tooth, lateral margin with rounded teeth (Fig. 14.63t). Female pleonites all free (Fig. 14.67d)�����������������������������������������������������Ihleus

14 – Brachyura – crabs

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27. Cheliped fingers shorter than dorsal margin of palm, opening almost horizontally (Fig. 14.64o)��������������������������� Nucia – Cheliped fingers longer than dorsal margin of palm, opening vertically or obliquely (Fig. 14.63r)��������������� Heteronucia 28. Carapace posterior half with single U-shaped, 2 or 4 deep cave-like hollows, most evident when viewed posteriorly��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 29 – Carapace surface variously eroded and sculptured, but without deep cave-like hollows on posterior half������������������� 30 29. Carapace subpentagonal–suboctagonal, with 2 or 4 deep hollows or cavities visible from posterior view (Fig. 14.65n, o). Male pleonites 3–5 fused, pleonite 6 with strong spine directed anteriorly (Fig. 14.68t)����������������������������������������������������������������������������������������������������������������������������������������������������������� Speloeophorus – Carapace subhexagonal, with 4 deep hollows on posterior half. Male pleonites 3–6 fused, pleonite 6 without spine (Fig. 14.68s)�����������������������������������������������������������������������������������������������������������������������������Speloeophoroides 30. Carapace dorsal surface with 8 raised bosses (cardiac, intestinal, 2 gastric, 4 branchial), each rimmed by tubercles, with 3 leaf-shaped branchial teeth similarly beaded (Fig. 14.64u)��������������������Orientotlos – Carapace otherwise ornamented�������������������������������������������������������������������������������������������������������������������������������������������������31 31. Carapace twice as wide as long (including produced intestinal region), anterior and posterolateral margins parallel, lateral margin evenly convex, surface honey-combed, without ridges (Fig. 14.63o)�������������������������������� Favus – Carapace at most 1.6 times as wide as long, not ornamented as above���������������������������������������������������������������������������������32 32. Antennular basis occupying entire fossa (Fig. 14.66k)�������������������������������������������������������������������������������������������������������������33 – Antennular basis occupying lower half of fossa (Fig. 14.66l)��������������������������������������������������������������������������������������������������35 33. Carapace with granule-filled groove parallel to anterolateral margin, with pair of postorbital prominences; lateral margin expanded, not thin and plate-like (Fig. 14.63b, c). Cheliped fingers ~1.0–1.6 times as long as palm. Gonopod l 1.6–4.3 times as long as gonopod 2 (Fig. 14.69b)������������������������������������������������������������������������� Alox – Carapace without submarginal groove; lateral and/or frontal margin, without pair of postorbital prominences; lateral margin expanded, thin and plate-like. Cheliped fingers various. Gonopod l 2.0–2.5 times as long as gonopod 2����������������������������������������������������������������������������������������������������������������������������������������������������� 34 34. Carapace margin upturned to form cup-shaped structure, median longitudinal ridge distinct, narrow; front entire; lateral margin with open sutures (Fig. 14.65p). Male pleon coarsely granular (Fig. 14.68w)������������������������� Tlos – Carapace margins not upturned, median longitudinal ridge indistinct, broad; front bilobed; lateral margin with closed sutures (Fig. 14.63l). Male pleon finely granular (Fig. 14.68e)����������������������������������������������������������������Dolos 35. Cheliped fingers shorter than dorsal margin of palm (Fig. 14.65r, s)���������������������������������������������������������������������������� Uhlias – Cheliped fingers longer than dorsal margin of palm�������������������������������������������������������������������������������������������������������������� 36 36. Carapace with thick, irregular, median and diagonal or transverse epibranchial ridges (Fig. 14.64c, i). Maxilliped 3 exopod with angular tooth at outer proximal margin (Fig. 14.66s)����������������������������������������������������������37 – Carapace sculpture irregular, eroded, but without median and epibranchial ridges. Maxilliped 3 exopod without tooth at outer proximal margin������������������������������������������������������������������������������������������������������������������������������ 38 37. Carapace anterolateral margins irregular but produced anteriorly to form an almost right angle giving carapace approximately rectangular outline (Fig. 14.64c)��������������������������������������������������������������������������������������� Kabutos – Carapace anterolateral margins irregular to rounded but sloping posteriorly giving carapace approximately triangular or polygonal outline (Fig. 14.64i)�����������������������������������������������������������������������Merocryptoides 38. Carapace dorsal surface with large fossae on anterior half, connected by subsurface channels, floor of fossae covered with mushroom-like granules; lateral margins not swollen to form rim (Fig. 14.64s). Cheliped fixed finger massive, blade-like (Fig. 14.66z). Gonopod l ~3 times as long as gonopod 2 (Fig. 14.69l)����������������������Oreophorus – Carapace dorsal surface without fossae on anterior half; lateral margins swollen to form rim or not. Cheliped fixed finger not massive or blade-like. Gonopod l 2.1–2.4 times as long as gonopod 2 (Fig. 14.69g, m)�������������������� 39 39. Male pleon with short transverse ridge on fused segment, telson arrow-shaped (Fig. 14.68d). Gonopod l short, stout, with densely setose distal half (Fig. 14.69g). Carapace lateral margins not expanded to form rim (Fig. 14.63j)����������������������������������������������������������������������������������������������������������������������������������������������������������������Cateios – Male pleon with tubercle on fused segment, telson triangular (Fig. 14.68l). Gonopod l long, narrow, sparsely setose distal half (Fig. 14.69m). Carapace lateral margins expanded to form conspicuous rim (Fig. 14.64t)���������������������������������������������������������������������������������������������������������������������������������������������������������������������� Oreotlos

556

Marine Decapod Crustacea

40. Buccal cavern narrowly triangular. Maxilliped 3 exopod narrower than endopod ischium at midlength (Fig. 14.66g, i)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������41 – Buccal cavern transversely broad. Maxilliped 3 exopod as wide as or wider than endopod ischium at midlength (Fig. 14.66j, t, u, w)����������������������������������������������������������������������������������������������������������������������������������������������� 60 41. Anterior margin of inhalant branchial channel with 3 teeth or lobes (Fig. 14.66m)���������������������������������������������������������� 42 – Anterior margin of inhalant branchial channel with no more than 2 teeth or lobes (Fig. 14.66n–p)����������������������������� 50 42. Carapace, cheliped merus and thoracic sternum covered with flat-topped mushroom-shaped tubercles. Carapace dorsal surface with broad median ridge, paired gastric and hepatic bosses, median cardiac and intestinal bosses; subhepatic margin visible in dorsal view (Fig. 14.63q)������������������������������������������� Heterolithadia – Carapace, cheliped merus and thoracic sternum smooth or covered with rounded granules, tubercles or spines; subhepatic margin not usually visible in dorsal view�������������������������������������������������������������������������������������������� 43 43. Carapace posterolateral and posterior margins together with 0, 2, 4 or 7 spines or projections�������������������������������������� 44 – Carapace posterolateral and posterior margins together with 3 spines or projections������������������������������������������������������ 47 44. Carapace posteriorly with 7 spines or protuberances (3 in posterior midline; 2 pairs around posterolateral and posterior margins) (Fig. 14.63i)��������������������������������������������������������������������������������������������������������������������Callidactylus – Carapace posteriorly with 0–5 spines or protuberances��������������������������������������������������������������������������������������������������������� 45 45. Carapace posterolateral and posterior margins together with 4 spines or protuberances (submedian spines sometimes fused into single plate) (Fig. 14.63u)�����������������������������������������������������������������������������������������������������������������Ilia – Carapace posterolateral and posterior margins together with 0, 2 or 5 spines or protuberances������������������������������������ 46 46. Carapace posteriorly with 2 prominent spines. Male cheliped merus longer than carapace (Fig. 14.65h)���������� Pseudomyra – Carapace with 5 posterior spines or protuberances (1 median, 2 pairs around posterior and posterolateral margins), rarely posteriorly unarmed. Male cheliped merus usually shorter than carapace (Fig. 14.65k)���������Randallia 47. Carapace frontal margin strongly produced anteriorly beyond hepatic margin; dorsal surface distinctly irregular and uneven, with cardiac and epibranchial bosses (Fig. 14.63n)�������������������������������������������������������� Ebaliopsis – Carapace frontal margin weakly produced beyond hepatic margin; dorsal surface with even contours, without bosses�������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 48 48. Male pleonites 3–5 fused������������������������������������������������������������������������������������������������������������������������������������������������ Acanthilia – Male pleonites 3–6 fused�������������������������������������������������������������������������������������������������������������������������������������������������������������� 49 49. Carapace lateral margin lined with closely-spaced granules; subhepatic tubercle present (Fig. 14.64k). Female maxilliped 3 with longitudinal row of upright setae��������������������������������������������������������������������������������������� Myra – Carapace lateral margin smooth; subhepatic tubercle absent. Female maxilliped 3 without longitudinal row of upright setae�����������������������������������������������������������������������������������������������������������������������������������������������������������������Myrine 50. Carapace posterior margin with single blunt median tubercle (Fig. 14.64f)���������������������������������������������������������� Leucosilia – Carapace posterior margin with 2 or more spines or lobes�����������������������������������������������������������������������������������������������������51 51. Anterior margin of inhalent branchial channel forming suborbital margin, margin curved, lateral corner forming blunt angular point���������������������������������������������������������������������������������������������������������������������������������������������������52 – Anterior margin of inhalent branchial channel abutting or separated from suborbital margin by distinct groove or larger space�������������������������������������������������������������������������������������������������������������������������������������������������������������� 54 52. Carapace posterior margin with 2 pairs of teeth and higher median tooth (Fig. 14.64l). Male pleonites 3–6 fused (Fig. 14.68i)������������������������������������������������������������������������������������������������������������������������Myropsis – Carapace posterior margin with pair of teeth and higher median tooth. Male pleonites 3–5 fused��������������������������������53 53. Cheliped fingers slender but distinctly tapering throughout length, opening in almost horizontal plane; merus of males as shorter than carapace (excluding posteromedian spine) (Fig. 14.65b)�����������������������������Persephona – Cheliped fingers styliform, tapering little throughout length, opening in vertical plane; merus of males as long or longer than carapace (excluding posteromedian spine) (Fig. 14.64a)���������������������������������������������������Iliacantha 54. Carapace anterior margin of inhalent branchial channel appressed to suborbital margin such that suborbital spine or lobe positioned immediately behind plane of inhalent branchial margin (when examined in anteroventral view) (Fig. 14.66n, p)�����������������������������������������������������������������������������������������������������������������55

14 – Brachyura – crabs

– 55.



56. – 57. – 58. – 59. – 60. – 61. – 62. – 63. – 64. – 65. –

557

Carapace anterior margin of inhalent branchial channel separated from suborbital margin by transverse groove or larger space (Fig. 14.66a, j)������������������������������������������������������������������������������������������������������������������������������������ 56 Carapace markedly wider than long, spindle-shaped, with prominent conical lateral process (Fig. 14.64b). Anterior margin of inhalent branchial channel formed of 2 adjacent subequal convex lobes; suborbital lobe a blunt process immediately behind plane of convex anterior lobes (when examined in anteroventral view) (Fig. 14.66p)������������������������������������������������������������������������������������������������������������������������������������������Ixa Carapace, excluding lateral spines little wider than long or longer than wide (Fig. 14.63e, f). Anterior margin of inhalent branchial channel with angular lobe or spine at lateral and mesial corner, separated by straight or weakly curved margin, suborbital lobe/spine visible behind plane of inhalent margin (when examined in anteroventral view) (Fig. 14.66n)�����������������������������������������������������������������������������������������������Arcania Carapace posterior margin with 3 blunt spines or rounded lobes in horizontal row (occasionally fused into single wide flange-like plate), with or without median lobe or spine on higher plane (intestinal spine) (Fig. 14.65l, q)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������57 Carapace posterior margin with 2 rounded lobes in horizontal row, with or without median lobe or spine on higher plane (intestinal spine) (Fig. 14.63p)������������������������������������������������������������������������������������������������������������������� 58 Carapace surface and margins finely and evenly granulate (Fig. 14.65q). Male pleonites 2–6 fused. Gonopod 1 with slender, hooked tip (Fig. 14.69t)������������������������������������������������������������������������������������������������������� Tokoyo Carapace surface and margins covered with large, sharp to blunt tubercles (Fig. 14.65l). Male pleonites 3–6 fused. Gonopod 1 with flared, triangular or trilobate tip (Fig. 14.69q)����������������������������������������������������������������� Raylilia Gonopod 1 straight, with short preapical process, apex blunt (Fig. 14.69s). Male telson triangular (Fig. 14.68u)���������������������������������������������������������������������������������������������������������������������������������������������������������������������� Tanaoa Gonopod 1 with long attenuate distal process at right angles to main axis (Fig. 14.69u). Male telson lingulate (Fig. 14.68v)���������������������������������������������������������������������������������������������������������������������������������������������������������������59 Carapace surface smooth of finely granular, with regions indistinct. Gonopod 1 with attenuate distal process sharply angled (Fig. 14.69u)��������������������������������������������������������������������������������������������������������������������������������� Toru Carapace surface covered by large, rounded, pustulose tubercles, regions distinct (Fig. 14.63p). Gonopod 1 with attenuate distal process gently angled (Fig. 14.69h)��������������������������������������������������������������������������������������������Galilia Carapace anterolateral margin straight, oblique, meeting posterolateral margins at right angle (Fig. 14.64g). Maxilliped 3 exopod distally swollen (Fig. 14.66t)������������������������������������������������������������������� Lissomorpha Carapace anterolateral margins convex, rounded, meeting posterolateral margin obtusely or with even curve, even if spine or tubercle present at junction. Maxilliped 3 exopod flat���������������������������������������������������������������61 Carapace covered with large rounded tubercles of different sizes (Figs 14.64m, 14.65e, t)������������������������������������������������62 Carapace dorsally smooth or evenly and finely granulate, sometimes with few rows of prominent granules�������������� 64 Maxilliped 3 exopod distinctly wider than ischium (Fig. 14.66u). Cheliped fingers distinctly longer than dorsal margin of palm (Fig. 14.64m). Male pleonites 3–6 fused but somites 5 and 6 demarcated by groove (Fig. 14.68j)������������������������������������������������������������������������������������������������������������������������������������������������������� Neparilia Maxilliped 3 exopod little wider than ischium. Cheliped fingers as long as or shorter than dorsal margin of palm. Male pleonites 3–6 indistinguishably fused������������������������������������������������������������������������������������������������������������� 63 Carapace dorsal surface with posteromedian row of 2–4 prominent tubercles, anterior 1–3 upright, posteriormost directed posteriorly (Fig. 14.65e). Anterior margin of inhalent branchial channel with 3 teeth or lobes�������������������������������������������������������������������������������������������������������������������������������������������������������������������� Phlyxia Carapace dorsal surface without posteromedian row of prominent tubercles (Fig. 14.65t). Anterior margin of inhalent branchial channel bilobed��������������������������������������������������������������������������������������� Urashima Cheliped slender, strongly elongate, male palm at least 5 times as long as wide (Fig. 14.65a)�����������������������������������Parilia Cheliped stout, male palm no more than 3 times as long as wide����������������������������������������������������������������������������������������� 65 Carapace front with median blunt lobe produced beyond epistome and anterior margin of inhalant branchial channel (Fig. 14.65i)���������������������������������������������������������������������������������������������������������������������������Pseudophilyra Carapace front straight, concave or with small tooth, not produced beyond epistome and anterior margin of inhalant branchial channel������������������������������������������������������������������������������������������������������������������������������������������������ 66

558

Marine Decapod Crustacea

Fig. 14.62.  Leucosiidae, Cryptocneminae. a, b, Cryptocnemus kavieng Galil & Ng, 2020, dorsal, ventral; c, Onychomorpha lamelligera Stimpson, 1858. d, e, Paranursia abbreviata (Bell, 1855), dorsal, ventral); Gonopod 1: f–h, Cryptocnemus; i, Paranursia.

66. Carapace front with medial incision; supraorbital margin with 2 fissures (Fig. 14.66q). Gonopod 2 filiform, as long as gonopod 1 (Fig. 14.69f)�������������������������������������������������������������������������������������������������������������������������������Bellidilia – Carapace front concave or with short medial tooth; supraorbital margin with 1 fissure, rarely without (Fig. 14.66r). Gonopod 2 with scoop-like apex, shorter than gonopod 1 (Fig. 14.69a)������������������������������������������������� 67 67. Male pleonites 1, 2 articulating; pleonites 3–5 or 3–6 fused (Fig. 14.68a, c)������������������������������������������������������������������������ 68 – Male pleonites 2–6 fused (Fig. 14.68h, q)���������������������������������������������������������������������������������������������������������������������������������� 73 68. Carapace longer than wide with tooth at junction between posterolateral and posterior margins (Fig. 14.64v)������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 69 – Carapace as wide as or wider than long, without lobiform tooth at junction between posterolateral and posterior margins��������������������������������������������������������������������������������������������������������������������������������������������������������������������� 69 69. Carapace length 1.2–1.3 times width; surface finely or indistinctly punctate; branchial surfaces each with 1 oblique row of granules. Gonopod 1 distal quarter sharply bent backwards from longitudinal axis; with curved, tapering subdistal process (Fig. 14.69n)�������������������������������������������������������������������������������������������������������� Ovilyra – Carapace length 1.1 times width; surface prominently punctate; branchial surfaces each with 2 oblique rows of granules. Gonopod 1 almost straight, without distal bend����������������������������������������������������������������������� Alcolyra 70. Male pleonites 3–5 fused (Fig. 14.68a). Gonopod 1 distally crook-shaped (Fig. 14.69a)����������������������������������������Afrophila – Male pleonites 3–6 fused. Gonopod 1 otherwise����������������������������������������������������������������������������������������������������������������������71 71. Male pleonites 3–6 constricted between fused pleonites (Fig. 14.68c). Gonopod 1 coiled twice (Fig. 14.69c)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Atlantolocia – Male pleonites 3–6 tapering, not constricted between fused pleonites (Fig. 14.68o). Gonopod 1 uncoiled������������������� 72 72. Carapace globose, pear-shaped. Epistome and lower hepatic margin not projecting. Maxilliped 3 exopod semicircular. Male pleonites 1, 2 narrow. Gonopod 1 flattened, apical process bifurcate, cornute (Fig. 14.69o)����������������������������������������������������������������������������������������������������������������������������������������������������������������������� Philyra – Carapace discoid. Epistome and lower hepatic margin projecting (Fig. 14.65m). Maxilliped 3 exopod subtriangular (Fig. 14.66w). Male pleonites 1, 2 laterally swollen. Gonopod 1 with apical process tubular (Fig. 14.69r)���������������������������������������������������������������������������������������������������������������������������������������������������������������������� Ryphila 73. Male pleonite 1 two-thirds width of pleonite 2. Gonopod 1 coiled 3 times, apical process with setose muff (Fig. 14.69d)������������������������������������������������������������������������������������������������������������������������������������������������������������� Atlantophila

14 – Brachyura – crabs

– 74. – 75. –

559

Male pleonite 1 almost as wide as pleonite 2 (Fig. 14.68h, q). Gonopod 1 not coiled (Fig. 14.69j, i, p)����������������������������74 Male pleonites 2–6 with denticle (Fig. 14.68h)���������������������������������������������������������������������������������������������������������������� Lyphira Male pleonites 2–6 without denticle (Fig. 14.68q)���������������������������������������������������������������������������������������������������������������������75 Gonopod 1 apical process long, tubular (Fig. 14.69p). Cheliped merus shorter than carapace in male, fixed finger without fringe of setae (Fig. 14.65j)��������������������������������������������������������������������������������������������������������������������Pyrhila Gonopod 1 apical process minute (Fig. 14.69i). Cheliped merus longer than carapace in male, fixed finger with fringe of setae on inner margin (Fig. 14.63s)����������������������������������������������������������������������������������������������������� Hiplyra

Subfamily Cryptocneminae Stimpson, 1907 Diagnosis. Carapace lateral and posterior margins expanded, with thin sharp edge, forming continuous laminar rim overhanging proximal part of pereopods and pleon; dorsal surface smooth, appearing polished, unornamented or at most with low, granular median and oblique epibranchial ridges; thoracic sinus absent. Endostome reaching beyond anterior margin of inhalant branchial channel. Cryptocnemus Stimpson, 1858 Diagnosis. Carapace wider than long; dorsal surface smooth, appearing polished, unornamented or with low, granular median and oblique epibranchial ridges; frontal region weakly produced, strongly produced or as two horn-like lobes (if carapace little wider than long and lateral margin with concavity behind midlength, then front forming two horn-like lobes); margins expanded, with thin sharp edge, forming continuous laminar rim overhanging proximal part of pereopods and pleon. Cheliped palm with inner lower surface glabrous. Male pleonites 3–6 fused. Gonopod 1 with 1 or 2 distal points or lobes. Maximum cl. 8 mm (Pl. 46a). Intertidal–slope (0–275 m). Temperate Northern E Pacific, Western and Central Indo-Pacific, Temperate Australia. 22 species (Galil and Ng 2020: commentary on polyphyly, list of species)

Onychomorpha Stimpson, 1858 Diagnosis. Carapace longer than wide, subtriangular, with convex lateral and posterior margins; dorsal surface smooth, with broad longitudinal ridge; lateral and posterior margins convex, meeting at posterolateral corner, laminar, with sharp edge. Cheliped palm with inner lower surface densely setose. Male pleonites 2–6 fused. Maximum cl. 6 mm. Subtidal–shelf (6–27 m). Western and Central Indo-Pacific. 1 species (Alcock 1896: description; Chen and Sun 2002: figures; Stimpson 1907: redescription; Walker 1887: figure).

Paranursia Serène & Soh, 1976 Diagnosis. Carapace subcircular; dorsal surface with prominent medial and paired oblique epibranchial ridges; frontal region triangular; lateral margin laminar, with 3 broad lobes, beaded. Cheliped palm with inner lower surface glabrous. Male

pleonites 3–6 fused. Gonopod 1 with 3 distal points. Maximum cl. 10 mm. Intertidal, subtidal (0–14 m). Western and Central IndoPacific, Temperate Australia. 1 species (Lee and Ng 2014; Serène and Soh 1976).

Subfamily Ebaliinae Stimpson, 1871 Diagnosis. Carapace variously globose or angular; lateral margins unexpanded, or if expanded, edge blunt, not thinsharp, often tubercular or beaded, rarely forming continuous rim overhanging branchial and posterior margins; dorsal surface granular or punctate, often sculptured; median and oblique epibranchial ridges, if present, usually prominent; thoracic sinus absent. Implicit generic attributes. Frontal region produced, bilobed. Female pleonites 1, 2 free, 3–6 fused. Telson tapering. Gonopod 2 shorter than gonopod 1. Acanthilia Galil, 2000 Diagnosis. Carapace subcircular, globose; dorsal surface with contours largely even, without bosses; anterolateral margin with broad lobe; posterior margin with 3 teeth, medial one sharp. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 3 teeth, contiguous with lower orbital margin. Cheliped fingers half as long as palm. Male pleonites 3–5 fused. Maximum cl. 15 mm (Pl. 46b). Subtidal–slope (10–329 m). Temperate Northern and Tropical W Atlantic. 1 species (Galil 2000).

Afrophila Galil, 2009 Diagnosis. Carapace globose, as long as wide; dorsal surface smooth, finely or coarsely granulate; frontal region upcurved, weakly bilobed, not exceeding epistome; margins beaded, hepatic margin produced; posterior margin rounded. Endostome reaching level of inhalant branchial channel. Supraorbital margin with 1 fissure. Cheliped fingers about as long as palm. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 1, 2 articulating, first anvil-shaped, second narrow; pleonites 3–5 fused. Gonopod 1 slender, sinuous, distally crook-shaped. Gonopod 2 with scoop-like apex, shorter than gonopod 1. Maximum cl. 20 mm. Intertidal, subtidal (0–50 m). Temperate Southern Africa. 1 species (Galil 2009).

560

Marine Decapod Crustacea

Fig. 14.63.  Leucosiidae, Ebaliinae. a, Acanthilia intermedia (Miers, 1886); b, Alox naispela Galil & Ng, 2015; c, Alox rugosum (Stimpson, 1858); d, Ancylodactyla elongata (Zarenkov, 1969); e, Arcania fungilifera Galil, 2001; f, Arcania heptacantha (De Man, 1907); g, Atlantotlos rhombifer Doflein, 1904; h, Bellidilia laevis (Bell, 1855); i, Callidactylus asper Stimpson, 1871; j, Cateios frontalis (Miers, 1884); k, Coralliocryptus caementa Komai & Ng, 2012; l, Dolos petraeus (A. Milne-Edwards, 1874); m, Ebalia dentifrons Miers, 1886; n, Ebaliopsis erosa (A. Milne-Edwards, 1873); o, Favus granulatus Lanchester, 1900; p, Galilia petricola Komai & Tsuchida, 2014; q, Heterolithadia fallax (Henderson, 1893); r, Heteronucia perlata (Sakai, 1963); s, Hiplyra longimana (A. Milne Edwards, 1874); t, Ihleus lanatus (Alcock, 1896); u, Ilia spinosa.

Alcolyra Trivedi, Mitra & Ng, 2022 Diagnosis. Carapace suborbicular, 1.1 times longer than wide; dorsal surface finely granulate, strongly punctate; frontal region upcurved, short median tooth, not exceeding epistome;

margins granular; posterior half of anterolateral margin with 2 teeth, posteriormost at junction with posterolateral margin; posterior margin with blunt, triangular tooth separating posterolateral margin. Endostome reaching level of inhalant branchial channel. Cheliped fingers about as long as palm.

14 – Brachyura – crabs

561

Fig. 14.64.  Leucosiidae, Ebaliinae. a, Iliacantha intermedia Miers, 1886; b, Ixa holthuisi Tirmizi, 1970; c, Kabutos durandi (Serène, 1955); d, e, Leucisca halimedophila Galil & Innocenti, 2019; f, Leucosilia jurinii (de Saussure, 1853); g, Lissomorpha haswelli Ward, 1936; h, Lithadia conica (Coelho, 1973); i, Merocryptoides frontalis Sakai, 1963; j, Merocryptus lambriformis A. Milne-Edwards, 1873; k, Myra mammillaris Bell, 1855; l, Myropsis quinquespinosa Stimpson, 1871; m, Neparilia tuberculata Sakai, 1961; n, Nobiliella jousseaumei (Nobili, 1905); o, Nucia speciosa Dana, 1852; p, Nuciops modesta (Ihle, 1918); q, Nursia phylloides Ihle, 1918; r, Nursilia dentata Bell, 1855; s, Oreophorus fenestrus Tan & Ng, 1995; t, Oreotlos etor Tan & Richer de Forges, 1993; u, Orientotlos iishiba Sakai 1980; v, Ovilyra fuliginosa (Targioni-Tozzetti, 1877).

562

Marine Decapod Crustacea

Fig. 14.65.  Leucosiidae, Ebaliinae. a, Parilia major Sakai, 1961; b, Persephona orbicularis Bell, 1855; c, d, Philyra globus (Fabricius, 1775); e, Phlyxia lambriformis Bell, 1855; f, Praebebalia fungifer Galil, 2015; g, Praosia punctata Tan & Ng, 1993; h, Pseudomyra mbizi Capart, 1951; i, Pseudophilyra punctulata Chen & Ng, 2003; j, Pyrhila carinata (Bell, 1855); k, Randallia americana (Rathbun, 1894); l, Raylilia coniculifera Galil, 2001; m, Ryphila cancellus (Herbst, 1783); n, o, Speloeophorus nodosus (Bell, 1855) with posterior view; p, Tlos muriger Adams & White, 1848; q, Tokoyo eburnea (Alcock, 1896); r, s, Uhlias ellipticus Stimpson, 1871; t, Urashima pustuloides Sakai, 1961.

Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shieldlike. Male pleonites 1, 2 articulating, narrow; fused pleonites 3–5 tapering, without constrictions. Gonopod 1 slender, elongate, almost straight; apical process spatuliform. Gonopod 2

apex tapering, sharp, shorter than gonopod 1. Maximum cl. 15 mm. Estuarine, subtidal (1.5–3 m). Western Indo-Pacific. 1 species (Trivedi et al. 2022).

14 – Brachyura – crabs

563

Fig. 14.66.  Leucosiidae, Ebaliinae. Epistome, inhalent channel, orbit (maxilliped 3 removed): a, Bellidilia; b, Ebalia; c, Alox; d, Oreophorus. Orbit, antennular fossa, antenna: e, Ebalia; f, Oreophorus. Anteroventral carapace, orbit, antennular fossae, maxillipeds 3: g, Arcania; h Leucisca; i, Myra; j, Parilia. Antennular bases, fossae: k, Alox; l, Oreophorus. Anterior inhalent channel margin (and suborbital lobe): m, Acanthilia; n, Arcania; o, Iliacantha; p Ixa. Front, orbits, dorsal view (fissures arrowed): q, Bellidilia; r, Hiplyra. Maxilliped 3: s, Kabutos; t, Lissomorpha; u, Neparilia; v, Oreotlos; w, Ryphila. Cheliped: x, Coralliocryptus; y, Nursilia; z, Oreophorus.

Fig. 14.67.  Leucosiidae, Ebaliinae. Female pleon: a, Coralliocryptus; b, Ebalia; c, Heteronucia; d, Ihleus; e, Kabutos; f, Nobiliella; g, Nursia.

Alox Tan & Ng, 1995 Diagnosis. Carapace suboval, 1.2–1.7 times as wide as long; dorsal surface rugose, with numerous mushroom-like granules and excavations, with granule-filled groove parallel to margin, with pair of postorbital prominences; frontal region narrow, moderately upturned; anterolateral and posterolateral margins expanded laterally, with or without rim. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closedoff from antennular fossa by immovable antennal basal article; antennular basis occupying entire fossa. Cheliped fingers ~1.5 times as long as palm. Male pleon covered with large mushroomlike granules; pleonites 3–5 fused. Gonopod 1 1.6–4.3 times as long as gonopod 2. Maximum cl. 14 mm (Pl. 46c).

Intertidal–shelf (0–200 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 14 species (Tan and Ng 1995: key to 8 species).

Ancylodactyla Galil, 2004 Diagnosis. Carapace subhexagonal, globose; dorsal surface with regions indistinct except for intestinal prominence; anterior and posterolateral margins separated by broad curve; posterior margin straight. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially open, not closed-off from antennular fossa, antennal basal article movable. Cheliped merus 1.5 times as long as carapace; fingers about half as long as

564

Marine Decapod Crustacea

Fig. 14.68.  Leucosiidae, Ebaliinae. Male pleon (pleonites 1, 2 not always shown): a, Afrophila; b, Ancylodactyla; c, Atlantolocia; d, Cateios; e, Dolos; f, Ihleus; g, Leucosilia; h, Lyphira; i, Myropsis; j, Neparilia; k, Nursia; l, Oreotlos; m, Parilia; n, Persephona; o, Philyra; p, Praosia; q, Pyrhila; r, Randallia; s, Speloeophoroides; t, Speloeophorus; u, Tanaoa; v, Toru; w, Tlos.

Fig. 14.69.  Leucosiidae, Ebaliinae. Gonopod 1: a, Afrophila; b, Alox; c, Atlantolocia; d, Atlantophila; e, Atlantotlos; f, Bellidilia; g, Cateios; h, Galilia; i, Hiplyra; j, Lyphira; k, Merocryptus; l, Oreophorus; m, Oreotlos; n, Ovilyra; o, Philyra; p, Pyrhila; q, Raylilia; r, Ryphila; s, Tanaoa; t, Tokoyo; u, Toru. a, b, f, l with gonopod 2.

14 – Brachyura – crabs

palm. Pereopods 1–4 smooth, slender. Male pleonites 3–6 fused. Maximum cl. 9.3 mm (Pl. 46d). Shelf, slope (156–618 m). Central Indo-Pacific, Tropical Eastern Pacific. 3 species (Galil 2004: key to species).

Arcania Leach, 1817 Diagnosis. Carapace subhexagonal, or globose, or pyriform (little wider than long or longer than wide (excluding lateral spines)); dorsal surface densely covered with granules, tubercles or spines; posterior margin with pair of teeth or lobes and higher median tooth. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with angular lobe or spine at lateral and mesial corner, separated by straight or weakly curved margin, suborbital lobe/spine visible behind plane of inhalent margin (in anteroventral view). Maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped elongate, palm thin, fingers about as long as palm, cutting edges with uneven short triangular teeth. Female pleonites 1–3 free, 4–6 fused, swollen. Male pleonites 3–5 fused; pleonite 6 without denticle. Maximum cl. 34 mm (Pl. 46f, g). Intertidal–slope (0–366 m). Temperate Northern W Pacific, Indo-West Pacific, Temperate Southern Africa, Temperate ­Australia. 22 species (Galil 2001c: key to 14 species, includes Raylilia; Naruse 2014: species resolved).

Atlantolocia Galil, 2009 Diagnosis. Carapace globose, subpentagonal; dorsal surface smooth, finely or coarsely granulate; frontal region upcurved, weakly bilobed, not exceeding epistome; margins beaded, hepatic margin produced; posterior margin rounded. Endostome reaching level of inhalant branchial channel. Supraorbital margin with 1 fissure. Cheliped fingers longer than palm. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 1, 2 articulating; fused pleonites 3–6 with constrictions between pleonites. Gonopod 1 elongate, coiled twice. Gonopod 2 with scooplike apex, shorter than gonopod 1. Maximum cl. 15 mm. Subtidal, shelf (4–86 m). Temperate Northern and Tropical E Atlantic. 1 species (Galil 2009).

Atlantophila Galil, 2009 Diagnosis. Carapace globose, as long as wide; dorsal surface smooth, finely or coarsely granulate; frontal region upcurved, weakly bilobed, not exceeding epistome; lateral margin beaded; posterior margin rounded. Endostome reaching level of inhalant branchial channel. Supraorbital margin with 1 fissure. Maxilliped 3 exopod flat, as wide as endopod ischium at midlength. Cheliped fingers shorter than palm. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 1 two-thirds width of pleonite 2; pleonites 2–6 fused, with subdistal denticle. Gonopod 1 elongate, coiled 3 times, apical process with setose muff. Gonopod 2 with scoop-like apex, shorter than gonopod 1. Maximum cl. 6.3 mm. Subtidal, shelf (4–25 m). Tropical E Atlantic. 1 species (Galil 2009).

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Atlantotlos Doflein, 1904 Diagnosis. Carapace subtriangular, 1.0–1.3 times as wide as long; dorsal surface eroded, sculptured, thick, irregular, with median and diagonal epibranchial ridges; anterolateral margins irregular but produced anteriorly to form an almost right angle giving carapace approximately rectangular outline. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially open, not closed-off from antennular fossa, antennal basal article movable. Cheliped fingers about as long as palm, cutting edges dentate or crenulated. Female pleonites 1–3 free, 4–6 fused. Male pleonites 3–6 fused. Gonopod 1 distally bulbous with slender apical process. Maximum cl. 12 mm. Subtidal–shelf (33–115 m). Tropical E Atlantic. 1 species (Capart 1951: figure; Manning and Holthuis 1981).

Bellidilia Kinahan, 1856 Diagnosis. Carapace subcircular; dorsal surface smooth; frontal region with 4 teeth separated by medial notch, not exceeding epistome; convex, with granules; posterior margin with pair of teeth or lobes and higher median tooth, or rounded. Endostome reaching level of inhalant branchial channel. Supraorbital margin with 2 fissures. Maxilliped 3 exopod flat, as wide as endopod ischium at midlength. Cheliped fingers longer than palm. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 3–6 fused. Gonopod 2 filiform, as long as gonopod 1. Maximum cl. 19 mm (Pl. 46h). Intertidal–slope (0–512 m). Temperate Australasia (Australia, New Zealand). 3 species (Tan 1995).

Callidactylus Stimpson, 1871 Diagnosis. Carapace subglobular, subhepatic margin not visible in dorsal view; dorsal surface covered with simple granules and short, capitate spinules; posterolateral and posterior margins together with 7 spines or protuberances (2 pairs around posterolateral and posterior margins; 3 in posterior midline). Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 3 teeth. Maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped elongate, palm swollen, fingers at least 1.5 times as long as palm, cutting edges with uneven short perpendicular teeth. Male pleonites 3–5 fused. Maximum cl. 18 mm (Pl. 46i). Shelf, slope (27–393 m). Tropical W Atlantic. 1 species (Rathbun 1937; Williams 1984).

Cateios Tan & Ng, 1995 Diagnosis. Carapace subtriangular, 1.5 times as wide as long; dorsal surface granular, with narrow median ridge, branchial region swollen; frontal region narrow, strongly produced, weakly upturned; anterolateral and posterolateral margins expanded, scalloped with large pointed granules. Endostome reaching level of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article; antennular basis occupying lower half of fossa. Cheliped fingers twice as long as palm. Male

566

Marine Decapod Crustacea

pleon coarsely granular; pleonites 3–5 fused, with short transverse ridge; telson arrow-shaped. Gonopod 1 twice as long as gonopod 2, stout, with densely setose distal half. Maximum cl. 8 mm. Shelf (24–64 m). Temperate Northern W Pacific, Central IndoPacific. 1 species (Tan and Ng 1995).

Coralliocryptus Komai & Ng, 2012 Diagnosis. Carapace subhexagonal, 1.6 times as wide as long; dorsal surface covered with tubercles, without ridges; epibranchial margin strongly expanded laterally, divided into 2 lobes. Endostome reaching beyond anterior margin of inhalant branchial channel. Cheliped fingers 1.6 times as long as palm, slender, curved in dorsal view, cutting edges with unequal perpendicular spines. Female pleonites 1–6 and telson all free, much longer than wide. Male pleonites 3–5 fused. Maximum cl. 5.4 mm. Subtidal, shelf (13–30 m). Western and Central Indo-Pacific. 1 species (Komai and Ng 2012).

Dolos Tan & Richer de Forges, 1993 Diagnosis. Carapace oval, 1.4 times as wide as long; dorsal surface finely granular, with indistinct median ridge, branchial region concave, with region of mushroom-shaped tubercles; frontal region produced, bilobed; anterolateral and posterolateral margins expanded, thin, plate-like, upturned, with 3 closed sutures. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article; antennular basis occupying entire fossa. Cheliped fingers about as long as palm. Male pleon finely granular; pleonites 3–5 fused. Gonopod 1 2.0–2.4 times as long as gonopod 2. Maximum cl. 7 mm. Subtidal–shelf (3–74 m). Central Indo-Pacific. 1 species (Tan and Ng 1995: rediagnosis).

Ebalia Leach, 1817 Diagnosis. Carapace oval or irregularly polygonal, highest point in midline; dorsal surface ranging from finely granular to coarsely tuberculate, lobate or spinous, but without substantial areas of deep, irregular erosions and cavities; not expanded, or if expanded, not thin-sharp, edge blunt; posterior margin rounded, or with 2 or 3 teeth. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially open, not closed-off from antennular fossa, antennal basal article movable. Cheliped merus in adults about as long as or shorter than carapace, propodus stout; fingers less than 1.3 times as long as palm, cutting edges dentate or crenulated. Pereopods 1–4 coarsely tuberculate. Female pleonites 1–3 free, 4–6 fused, or 1, 2 free, 3–6 fused. Male pleonites 3–5 fused. Maximum cl. 12 mm (Pl. 46j). Intertidal–bathyal (0–2983 m). Cosmopolitan except polar seas. 68 species (Miers 1886: grouped species; Rathbun 1937: key to 7 American species). Ebalia has been a catch-all genus since its inception but its taxonomy is unstable (Komai and Ng 2012; Naruse and Ng 2006)

Ebaliopsis Ihle, 1918 Diagnosis. Carapace longer than wide; dorsal surface distinctly irregular and uneven, with low cardiac and epibranchial bosses, granulate; frontal region strongly produced beyond hepatic margin; hepatic margin obtuse, lateral margin convex, with posterolateral blunt tooth; posterior margin with pair of teeth or lobes and higher median tooth. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 3 teeth or lobes. Maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped fingers shorter than palm; male merus shorter than carapace. Female pleonites 1–3 free, 4–6 fused, swollen. Male pleonites 3–5 fused. Maximum cl. 12 mm. Intertidal, subtidal (0–20 m). Indo-West Pacific, Temperate Australia. 2 species (Komatsu et  al. 2004: figures; Naruse et  al. 2017: discussed 2 species, figures).

Favus Lanchester, 1900 Diagnosis. Carapace twice as wide as long (including produced intestinal region); dorsal surface honey-combed, without ridges; frontal region not produced; anterior and posterolateral margins parallel, lateral margin evenly convex. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article. Cheliped fingers shorter than palm. Maximum cl. 7.5 mm. Intertidal, subtidal. Central Indo-Pacific (Singapore). 1 species (Lanchester 1900).

Galilia Ng & Richer de Forges, 2007 Diagnosis. Carapace subglobular; dorsal surface covered by large, rounded, pustulose tubercles, regions distinct; posterior margin with 2 rounded lobes in horizontal row, with low median lobe at higher plane (intestinal spine). Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 2 lobes, separated from suborbital margin by transverse groove or larger space. Orbit mesially closed-off from antennular fossa by slightly movable antennal basal article. Maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped fingers as long as palm, curved; male merus two-thirds carapace length. Pereopods 1–4 coarsely tuberculate. Female pleonites 1–3 free, 4–6 fused, swollen. Male pleonites 3–6 fused; telson lingulate. Gonopod 1 slender, distal third bent. Maximum cl. 31 mm (Pl. 46k). Slope (377–595 m; seamounts). Central Indo-Pacific. 2 species (Komai and Tsuchida 2014; Ng and Richer de Forges 2007; Shih et al. 2015).

Heterolithadia Alcock, 1896 Diagnosis. Carapace subglobular; dorsal surface covered with mushroom-shaped tubercles, with broad median ridge with smooth paired gastric and hepatic bosses, and smooth median cardiac and intestinal bosses; pterygostomial margin ending in

14 – Brachyura – crabs

lateral tubercle, visible in dorsal view; posterior margin narrow, bilobed. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 3 teeth or lobes. Maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped elongate, palm swollen, fingers at least 1.5 times as long as palm, cutting edges with uneven short erect teeth; merus covered with mushroom-shaped tubercles. Thoracic sternum covered with mushroom-shaped tubercles. Male pleonites 3–5 fused, with prominent tubercle. Maximum cl. 13 mm (Pl. 46l). Subtidal–slope (2–382 m). Western and Central Indo-Pacific. 1 species (Galil and Ng 2015: distribution, figures; Henderson 1893: figure; Tan 1996: rediagnosis). .

Heteronucia Alcock, 1896 Diagnosis. Carapace subcircular-polygonal in outline, globose; dorsal surface and margins coarsely tuberculate to multilobate or multispinous; lateral margin convex, tuberculate; posterior margin with 2 teeth, or rounded. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by slightly movable antennal basal article (slightly movable in H. laminata). Cheliped merus shorter than carapace length; fingers slender, longer than upper margin of palm. opening vertically or obliquely. Pereopods 1–4 coarsely tuberculate. Female pleonites 1–3 free, 4–6 fused. Male pleonites 3–5 fused. Maximum cl. 21 mm (Pl. 46m). Intertidal–shelf (0–220 m). Temperate Northern W Pacific, Indo-West Pacific, Temperate Southern Africa. 17 species (Komatsu and Takeda 2005: 2 species described; Tan 1996: discussion).

Hiplyra Galil, 2009 Diagnosis. Carapace subcircular, globose; dorsal surface smooth, finely or coarsely granulate; frontal region upcurved, with short medial tooth, not exceeding epistome; lateral margin beaded; posterior margin rounded. Endostome reaching level of inhalant branchial channel. Supraorbital margin with 1 fissure. Maxilliped 3 exopod flat, as wide as endopod ischium at midlength. Cheliped merus longer than carapace in male, fixed finger with fringe of setae. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 1 almost as wide as pleonite 2; pleonites 2–6 fused, without subdistal denticle. Gonopod 1 elongate, flattened, apical process minute. Gonopod 2 with scoop-like apex, shorter than gonopod 1. Maximum cl. 19 mm (Pl. 46n). Intertidal–slope (0–150 m). Western and Central IndoPacific, Temperate Southern Africa, Temperate Australia. 7 species (Galil 2009).

Ihleus Ovaere, 1989 Diagnosis. Carapace suboval, 1.2 times as long as wide; dorsal surface slightly pubescent, regions well defined, tubercular especially on cardiac and intestinal regions; lateral margin convex, with short teeth; posterior margin with 4 rounded teeth, or with 2 teeth plus medial intestinal tooth. Endostome reaching beyond

567

anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article. Cheliped fingers shorter than palm. Pereopods 1–4 smooth, stocky. Female pleonites 1–6 and telson all free, much longer than wide. Male pleonites 3–5 fused. Maximum cl. 10 mm. Subtidal–shelf (15–100 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 2 species (Ovaere 1989).

Ilia Leach, 1817 Diagnosis. Carapace subglobular; dorsal surface granular; subhepatic margin not visible in dorsal view; margins with 4 spines or protuberances (submedian spines sometimes fused into single plate) around posterolateral and posterior margins, median spine or protuberance absent. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 3 lobes, separated from suborbital margin by transverse groove. Maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped elongate, palm thin, fingers about as long as palm, cutting edges with uneven short triangular teeth. Male pleonites 3–5 fused. Maximum cl. 28 mm. Intertidal–shelf (0–162 m). Tropical E Atlantic, including Mediterranean. 2 species (Manning and Holthuis 1981: ecology, distribution; Zariquiey Alvarez 1968: rediagnosis).

Iliacantha Stimpson, 1871 Diagnosis. Carapace subcircular, globose; dorsal surface smooth; anterolateral margin with broad lobe; posterior margin with 3 teeth, medial one sharp. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 2 lobes, separated by narrow groove from lower orbital margin. Cheliped fingers about as long as palm, or fingers longer than palm. Male pleonites 3–5 fused. Maximum cl. 31 mm (Pl. 47a). Subtidal–slope (11–915 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 4 species (Rathbun 1937: key to 4 species with Acanthilia).

Ixa Leach, 1816 Diagnosis. Carapace markedly wider than long, spindle-shaped, with prominent conical lateral process; dorsal surface granular, or smooth; posterior margin with pair of teeth or lobes and higher median tooth. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 2 adjacent subequal convex lobes. Suborbital lobe a blunt process placed immediately behind plane of convex anterior lobes inhalent branchial channel (in anteroventral view). Maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped exceptionally slender, fingers much shorter than palm. Male pleonites 3–5 fused. Maximum cl. 31 mm (Pl. 47b). Subtidal–shelf (16–84 m). Temperate Northern Atlantic, Western and Central Indo-Pacific. 9 species (Tyndale-Biscoe and George 1962: 3 species compared). Ixa monodi Holthuis & Gottlieb, 1956 is an exotic species in the eastern Mediterranean (Holthuis and Gottlieb 1956).

568

Marine Decapod Crustacea

Kabutos Komatsu & Ng, 2011 Diagnosis. Carapace 1.3 times as wide as long; dorsal surface eroded, sculptured, thick, irregular, with median and diagonal epibranchial ridges; frontal region short, with pair of subparallel gastric carinae; anterolateral margins irregular but produced anteriorly to form an almost right angle giving carapace approximately rectangular outline. Endostome reaching level of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article; antennular basis occupying lower half of fossa. Maxilliped 3 exopod with angular tooth at outer proximal margin. Cheliped fingers 1.1–1.2 times as long as palm, cutting edges dentate or crenulated. Female pleonites 1–3 free, 4–6 fused. Male pleonites 3–5 fused. Maximum cl. 4.1 mm. Subtidal–shelf (2–60 m). Central Indo-Pacific. 1 species (Komatsu and Ng 2011).

Leucisca MacLeay, 1838 Diagnosis. Carapace transversely ovate to semicircular; dorsal surface finely and evenly granular; frontal region produced beyond orbits, truncate or concave; margins expanded laterally and posteriorly forming continuous laminar rim overhanging meri of pereopods 2–5, edges blunt, beaded; posterior margin broad, bilobed. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article. Cheliped fingers about as long as palm. Male pleonites 3–6 fused. Maximum cl. 9 mm. Intertidal–subtidal (0–1 m). Western Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 4 species (Barnard 1950; Galil and Innocenti 2019).

Leucosilia Bell, 1855 Diagnosis. Carapace globose, as long as wide; dorsal surface granular, with low hepatic bosses; posterior margin with short blunt tooth. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with no more than 2 teeth or lobes. Maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped fingers slender, about as long as palm. Male pleonites 3–5 fused; pleonite 6 with spine overlapping pleonite 5. Maximum cl. 20 mm. Intertidal, subtidal (0–17 m). Tropical Eastern Pacific. 1 species (Rathbun 1937).

Lissomorpha Ward, 1933 Diagnosis. Carapace wider than long; dorsal surface smooth, with smooth longitudinal ridge; frontal region strongly produced; anterolateral margin straight, oblique, meeting posterolateral margins at right angle. Endostome reaching level of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article. Maxilliped 3 exopod as wide as or wider than endopod ischium at midlength, distally

swollen. Cheliped fingers shorter than palm. Female pleonites 2–6 fused. Maximum cl. 8 mm. Subtidal, shelf. Central Indo-Pacific (NE Australia). 1 species (Ward 1933).

Lithadia Bell, 1855 Diagnosis. Carapace subhexagonal-ovoid, highest point on branchial regions; dorsal surface with substantial areas of deep, irregular or convoluted excavations, often with deep pits and erosions; hepatic region depressed; lateral margin with 1 or 2 branchial lobes; posterior margin with 2 cardiac lobes. Endostome reaching beyond anterior margin of inhalant branchial channel. Cheliped shorter than palm; merus in adults about as long as or shorter than carapace, propodus stout. Female pleonites 1–3 free, 4–6 fused. Male pleonites 3–5 fused. Maximum cl. 18 mm (Pl. 47c). Subtidal, shelf (4–210 m). Tropical W Atlantic, Tropical Eastern Pacific. 7 species (de Melo 1996: key to 4 Brazilian species; Rathbun 1937: key to 3 species).

Lyphira Galil, 2009 Diagnosis. Carapace subcircular, globose; dorsal surface smooth, finely or coarsely granulate; frontal region upcurved, weakly bilobed, not exceeding epistome; lateral margin beaded; posterior margin rounded. Endostome reaching level of inhalant branchial channel, anterior margin forming lower margin of orbit. Supraorbital margin with 1 fissure. Maxilliped 3 exopod flat, as wide as endopod ischium at midlength. Cheliped fingers about as long as palm. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 1 almost as wide as pleonite 2; pleonites 2–6 fused, with subdistal denticle. Gonopod 1 elongate, flattened, apical process cornute. Gonopod 2 with scoop-like apex, shorter than gonopod 1. Maximum cl. 20 mm (Pl. 47d). Subtidal, shelf (1–108 m). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Southern Africa. 5 species (Galil 2009).

Merocryptoides Sakai, 1963 Diagnosis. Carapace 1.0–1.2 times as wide as long; dorsal surface eroded, sculptured, with thick, irregular median and diagonal or transverse epibranchial ridges; frontal region grossly produced, with pair of carinae diverging anteriorly; anterolateral margins irregular to rounded but sloping posteriorly giving carapace approximately triangular or polygonal outline. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article; antennular basis occupying lower half of fossa. Maxilliped 3 exopod with angular tooth at outer proximal margin. Cheliped slender, elongate, more than 3 times carapace length, palm at least 5 times as long as wide; fingers as long as or shorter than dorsal margin of palm. Male pleonites 3–5 fused. Maximum cl. 3.7 mm. Intertidal–shelf (18–45 m). Temperate Northern W Pacific. 3 species (Komatsu and Takeda 2001b: comparison of 3 species).

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569

Merocryptus A. Milne-Edwards, 1873

Myropsis Stimpson, 1871

Diagnosis. Carapace subtriangular-rhomboid, 1.0–1.3 times as wide as long; dorsal surface branchial region with strong oblique or transverse epibranchial ridge extending posterolaterally, widest posterolaterally; anterolateral margin with 1 or 2 wide blunt triangular lobes, epibranchial margin expanded as prominent rounded tooth or lobe. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially open, not closed-off from antennular fossa, antennal basal article movable. Cheliped fingers less than 1.6 times as long as palm, cutting edges dentate or crenulated. Male pleonites 3–6 fused. Gonopod 1 evenly tapering to sharp point. Maximum cl. 14 mm (Pl. 47e). Subtidal–slope (40–668 m). Temperate Northern and Tropical E Atlantic, Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Australasia. 4 species (Serène 1955).

Diagnosis. Carapace globose, as long or little longer than wide; dorsal surface smooth; posterior margin with 2 pairs of teeth and higher median tooth. Endostome reaching level of inhalant branchial channel, buccal cavern narrowly triangular. Inhalent branchial channel anterior margin forming suborbital margin, margin curved, lateral corner forming blunt angular point. Cheliped fingers slender, about as long as palm. Male pleonites 3–6 fused. Maximum cl. 69 mm (Pl. 47g). Shelf, slope (84–1048 m). Temperate Northern and Tropical W Atlantic. 1 species (Rathbun 1937).

Myra Leach, 1817 Diagnosis. Carapace ovate, or subcircular, globose; dorsal surface with even contours, without bosses, finely granulate; frontal region weakly produced beyond hepatic margin; subhepatic margin visible in dorsal view, with tubercle; posterolateral and posterior margins together with pair of teeth and higher median tooth, with line of closely-spaced granules. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 3 lobes, separated from suborbital margin by transverse groove or forming lower margin of orbit. Female maxilliped 3 endopod with longitudinal setose fringe; maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped elongate, finely granulate, fingers shorter or longer than palm, cutting edges with uneven spiniform or short triangular teeth. Male pleonites 3–6 fused, with posterior denticle. Maximum cl. 43 mm (Pl. 47f). Intertidal–slope (0–302 m). Temperate Northern Atlantic, Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. Myra subgranulata Kossmann, 1877 is an exotic species in the eastern Mediterranean. 18 species (Galil 2001b: key to 13 species).

Myrine Galil, 2001 Diagnosis. Carapace subcircular, globose; dorsal surface with even contours, without bosses, finely granulate; frontal region weakly produced beyond hepatic margin; subhepatic margin visible in dorsal view, without tubercle; lateral surface and margin evenly granulate, without line of closely-spaced granules, posterolateral and posterior margins together with pair of teeth and higher median tooth. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 3 lobes, separated from suborbital margin by transverse groove. Female maxilliped 3 endopod without longitudinal setose fringe; maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped elongate, palm thin, fingers about as long as palm, cutting edges with uneven short triangular teeth. Male pleonites 3–6 fused, with posterior denticle. Maximum cl. 28 mm. Subtidal–slope (15–320 m). Western and Central Indo-Pacific. 2 species (Galil 2001b: key to species).

Neparilia Ng, Devi & Kumar, 2018 Diagnosis. Carapace subglobular; dorsal surface densely covered with large rounded tubercles of different sizes; posterior margin with pair of teeth or lobes and higher median tooth. Endostome reaching level of inhalant branchial channel. Maxilliped 3 exopod much wider than ischium, outer margin convex, coxa reaching halfway to margin of exopod. Cheliped of male about twice carapace length, fingers longer than palm. Male pleonites 3–5 fused. Maximum cl. 41 mm (Pl. 47h). Shelf (100–220 m). Temperate Northern W Pacific, Central Indo-Pacific. 1 species (Ng et al. 2018).

Nobiliella Komatsu & Takeda, 2003 Diagnosis. Carapace angular, polygonal, slightly wider than long; dorsal surface variously tuberculate, with broad tuberculate median longitudinal ridge and diagonal epibranchial ridges; frontal region not produced beyond orbits, transversely sinuous; anterolateral margin produced as broad, thickened, granular-tuberculate lobe, posterolateral margin straight; posterior margin narrow, trilobed. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article; supraorbital margin with 1 fissure. Cheliped fingers about as long as palm. Female pleonites 1–3 free, 4–6 fused, pleonite 1 concealed medially by overlap of pleonite 2. Male pleonites 2–6 fused, with posterior denticle. Maximum cl. 7.3 mm. Intertidal, subtidal. Western Indo-Pacific. 3 species (Galil and Innocenti 2019: comparison of species; Komatsu and Takeda 2003).

Nucia Dana, 1852 Diagnosis. Carapace subcircular-polygonal in outline, globose; dorsal surface and margins coarsely tuberculate to multilobate; lateral margin convex, with 3 or 5 broad triangular teeth; posterior margin rounded. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article. Cheliped merus shorter than carapace length; fingers slender, shorter than upper margin of palm, usually opening almost horizontally. Pereopods 1–4 coarsely tuberculate. Female pleonites 1–3 free, 4–6 fused. Male pleonites 3–5 fused. Gonopod 1 longer than gonopod 2. Maximum cl. 11 mm (Pl. 47i).

570

Marine Decapod Crustacea

Intertidal–shelf (0–56 m). Temperate Northern Pacific, IndoWest Pacific, Tropical Eastern Pacific. 8 species (Serène 1954: diagnosis; 1955: key to species, one now in Nuciops, one in Heteronucia).

Nuciops Serène & Soh, 1976 Diagnosis. Carapace subhexagonal; dorsal surface tuberculate, uneven, regions distinct; lateral margin convex, tuberculate; posterior margin straight. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article. Cheliped fingers about as long as palm. Pereopods 1–4 smooth, stocky. Male pleonites 3–5 fused. Gonopod 1 as long as gonopod 2. Maximum cl. 5 mm. Subtidal (11–17 m). Central Indo-Pacific. 1 species (Serène and Soh 1976).

Nursia Leach, 1817 Diagnosis. Carapace angular, polygonal or deltoid, as wide as or wider than long; dorsal surface with median longitudinal ridge and diagonal epibranchial ridges, variously tuberculate; frontal region at most weakly produced beyond orbits, transversely straight or sinuous; anterolateral margin produced, broad, edge thickened, blunt often granular-tuberculate; posterior margin narrow, weakly bi- or trilobed. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closedoff from antennular fossa by immovable antennal basal article (or slightly movable); supraorbital margin with 2 fissures. Cheliped fingers about as long as palm. Female pleonites 1–3 free, 4–6 fused, or 1, 2 free, 3–6 fused. Male pleonites 3–6 fused. Maximum cl. 15 mm. Subtidal, shelf (1–94 m). Central and Eastern Indo-Pacific, Tropical Eastern Pacific. 21 species (Chen 1989: descriptions of 5 species; Komatsu and Takeda 2001a; Ng and Komatsu 2016: discussion).

Nursilia Bell, 1855 Diagnosis. Carapace subhexagonal, widest at epibranchial tooth; dorsal surface with ridges and spines, with 3 vertical cardiac spines, 1 or all prominent; anterolateral margin slightly expanded, with blunt edge, with 3 or 4 irregular obtuse teeth; posterior margin narrow, bilobed. Endostome reaching beyond anterior margin of inhalant branchial channel. Cheliped elongate, palm swollen, fingers at least 1.5 times as long as palm, cutting edges with uneven short perpendicular teeth. Male pleonites 3–6 fused. Maximum cl. 11 mm (Pl. 47j). Intertidal–shelf (0–229 m). Western and Central Indo-Pacific. 3 species (Chen 1989: comparison of species).

Oreophorus Rüppell, 1830 Diagnosis. Carapace subtriangular–oval, 1.2–1.5 times as wide as long; dorsal surface finely granular, with 6 large uneven oval fossae near anterolateral margins, connected by subsurface channels,

floor of fossae covered with mushroom-like granules; frontal region narrow, moderately upturned; anterolateral and posterolateral margins expanded, granule-lined, sinuous. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article; antennular basis occupying lower half of fossa. Cheliped fixed finger blade-like, twice as long as palm. Male pleon coarsely granular; pleonites 3–5 fused. Gonopod 1 2.6–3.2 times as long as gonopod 2. Maximum cl. 16 mm. Shelf (20–101 m). Western and Central Indo-Pacific. 7 species (Tan and Ng 1995: key to 4 species).

Oreotlos Ihle, 1918 Diagnosis. Carapace suboval, 1.3–1.5 times as wide as long; dorsal surface granular, with broad shallow groove parallel to margin; frontal region narrow, moderately upturned; anterolateral and posterolateral margins expanded to form conspicuous granulated rim. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article; antennular basis occupying lower half of fossa. Cheliped fingers as long to twice as long as palm. Male pleon coarsely granular; pleonites 3–5 fused, with prominent tubercle. Gonopod 1 2.0–2.4 times as long as gonopod 2. Maximum cl. 9.5 mm. Intertidal–shelf (0–140 m). Temperate Northern W Pacific, Indo-West Pacific. 12 species (Tan and Ng 1995: key to 11 species).

Orientotlos Sakai, 1980 Diagnosis. Carapace subtriangular, 1.2–1.3 times as wide as long; dorsal surface with 8 raised bosses (cardiac, intestinal, 2 gastric, 4 branchial), each rimmed by tubercles; anterolateral and posterolateral margins with 2 beaded lobes and 3 leaf-shaped branchial teeth. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article. Cheliped palm swollen, fingers as long as palm. Maximum cl. 5.5 mm. Shelf (116–159 m). Temperate Northern W Pacific, Central Indo-Pacific. 1 species (Ng and Chan 2021; Sakai 1980).

Ovilyra Ng, 2021 Diagnosis. Carapace globose, 1.2–1.3 times longer than wide; dorsal surface smooth, finely or coarsely granulate; frontal region upcurved, weakly angular medially, not exceeding epistome; margins beaded; posterior margin with lateral tooth separating posterolateral margin. Endostome reaching level of inhalant branchial channel. Cheliped fingers about as long as palm. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 1, 2 articulating, narrow; fused pleonites 3–5 tapering, without constrictions. Gonopod 1 slender, elongate, distal quarter sharply bent backwards from longitudinal axis; with curved, tapering subdistal process. Gonopod 2 apex tapering, sharp, shorter than gonopod 1. Maximum cl. 16 mm. Estuarine, intertidal, subtidal (0–2 m). Central Indo-Pacific. 1 species (Ng 2021).

14 – Brachyura – crabs

Parilia Wood-Mason & Alcock, 1891 Diagnosis. Carapace transversely or longitudinally ovate, subglobular; dorsal surface smooth, or finely and evenly granular; anterolateral-posterolateral margins forming even curve; posterior margin with pair of teeth or lobes and higher median tooth. Endostome reaching level of inhalant branchial channel. Maxilliped 3 exopod flat, little wider than ischium. Cheliped slender, elongate, more than 3 times carapace length, palm at least 5 times as long as wide; fingers as long as or shorter than dorsal margin of palm. Male pleonites 3–5 fused, with prominent cup-shaped tubercle. Maximum cl. 70 mm (Pl. 47k). Shelf, slope (90–549 m). Temperate Northern Pacific, Western and Central Indo-Pacific. 4 species (Ng et  al. 2018: rediagnosis, key to species).

Persephona Leach, 1817 Diagnosis. Carapace globose, as long or little longer than wide; dorsal surface smooth or granular, regions well defined, sometimes with low bosses; posterior margin with pair of teeth or lobes and higher median tooth. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin forming suborbital margin, margin curved, lateral corner forming blunt angular point. Maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped fingers slender, about as long as palm. Female pleonites 1–3 free, 4–6 fused. Male pleonites 3–5 fused. Maximum cl. 49 mm (Pl. 47m). Subtidal–shelf (9–112 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific, Temperate South America. 8 species (Rathbun 1937: rediagnosis, key to species).

Philyra Leach, 1817 Diagnosis. Carapace globose; dorsal surface smooth, finely or coarsely granulate; frontal region upcurved, with short medial tooth, not exceeding epistome; margins beaded, hepatic margin produced; posterior margin rounded. Endostome reaching level of inhalant branchial channel, anterior margin forming lower margin of orbit. Supraorbital margin without fissure. Cheliped fingers about as long as palm. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 1, 2 articulating, narrow; fused pleonites 3–6 tapering, without constrictions. Gonopod 1 elongate, flattened, apical process bifurcate, cornute. Gonopod 2 with scooplike apex, shorter than gonopod 1. Maximum cl. 22 mm. Intertidal–shelf (0–85 m). Western and Central Indo-Pacific, Temperate Australasia. 31 species (Galil 2009: key to 8 species).

Phlyxia Bell, 1855 Diagnosis. Carapace subrhomboidal, depressed; dorsal surface with large rounded tubercles of different sizes; posteromedian row of 2–4 prominent tubercles, anterior 1–3 upright, posteriormost directed posteriorly; lateral margins convex, rounded, with tubercles or spines. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 3 teeth or lobes. Orbit mesially open, not closed-off from

571

antennular fossa, antennal basal article movable. Cheliped fingers slender, about as long as palm. Male pleonites 3–5 fused. Maximum cl. 13 mm (Pl. 47l). Intertidal–shelf (0–99 m). Central Indo-Pacific, Temperate Australasia (NE and S Australia). 2 species. We resurrect Phlyxia for two Australian species originally described in this genus (see Introduction). For figures of P. crassipes Bell, 1855 see Poore (2004 as Ebalia) and for P. lambriformis Bell, 1855, see Bell (1855).

Praebebalia Rathbun, 1911 Diagnosis. Carapace subcircular, globose; dorsal surface with regions indistinct except for intestinal prominence; lateral margin convex with tooth at midpoint; posterior margin with 2 or 3 teeth. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially open, not closed-off from antennular fossa, antennal basal article movable. Cheliped merus 1.5 times as long as carapace; fingers about half as long as palm. Pereopods 1–4 smooth, slender. Male pleonites 3–5 fused. Maximum cl. 31 mm (Pl. 48a). Shelf, slope (156–400 m). Western and Central Indo-Pacific. 15 species (Galil 2001d: key to 4 species; Galil 2015: rediagnosis).

Praosia Tan & Ng, 1993 Diagnosis. Carapace angular, polygonal, slightly wider than long; dorsal surface strongly punctate, with 4 parallel, granular longitudinal ridges, epibranchial ridge short; frontal region produced, trilobed; anterolateral and posterolateral margins expanded, edges unevenly sinuous, thin, edges sharp, forming continuous laminar rim around carapace. Endostome reaching beyond anterior margin of inhalant branchial channel. Cheliped fingers shorter than palm. Female pleonites 1–3 free, 4–6 fused. Male pleonites 3–5 fused. Maximum cl. 10 mm (Pl. 48b). Intertidal, subtidal (mangroves). Central Indo-Pacific (Singapore). 1 species (Tan and Ng 1993).

Pseudomyra Capart, 1951 Diagnosis. Carapace subglobular; dorsal surface finely and evenly granular; subhepatic margin not visible in dorsal view; posterior margin with 2 prominent spines in horizontal row, without median spine or protuberance. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 3 teeth. Maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped elongate, palm thin, fingers half as long as palm, cutting edges with minute teeth; merus longer than carapace. Male pleonites 3–6 and telson fused. Maximum cl. 27 mm. Subtidal–slope (12–220 m). Tropical E Atlantic. 1 species (Capart 1951).

Pseudophilyra Miers, 1879 Diagnosis. Carapace longitudinally ovate, subglobose; dorsal surface smooth or evenly and finely granulate, with weak hepatic boss; frontal region with median blunt lobe produced beyond

572

Marine Decapod Crustacea

epistome and pterygostomial margin; lateral margins and posterior margins convex, rounded. Endostome reaching level of inhalant branchial channel. Maxilliped 3 exopod flat, as wide as endopod ischium at midlength. Cheliped stout, male palm no more than 3 times as long as wide, fingers shorter than palm. Male pleonites 3–6 fused. Maximum cl. 8 mm. Intertidal–shelf (0–70 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 15 species (Komai et al. 2019: list of species; Miers 1879).

Pyrhila Galil, 2009 Diagnosis. Carapace subcircular, globose; dorsal surface smooth, finely or coarsely granulate; frontal region upcurved, weakly bilobed, not exceeding epistome; lateral margin beaded; posterior margin rounded. Endostome reaching level of inhalant branchial channel, anterior margin forming lower margin of orbit. Inhalent branchial channel anterior margin with no more than 2 teeth or lobes. Supraorbital margin with 1 fissure. Maxilliped 3 exopod flat, as wide as endopod ischium at midlength. Cheliped merus shorter than carapace in male, fixed finger without fringe of setae. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 1 almost as wide as pleonite 2; pleonites 2–6 fused, without subdistal denticle. Gonopod 1 elongate, flattened, apical process tubular. Gonopod 2 with scoop-like apex, shorter than gonopod 1. Maximum cl. 23 mm (Pl. 48c). Intertidal–shelf (0–25 m). Temperate Northern W Pacific, Central Indo-Pacific. 3 species (Galil 2009).

Randallia Stimpson, 1857 Diagnosis. Carapace subglobular; dorsal surface smooth, finely or coarsely granulate; subhepatic margin not visible in dorsal view; posterior margin with 5 spines or protuberances (1 dorsomedian, 2 pairs around posterior and posterolateral margins), or rounded (R. curacaoensis Rathbun, 1922, only). Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 3 teeth or lobes. Maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped fingers longer than palm. Female pleonites 1–3 free, 4–6 fused, swollen. Male pleonites 3–5 fused; telson tapering. Gonopod 1 distally club-shaped. Maximum cl. 56 mm. Subtidal–shelf (4–93 m). Tropical W Atlantic, Eastern IndoPacific. 5 species (Galil 2003c; Rathbun 1937: key to species). Galil (2003c) restricted Randallia to American species. She left as incertae sedis several Indo-West Pacific species described within this genus that do not fall within her restricted diagnosis. Randallia gilberti Rathbun, 1906, is herein included in Toru; and Randallia agaricias Rathbun, 1898, and R. americana (Rathbun, 1894) are included in Tanaoa.

Raylilia Galil, 2001 Diagnosis. Carapace subglobular; dorsal surface and margins covered with large, sharp to blunt tubercles; posterior margin with 3 blunt spines or rounded lobes in horizontal row, with or without

median lobe or spine on higher plane. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin divided by narrow fissure; separated from suborbital margin by transverse groove. Orbit mesially closed-off from antennular fossa by immovable antennal basal article. Maxilliped 3 exopod narrower than endopod ischium at midlength. Cheliped elongate, palm thin, fingers about as long as palm, cutting edges with uneven short triangular teeth. Female pleonites 1–3 free, 4–6 fused, swollen. Male pleonites 3–6 fused, with posterior denticle. Gonopod 1 with flared, triangular or trilobed tip. Maximum cl. 13 mm (Pl. 48d). Intertidal–shelf (0–97 m). Western and Central Indo-Pacific. 5 species (Galil 2001a: key to 4 species).

Ryphila Galil, 2009 Diagnosis. Carapace globose, subpentagonal; dorsal surface smooth, finely or coarsely granulate; frontal region upcurved, weakly bilobed, not exceeding epistome; epistome and subhepatic margin projecting beyond front; lateral margin beaded; posterior margin rounded. Endostome reaching level of inhalant branchial channel, anterior margin forming lower margin of orbit. Supraorbital margin with 1 fissure. Cheliped fingers about as long as palm. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 1, 2 articulating, narrow, laterally swollen; pleonites 3–6 fused. Gonopod 1 elongate, flattened, apical process tubular. Gonopod 2 with scooplike apex, shorter than gonopod 1. Maximum cl. 14 mm (Pl. 48e). Subtidal (2–25 m). Western and Central Indo-Pacific, Temperate Australia. 3 species (Galil 2009).

Speloeophoroides de Melo & Torres, 1998 Diagnosis. Carapace subhexagonal; dorsal surface with 4 deep cave-like hollows on posterior half, most evident when viewed posteriorly; frontal region not produced; lateral margin with 1 or 2 branchial lobes. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article. Cheliped fingers shorter than palm. Female pleonites 1–3 free, 4–6 fused. Male pleonites 3–6 fused. Maximum cl. 8 mm. Shelf (41 m). Tropical W Atlantic. 1 species (de Melo and Torres 1998a).

Speloeophorus A. Milne-Edwards, 1865 Diagnosis. Carapace subcircular-subhexagonal; dorsal surface with 2 or 4 deep cave-like hollows on posterior half, most evident when viewed posteriorly; frontal region not produced; lateral margin with 1 or 2 branchial lobes. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article. Cheliped fingers shorter than palm. Female pleonites 1–3 free, 4–6 fused. Male pleonites 3–5 fused; pleonite 6 with strong spine directed anteriorly. Maximum cl. 27 mm (Pl. 48f). Intertidal, shelf (0–223 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 9 species (de Melo and Torres 1998b: rediagnosis key to 3 species; Rathbun 1937: key to 5 species; Williams 1984).

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573

Tanaoa Galil, 2003

Toru Galil, 2003

Diagnosis. Carapace subglobular; dorsal surface variously granular, regions demarcated; posterior margin with pair of rounded or angular lobes in horizontal row, with or without pair of branchial lobes, blunt median lobe or spine on higher plane (intestinal spine). Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 2 lobes, separated from suborbital margin by transverse groove or larger space. Orbit mesially open, not closed-off from antennular fossa, antennal basal article movable. Maxilliped 3 exopod narrower than endopod ischium at midlength. Pereopods 1–4 smooth, slender. Female pleonites 1–3 free, 4–6 fused, swollen. Male pleonites 3–6 fused; telson tapering. Gonopod 1 tapering, bearing short preapical process. Maximum cl. 44 mm (Pl. 48g). Subtidal–slope (5–1250 m). Indo-West Pacific, Tropical Eastern Pacific, New Zealand. 11 species (Galil 2003c). We include Randallia agaricias Rathbun, 1898, and R. americana (Rathbun, 1894) in Tanaoa.

Diagnosis. Carapace subglobular; dorsal surface smooth, finely or coarsely granulate; posterior margin with 2 rounded lobes in horizontal row, with low median lobe on higher plane (intestinal spine). Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 2 lobes, separated from suborbital margin by transverse groove or larger space. Orbit mesially open, not closed-off from antennular fossa, antennal basal article movable. Maxilliped 3 exopod narrower than endopod ischium at midlength. Female pleonites 1–3 free, 4–6 fused, swollen. Male pleonites 2–6 fused, with posterior denticle; telson lingulate. Gonopod 1 tapering, L-shaped, with long apical process at right angle to main axis. Maximum cl. 36 mm (Pl. 48i). Shelf, slope (35–648 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 6 species (Galil 2003c: key to 5 species). We include Randallia gilberti Rathbun, 1906, in Toru.

Tlos Adams & White, 1849 Diagnosis. Carapace oval, 1.5 times as wide as long; dorsal surface finely granular, with narrow median ridge, branchial region concave, with large D-shaped projection; frontal region strongly upturned, flat; anterolateral and posterolateral margins expanded, thin, plate-like, upturned, with 3 deep sutures. Endostome reaching level of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article; antennular basis occupying entire fossa. Cheliped fingers about as long as palm. Male pleon coarsely granular; pleonites 3–5 fused. Gonopod 1 2.0–2.4 times as long as gonopod 2. Maximum cl. 10 mm. Subtidal–shelf (7–55 m). Central Indo-Pacific. 1 species (Tan and Ng 1995: rediagnosis). The species mimics the dead, white fragments of the macroalga Halimeda (Tan and Richer de Forges 1993).

Tokoyo Galil, 2003 Diagnosis. Carapace subglobular; dorsal surface and margins finely and evenly granular; posterior margin with 3 blunt spines or rounded lobes in horizontal row, without median lobe or spine on higher plane. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin divided by narrow fissure; separated from suborbital margin by transverse groove. Orbit mesially open, not closed-off from antennular fossa, antennal basal article movable. Maxilliped 3 exopod narrower than endopod ischium at midlength. Female pleonites 1–3 free, 4–6 fused, swollen. Male pleonites 2–6 fused, with posterior denticle; telson lingulate. Gonopod 1 with slender, hooked tip. Maximum cl. 35 mm (Pl. 48h). Shelf, slope (35–366 m). Temperate Northern W Pacific, Central Indo-Pacific, Temperate Australia. 2 species (Galil 2003c: key to species).

Uhlias Stimpson, 1871 Diagnosis. Carapace oval, 1.6 times as wide as long; dorsal surface with broad elevated median regions surrounded by large pits; frontal region not produced; anterior and posterolateral margins more or less continuous. Endostome reaching beyond anterior margin of inhalant branchial channel. Orbit mesially closed-off from antennular fossa by immovable antennal basal article; antennular basis occupying lower half of fossa. Cheliped fingers shorter than palm. Male pleonites 3–5 fused. Maximum cl. 5.6 mm. Intertidal, shelf (0–64 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Rathbun 1937: key to species).

Urashima Galil, 2003 Diagnosis. Carapace subrhomboidal, subglobose; dorsal surface rugose, irregularly granular-tuberculate, regions swollen, demarcated; lateral margins convex-angular, irregularly lobate, tuberculate, spinous. Endostome reaching level of inhalant branchial channel. Inhalent branchial channel anterior margin with 2 lobes, separated by narrow groove from lower orbital margin. Orbit mesially open, not closed-off from antennular fossa, antennal basal article movable. Pereopods 1–4 coarsely tuberculate. Female pleonites 1–3 free, 4–6 fused, swollen. Male pleonites 2–6 fused, with posterior denticle; telson tapering. Gonopod 1 stout, distally flattened. Maximum cl. 40 mm (Pl. 48j). Shelf, slope (50–839 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 2 species (Galil 2003c: key to species).

Subfamily Leucosiinae Samouelle, 1819 Leucosiines are possibly the only monophyletic subfamily, uniquely recognised by the possession of a thoracic sinus, a groove on the anterolateral carapace (Ng et al. 2008). Diagnosis. Carapace more or less spherical, often with prominent anterior region; dorsal surface highly polished; thoracic sinus present. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like.

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Fig. 14.70.  Leucosiidae, Leucosiinae. a, Leucosia anatum (Herbst, 1783); b, Urnalana cheverti (Haswell, 1879). Carapace, lateral: c, Euclosiana crosnieri (Chen, 1989); d, Leucosia craniolaris (Linnaeus, 1758). Left thoracic sinus: e, Euclosiana; f, Leucosia; g, Seulocia. Left anterior front, thoracic sinus, maxilliped 3, base of cheliped: h, Leucosia. Female maxilliped 3: i, Urnalana. Male pleon: j, Coleusia; k, Euclosiana; l, Liusius; m, Seulocia; n, Urnalana. Gonopod 1: o, Coleusia; p, Euclosiana; q, Leucosia; r, Liusius; s, Seulocia; t, Soceulia; u, Urnalana.

Coleusia Galil, 2006 Diagnosis. Carapace subpentagonal, globose, regions indistinct; dorsal surface glabrous, smooth or minutely punctate; thoracic sinus anterior margin defined by overhanging margin of pterygostomian region, ventral margin with row of granules. Female maxilliped 3 endopod with longitudinal setose fringe. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 3–5 fused, anterolaterally inflated, pleonite 6 with medial denticle. Gonopod 1 sinuous, coiled 3 times, ultimate coil swollen, setose, apical process cornute. Maximum cl. 45 mm (Pl. 48k). Intertidal–shelf (0–110 m). Temperate Northern Atlantic (Coleusia signata (Paulson, 1875) is an exotic species in the Mediterranean), Western and Central Indo-Pacific, Temperate Southern Africa. 7 species (Galil 2006a: key to 5 species; Giraldes et al. 2017: key to species, distribution).

Euclosiana Galil & Ng, 2010 Diagnosis. Carapace subpentagonal, globose, regions indistinct; dorsal surface glabrous, smooth or minutely punctate; thoracic

sinus anterior margin defined by beaded loop, ventral margin with row of granules. Female maxilliped 3 endopod with longitudinal setose fringe. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 3–5 fused, anterolaterally inflated, pleonite 6 with medial denticle. Gonopod 1 sinuous, coiled 3 times, with preapical ruff of setae, apical process cornute. Maximum cl. 37 mm (Pl. 48l). Shelf, slope (30–366 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia. 11 species (Galil 2003b: key to 9 species as Euclosia; Galil and Ng 2010: replacement name).

Leucosia Weber, 1795 Diagnosis. Carapace subovoid, globose, hepatic region tumescent, regions indistinct; dorsal surface glabrous, smooth or minutely punctate; thoracic sinus deep, anteriorly Y-shaped, ventral margin with row of granules. Female maxilliped 3 endopod with longitudinal setose fringe. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 3–5 fused, anterolaterally inflated, pleonite 6 with medial denticle. Gonopod 1 slightly sinuous,

14 – Brachyura – crabs

tightly-coiled 5 or more times, apical process conical, with swollen setose muff. Maximum cl. 24 mm (Pl. 48m). Subtidal–shelf (2–80 m). Central Indo-Pacific, Temperate ­Australasia. 23 species (Galil 2003a: key to 4 species). Although several species have been removed to other genera Leucosia remains diverse.

Liusius Shih, Ho & Galil, 2020 Diagnosis. Carapace subpentagonal, globose; dorsal surface glabrous, smooth or minutely punctate; thoracic sinus with row of 6 granules above cheliped basis, median 4 largest. Female maxilliped 3 endopod with longitudinal setose fringe. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 3–5 fused, anterolaterally inflated, pleonite 6 without medial denticle. Gonopod 1 coiled 8 times, apex with setose lobe with slender process. Maximum cl. 31 mm. Shelf (48–82 m). Central Indo-Pacific. 1 species (Shih et  al. 2020).

Seulocia Galil, 2005 Diagnosis. Carapace subpentagonal, globose, regions indistinct; dorsal surface glabrous, smooth or minutely punctate; thoracic sinus anterior margin defined by overhanging margin of pterygostomian region, ventral margin with row of granules. Female maxilliped 3 endopod with longitudinal setose fringe. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 3–6 fused, with constriction at suture 5/6. Gonopod 1 straight, coiled once, distally club-like, bilobed. Maximum cl. 25 mm (Pl. 48n). Subtidal, shelf (5–91 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia. 11 species (Galil 2005b: key to 10 species).

Soceulia Galil, 2006 Diagnosis. Carapace subpentagonal, globose, regions indistinct; dorsal surface glabrous, smooth or minutely punctate; thoracic sinus anterior margin defined by overhanging margin of pterygostomian region, ventral margin with row of granules. Female maxilliped 3 endopod with longitudinal setose fringe. Female pleonites 1, 2, free, 3–6 fused, greatly enlarged, shield-like. Male pleonites 3–5 fused, anterolaterally inflated, pleonite 6 with medial denticle. Gonopod 1 coiled once, apical process encased in setose muff. Maximum cl. 39 mm. Subtidal, shelf (3–88 m). Central Indo-Pacific. 4 species (Galil 2006b: key to 4 species).

Urnalana Galil, 2005 Diagnosis. Carapace subpentagonal, globose, regions indistinct; dorsal surface sparsely setose, punctate; thoracic sinus anterior margin defined by overhanging margin of pterygostomian region, ventral margin with row of granules. Female maxilliped 3 endopod without longitudinal setose fringe. Female pleonites 1, 2 free,

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3–6 fused, greatly enlarged, shield-like. Male pleonites 3–5 fused, anterolaterally inflated, pleonite 6 with medial denticle. Gonopod 1 straight, with distal setae, apical process rounded or cornute. Maximum cl. 18 mm (Pl. 48o). Intertidal–shelf (0–200 m). Temperate Northern W Pacific, Indo-West Pacific. 22 species (Galil 2005a: key to 15 species). References Alcock A (1896) Materials for a carcinological fauna of India. No. 2. The Brachyura Oxystoma. Journal of the Asiatic Society of Bengal 65, 134–296, pls 6–8. Barnard KH (1950) Descriptive catalogue of South African decapod Crustacea (crabs and shrimps). Annals of the South African Museum 38, 1–837. Bell T (1855) Horae carcinologicae, or notices of crustacea. I. A monograph of the Leucosiidae, with observations on the relations, structure, habits and distribution of the family; a revision of the generic characters; and descriptions of new genera and species. Transactions of the Linnean Society of London 21, 277–314, pls 30–34. Capart A (1951) Crustacés Decapodes Brachyures. Expedition Océanographique Belge dans les Eaux Côtières Africaines de I’Atlantique Sud (1948–1949), Résultats Scientifiques 3, 11–205, pls 1–3. Chen H-L (1989) Leucosiidae (Crustacea, Brachyura). In: Forest J (Ed.), Résultats des Campagnes MUSORSTOM, vol. 5. Mémoires du Muséum National d’Histoire Naturelle, Paris 144, 181–264. Chen HL, Sun H (2002) Arthropoda Crustacea. Brachyura. Marine primitive crabs. Fauna Sinica. Invertebrata 30, 1–597, pls 1–4, 1–16. Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology - Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. de Melo GAS (1996) Manual de identifição dos Brachyura (Caranguejos e siris) do litoral Brasileiro. Editora Plêiade Ltda, Sao Paulo. de Melo GAS, Torres MFA (1998a) Novo gênero e espécie de Leucosiidae (Decapoda, Brachyura) no litoral brasileiro. Iheringia, Série Zoologia, Porto Alegre 84, 129–132. de Melo GAS, Torres MFA (1998b) O gênero Speloeophorus (Decapoda, Brachyura, Leucosiidae) no litoral brasileiro, com descrição de uma nova espécie. Iheringia, Série Zoologia 84, 121–128. Galil BS (2000) Acanthilia, a new genus of leucosiid crabs (Crustacea: Brachyura) from the Atlantic coast of the Americas. Proceedings of the Biological Society of Washington 113, 426–430. Galil BS (2001a) A new genus and species of leucosiid crab (Crustacea, Decapoda, Brachyura) from the Indo-Pacific Ocean. Zoosystema 23, 65–75. Galil BS (2001b) A revision of Myra Leach, 1817 (Crustacea: Decapoda: Leucosioidea). Zoölogische Mededeelingen 75, 409–446. Galil BS (2001c) A revision of the genus Arcania Leach, 1817 (Crustacea: Decapoda: Leucosioidea). Zoölogische Mededeelingen 75, 169–206. Galil BS (2001d) A revision of the genus Praebebalia Rathbun, 1911 (Brachyura, Leucosioidea). Journal of Crustacean Biology 21, 266–274. doi:10.1163/20021975-99990122 Galil BS (2003a) Contributions to the knowledge of Leucosiidae I. The identity of Leucosia craniolaris (Linnaeus, 1758), and redefinition of the genus Leucosia Weber, 1795 (Crustacea: Brachyura). Zoölogische Mededeelingen 77, 181–191. Galil BS (2003b) Contributions to the knowledge of Leucosiidae II. Euclosia gen. nov., (Crustacea: Brachyura). Zoölogische Mededeelingen 77, 331–347.

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Williams AB (1984) Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C. Zariquiey Alvarez R (1968) Crustáceos decápodos ibéricos. Investigaciones Pesqueras 32, 1–510.

Majoidea Samouelle, 1819 spider crabs, decorator crabs Majoids are commonly known as spider crabs or decorator crabs because some have long thin legs and others are able to camouflage themselves. Naturally, in a superfamily so large as this one not all species have long legs and not all use camouflage. The most easily recognised members of this superfamily have a pyriform body, tapering anteriorly to a more or less prominent rostrum, often bifurcated. Some species have a spiny carapace and the eyestalks are usually protected by spines whose arrangement is critical to their classification. Of course there are exceptions, some species are flattened, others are more rectangular or round, and a few are poorly spinose. The chelipeds are never massive, usually only slightly stronger and shorter than the walking legs. Majoids are unique in possessing stiff hooked setae on the exoskeleton that are used to attach camouflage material such as algae, sponges, hydroids, bryozoans or whatever is loose or readily detached in their environment. The crabs use their chelipeds to select and attach these extraneous materials (see review by Guinot and Wicksten 2015). The behaviour is plesiomorphic within the superfamily but has been lost in some groups, notably in deep-sea genera (­Hultgren and Stachowicz 2009, 2011). Majoid crabs are especially diverse in tropical IndoPacific waters with most of the more than 930 species occurring there. While a few species can be found in the lower intertidal, usually camouflaged species in beds of algae, many are from shelf depths on sedimentary and reef environments. The taxonomy of the superfamily is fluid with more than 60 family, subfamily and tribe names proposed (Ng et  al. 2008) but traditionally only six or seven groups have been recognised. This guide recognises six families without subfamilies. These have been separated on the degree of protection offered to the eye by the orbit, expansions of the carapace above the eye and the basal article of the antenna below. Elevation of the subfamilies of Majidae, especially Epialtine, Inachinae, Inachoidinae, Majinae, Mithracinae and Oregoniinae, to family rank (Drach and Guinot 1983) has long been an option and was adopted by several authors (Martin and Davis 2001; Ng et al. 2008) but not all (Poore 2004). Research on the taxonomies of the American species (Garth 1958; Guinot and Richer de Forges 1997) and those

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of the Indo-West Pacific (Griffin and Tranter 1986) have progressed more or less independently. Ng et al. (2008) went some way towards reconciling these, adopting some of the suprageneric names proposed by Števčić (2005). Early studies of numerous majid species reared in the laboratory shed some light on the phylogeny of the superfamily (Webber and Wear 1981) but more recent work casts doubt on the monophyly of some of the subfamilies/­ families. Building on studies by Marques and Pohle (1998), these same authors used larval features to investigate cladistic relationships of what were then subfamilies of Majidae (Marques et al. 2003; Pohle and Marques 2000). The found that not all families are monophyletic but were uncertain of their precise relationships. In a study relying on molecular data, Oregoniidae (three genera included) and Inachidae (two genera included) were more or less supported (­Hultgren and Stachowicz 2008) while other subfamilies were found to be paraphyletic. Another study of molecular and larval morphological characters of 14 genera (Hultgren et  al. 2009) was based on too few genera to draw conclusions about subfamily relationships. The monophyly of Inachoididae was first proposed after examination of just eight genera (Drach and Guinot 1983) and confirmed by Santana (2008) who reviewed most of the 14 now included. Molecular support is based on few genera. The relationship between Oregoniidae and Inachidae was addressed using larval and molecular data from few genera (Marco-Herrero et  al. 2013). The molecular data did not support a close relationship between these two families, a result supported by Lee (2020) using mitogenomes. Molecular affinities and several similarities between zoeae of Oregoniidae (s.s.) and Ergasticus prompted Marco-­Herrero et  al. (2013) to remove five genera, Ergasticus, Bothromaia, Pleisticanthoides, Parapleisticantha and Pleistacantha from Inachidae and place them within Oregoniidae as a subfamily, Pleistacanthinae, an option with some support (Ahyong et al. 2019). Marco-Herrero et al. (2013) also reported larval similarities between oregoniids and three other inachid genera, Eurypodius, Platymaia and Cyrtomaia, reinforcing the superficial resemblance between Chionoecetes (Oregoniidae) and Cyrtomaia and Platymaia (until now in Inachidae) noted by Griffin and Tranter (1986). They made no taxonomic recommendations. Adults of Cyrtomaia, Platymaia and Vitjazmaia have a circular to subcircular carapace, orbits lying more or less transversely, and a prominent postorbital spine. The zoea of another inachid genus, Macrocheira, is more similar to those of Oregoniidae and Majidae than to other inachids (Clark and Webber 1991). We follow the proposal to exclude Macrocheira from Inachidae and include it ­Oregoniidae (Clark and Webber 1991; Guinot 2019; Guinot and Van Bakel 2020).

Windsor and Felder (2014) found Mithracidae to be monophyletic and confined to the Americas and East Atlantic but only after excluding the six Indo-West Pacific genera included by earlier authors (e.g. Griffin and Tranter 1986). While Windsor and Felder (2014) made no recommendations on which family they belonged, their molecular phylogram suggested affinities with a paraphyletic Epialtidae. Three other American genera hitherto included in Mithracidae were also excluded and placed in Pisinae, a subfamily of Epialtidae. The paraphyly of Epialtidae, comprising subfamilies Epialtinae, Pisinae and Tychinae, is well documented (Hultgren et al. 2009; Lee 2020; Tsang et al. 2014). The subfamilies too are not monophyletic (Hultgren et  al. 2009; Hultgren and S­ tachowicz 2008; Lee 2020; Marco-Herrero et  al. 2013). The subfamilies are not recognised in this guide. The unusual genus Micippa was excluded from Mithracidae by Windsor and Felder (2014) without deciding where it should belong. Lee (2020) found on the basis of its mitogenome that it belonged in a clade with four other epialtids plus one Atlantic mithracid. For the time being we include Micippa in Epialtidae. The Majidae s.s. scarcely feature in recent molecular phylogenies (Hultgren et al. 2009; Hultgren and Stachowicz 2008; Lee 2020) – their monophyly has not been tested. The degree of protection offered to the eyestalk by ridges, spines and lobes is the critical feature distinguishing the families and genera. This key to families is built on earlier ones to subfamilies (Griffin 1966; Griffin and Tranter 1986) which non-specialists find exceptionally hard to master. This is in part because of the difficulty in understanding the terminology applied to the orbit and the armature surrounding it, and different interpretations in the literature of the ‘rostrum’. It is essential to clean setae or camouflaging algae and encrusting fauna away from the rostrum and orbit of decorating species before attempting an identification. Read the couplets carefully and refer to the illustrations where the features are labelled (Figs 14.78, 14.82, 14.87, 14.90, 14.92). The eyestalk sits on the side or front of the carapace in an orbit which is more or less complete. The orbit comprises four components which together offer the eye various degrees of concealment. First, dorsally is a supraorbital eave extending laterally to more or less obscure the eye from above. The supraorbital eave may be simple but more often is ornamented anteriorly by a preorbital spine at the front and an antorbital spine behind (be sure not to confuse the preorbital spine with a spine of the basal antennal article that may protrude laterally from beneath the anterior end of the orbit). Second is a postorbital spine or lobe sitting behind the eyestalk and sometimes cupped to receive it. The third component is the so-called intercalated spine which lies between the supraorbital eave and the postorbital spine. The orbit is fully completed underneath only in Mithracinae where the

14 – Brachyura – crabs

antennal basal article is expanded laterally as the fourth component of the orbit. The antennal basal article is homologous to articles 2 and 3 of the antennal peduncle – article 1 bears the nephridiopore (antennal gland). Peduncular articles 4 and 5 articulate with the antennal basal article and the terminal flagellum. Typically the antennal basal article is fused basally and medially to the epistome, exceptions being seen only in oregoniids and some inachids. The shape and armature of the basal antennal article feature in many keys. When the orbit is incomplete the eye is unprotected by a wide eave or spines and is clearly visible from above and below. The rostrum sits dorsally between the eyestalks and orbits, and is variously single, bifid or trifid. Some authors

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have referred to paired spines or teeth as ‘pseudorostra’, treating the lower interantenular spine which attaches to the interantennular septum, as the true rostrum (Garth 1958; Guinot and Van Bakel 2020). Our use of ‘rostrum’ refers to the dominating process or paired processes, sometimes incorporating the interantennular spine. Diagnosis. Carapace usually pyriform, usually longer than wide, sometimes squarish; carapace, chelipeds and walking legs usually with hooked setae (sometimes very dense) used in camouflage. Buccal cavity approximately quadrate. Antennules folding longitudinally or almost so. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular. Gonopod 1 much longer than gonopod 2.

Key to families of Majoidea 1. – 2. – 3. – 4. – 5. –

Antennal basal article broad, at most twice as long as wide. Orbits present, formed by supraorbital eave, adjacent spines and postorbital spine or lobe (Figs 14.87j–x, 14.90g–m)����������������������������������������������������������������������������2 Antennal basal article slender, usually at least twice as long as wide. Orbits usually absent, rarely with narrow, weakly developed supraorbital eave and small postorbital lobe (Figs 14.78, 14.82a–h, 14.84p–w, 14.92i–n)���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Antennal basal article broad, not expanded to form floor of orbit (Fig. 14.87j–x)��������������������������������������������������� Majidae Antennal basal article broad, expanded to form floor of orbit (Fig. 14.90g–m)�������������������������������������������������Mithracidae Orbits with narrow, weakly developed supraorbital eave partially overhanging eyes; with or without small postorbital lobe (Fig. 14.78)���������������������������������������������������������������������������������������������������������������������������������������Epialtidae Orbits absent, eyes unprotected though orbital margin usually with several small spines and postorbital spine����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������4 Male pleonite 6 posterior margin concave, telson recessed into concavity (if pleonite 6 and telson fused, traces of concave posterior margin of pleonite 6 visible); pleonites 3–5 always free (Fig. 14.93a–k). Gonopod 1 with prominent longitudinal groove (Fig. 14.93l–t)���������������������������������������������������������������������Oregoniidae Male pleonite 6 distal margin straight, telson not recessed into margin, or recessed, pleonites 3–5 fused (Paratymolus, Dumea). Gonopod 1 without prominent longitudinal groove�������������������������������������������������������������������5 Lateral parts of thoracic pleura 5–8 and often pleonite 1 integrated with carapace and fully visible in dorsal view. Pleotelson formed by fused pleonite 6 and telson in males, fused pleonites 5–6 and telson in females (Fig. 14.84o)��������������������������������������������������������������������������������������������������������������������������������������������� Inachoididae Lateral parts of thoracic pleura 5–8 and pleonite 1 entirely separate from carapace and not visible or only partially visible in dorsal view. Pleonite 6 and telson usually freely articulating in both sexes (Fig. 14.82u, v)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Inachidae

References Ahyong S, Mitra S, Ng PKL (2019) Redescription of Echinoplax rubida Alcock, 1895, a valid species of Pleistacantha from the Andaman Sea (Crustacea: Brachyura: Oregoniidae). Raffles Bulletin of Zoology 67, 391–395. Clark PF, Webber WR (1991) A redescription of Macrocheira kaempferi (Temminck, 1836) zoeas with a discussion of the classification of the Majoidea Samouelle, 1819 (Crustacea: Brachyura). Journal of Natural History 25, 1259–1279. doi:10.1080/00222939100770781 Drach P, Guinot D (1983) Les Inachoididae Dana, famille de Majoidea caractérisée par des connexions morphologiques d’un type nouveau entre carapace, pleurites, sternites, pléon. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 297, 37–42.

Garth JS (1958) Brachyura of the Pacific coast of America. Oxyrhyncha. Allan Hancock Pacific Expeditions 21, 1–854. Griffin DJG (1966) A review of the Australian majid spider crabs (Crustacea, Brachyura). Australian Zoologist 13, 259–98, pls 15–17. Griffin DJG, Tranter HA (1986) The Decapoda Brachyura of the Siboga Expedition. Part VIII Majidae. Siboga-Expéditie 39 c4, 1–335. Guinot D (2019) New hypotheses concerning the earliest brachyurans (Crustacea, Decapoda, Brachyura). Geodiversitas 41, 747–796. doi:10.5252/geodiversitas2019v41a22 Guinot D, Richer de Forges B (1997) Affinités entre les Hymenosomatidae MacLeay, 1838 et les Inachoididae Dana, 1851 (Crustacea, Decapoda, Brachyura). Zoosystema 19, 453–502. Guinot D, Van Bakel B (2020) Extraordinary majoid crabs: the genus Esopus A. Milne-Edwards, 1875 in the new subfamily Esopinae

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subfam. nov., and erection of Paulitinae subfam. nov. (Crustacea, Decapoda, Brachyura, Majoidea, Inachoididae Dana, 1851). Zootaxa 4766, 101–127. doi:10.11646/zootaxa.4766.1.5 Guinot D, Wicksten MK (2015) Chapter 71–11. Camouflage: carrying behaviour, decoration behaviour, and other modalities of concealment in Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 583–638. Brill, Leiden. Hultgren KM, Stachowicz JJ (2008) Molecular phylogeny of the brachyuran crab superfamily Majoidea indicates close congruence with trees based on larval morphology. Molecular Phylogenetics and Evolution 48, 986–996. doi:10.1016/j.ympev.2008.05.004 Hultgren KM, Stachowicz JJ (2009) Evolution of decoration in majoid crabs: a comparative phylogenetic analysis of the role of body size and alternative defensive strategies. American Naturalist 173, 566–578. doi:10.1086/597797 Hultgren KM, Stachowicz JJ (2011) Camouflage in decorator crabs. In Animal camouflage. (Eds Stevens MA, Merilaita S) pp. 214–238. Cambridge University Press, Cambridge. Hultgren KM, Guerao G, Marques FPL, Palero FP (2009) Assessing the contribution of molecular and larval morphological characters in a combined phylogenetic analysis of the superfamily Majoidea. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 437–455. CRC Press, Boca Raton. Lee S-H (2020). Systematic study on the superfamily Majoidea (Crustacea: Malacostraca: Decapoda) from Korean waters. Ph.D thesis. Seoul National University, College of Natural Sciences, Seoul. Marco-Herrero E, Torres AP, Cuesta JA, Guerao G, Palero F, et al. (2013) The systematic position of Ergasticus (Decapoda, Brachyura) and allied genera, a molecular and morphological approach. Zoologica Scripta 42, 427–439. doi:10.1111/zsc.12012 Marques FPL, Pohle G (1998) The use of structural reduction in phylogenetic reconstruction of decapods and a phylogenetic hypothesis for fifteen genera of Majidae: testing previous hypotheses and assumptions. Invertebrate Reproduction & Development 33, 241–262. doi:10.1 080/07924259.1998.9652636 Marques FPL, Pohle GW, Vrbova L (2003) On the larval stages of Macrocoeloma diplacanthum (Decapoda: Brachyura: Majidae), with a review of mithracine phylogenetic aspects. Journal of Crustacean Biology 23, 187–200. doi:10.1163/20021975-99990326 Martin JW, Davis GE (2001) An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County. Science Series 39, 1–124. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Pohle G, Marques FPL (2000) Larval stages of Paradasygyius depressus (Bell, 1835) (Crustacea: Decapoda: Brachyura: Majidae) and a phylogenetic analysis for 21 genera of Majidae. Proceedings of the Biological Society of Washington 113, 739–760. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Santana WRA (2008) Revisão taxonômica e relações filogenéticas em Inachoididae Dana, 1851 (Crustacea, Brachyura, Majoidea). PhD thesis. Universidade de São Paulo, São Paulo. doi:10.11606/T.41.2008. tde-29012009-100852 Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Tsang LM, Schubart CD, Ahyong ST, Lai JCY, Au EYC, et al. (2014) Evolutionary history of true crabs (Crustacea: Decapoda: Brachyura) and

the origin of freshwater crabs. Molecular Biology and Evolution 31, 1173–1187. doi:10.1093/molbev/msu068 Webber WR, Wear RG (1981) Life history studies on New Zealand Brachyura 5. Larvae of the family Majidae. New Zealand Journal of Marine and Freshwater Research 15, 331–383. doi:10.1080/00288330.1 981.9515929 Windsor AM, Felder DL (2014) Molecular phylogenetics and taxonomic reanalysis of the family Mithracidae MacLeay (Decapoda : Brachyura : Majoidea). Invertebrate Systematics 28, 145–173. doi:10.1071/ IS13011

Epialtidae Macleay, 1838 Figures 14.71–14.79, Plates 49–51 The orbits of members of this subfamily are undeveloped, with the often scarcely movable eyestalks sunken into the sides of the carapace. Many shallow-water species do not decorate but use their leaf-like body form as camouflage in algal habitats. Ng et al. (2008) and Davie et al. (2015) recognised four subfamilies, Epialtinae MacLeay, 1838, Pisinae Dana, 1851, Tychinae Dana, 1851 and Pliosomatinae Števčić, 1994. The first three were recognised as subfamilies of Majidae s.l. in the most comprehensive treatments, of American species (Garth 1958; Rathbun 1925) and of Indo-West Pacific species (Griffin and Tranter 1986). Pliosoma was recognised only more recently as a majoid (Guinot 1979) and later placed in its own subfamily. All Indo-West Pacific genera of Mithracinae listed by Griffin and Tranter (1986) and some American genera have been excluded from Mithracidae and included for the time being in ‘Pisinae’ (Windsor and Felder 2014). Taking these reallocations into account has made Epialtidae even more diverse than it was before. Further, the majid genus Thersandrus is now included (Colavite et  al. 2020). Other genera, such as Delsolaria, share features with epialtids and majids (Garth 1973). The subfamilies are unlikely to be monophyletic. We found that we could not rely on existing keys to subfamilies that depend on the nature of the orbit and eyestalks. Our key takes a more practical approach but could still be misinterpreted. Readers are advised to test both options, especially for couplet 42. Diagnosis. Orbit with narrow, weakly developed supraorbital eave partially overhanging eyes; with or without small postorbital lobe (or eyes sunk in sides of huge beak-like rostrum); intercalated spine absent; basal antennal article slender, usually more than twice as long as wide, always fused distally (rarely expanded to form floor of orbit, Delsolaria, Tiarinia). Thoracic pleurites 5–8 not visible dorsally, covered by posterolateral margin of carapace. Eyestalk short, stout, not completely concealed in dorsal view. Pleonite 1 free from to carapace. Maxilliped 3 merus as wide as ischium. Gonopod 1 slender, weakly curved,

14 – Brachyura – crabs

aperture usually terminal, apex simple, curved or expanded into lobes, rarely with slender process. Implicit generic attributes. Carapace pyriform. Postorbital lobe cupped. Antennal basal article narrow, with

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anterolateral tooth or spine invisible in dorsal view. Eyestalk protected on dorsal surface, exposed ventrally. Pereopods 2–5 not subchelate. Pleonites free in both sexes. Sexes similar.

Key to genera of Epialtidae 1.

Carapace branchial and/or hepatic margin flat, produced laterally, triangular or lobe-like, sometimes jointly or separately; carapace shape more or less rectangular or irregular����������������������������������������������������������������������2 – Carapace branchial margin convex, with or without spines or conical projections; carapace shape pyriform, ovoid, subcircular or subtriangular���������������������������������������������������������������������������������������������������������������������14 2. Carapace subrectangular, lateral margin broad, convex, entire. Rostrum semicircular, strongly produced laterally, concealing antennae (Fig. 14.75i)���������������������������������������������������������������������������������������������������������������Cyclonyx – Carapace subrectangular, triangular, irregular. Rostrum acute or bifid���������������������������������������������������������������������������������3 3. Rostrum bifid, split over most of length���������������������������������������������������������������������������������������������������������������������������������������4 – Rostrum acute, short and bilobed, or truncate����������������������������������������������������������������������������������������������������������������������������5 4. Carapace hepatic and branchial margins continuous, branchial margin expanded as triangular lamina. Rostrum margins converging (Fig. 14.72c)���������������������������������������������������������������������������������������������������������������Leucippa – Carapace hepatic and branchial margins separated by notch; branchial region with large plate arching beyond lateral margin. Rostrum comprising 2 flat plates with rounded apices (Fig. 14.77m)��������������Stegopleurodon 5. Rostrum truncate, shorter than wide, apex slightly excavate if at all, or contiguous plates. Carapace triangular�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������6 – Rostrum acute or apically bifid or bilobed. Carapace subquadrate, or triangular with triangular hepatic lobes�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������7 6. Carapace hepatic margin with concave flange; branchial margin with 2 rounded lobes, second larger (Fig. 14.76e). Pereopods 2–5 tuberculate. Maximum cl. 50 mm�����������������������������������������������������������������������������������Lissa – Carapace hepatic margin smooth; branchial margin with 2 rounded lobes, second larger. Pereopods 2–5 carinate (Fig. 14.72h). Maximum cl. 25 mm�����������������������������������������������������������������������������������������������������������Microlissa 7. Carapace preorbital angle absent (rarely present but not protruding laterally). Rostrum tapering or with parallel sides, truncate or with apical notch���������������������������������������������������������������������������������������������������������������������������8 – Carapace preorbital angle square. Rostrum acute, bilobed, rarely with apical notch�������������������������������������������������������� 12 8. Carapace without or with slight hepatic lobes. Rostrum truncate (Fig. 14.77l)����������������������������������������������� Simocarcinus – Carapace with prominent hepatic lobes. Rostrum with apical notch��������������������������������������������������������������������������������������9 9. Pleonites 4, 5 fused in male, pleonites 3–5 fused in female������������������������������������������������������������������������������������������ Epialtus – Pleonites 3–5 fused in both sexes�������������������������������������������������������������������������������������������������������������������������������������������������10 10. Eyestalks short, immovable. Carapace with dorsal bosses, hepatic margin obtusely rounded (Fig. 14.76o)�������� Mocosa – Eyestalks movable. Carapace smooth or tuberculate, hepatic margin with prominent lobe���������������������������������������������11 11. Carapace with strong tubercles (Fig. 14.75o). Pereopods 2–5 propodi with proximal thumb opposing dactylus (Fig. 14.79d)��������������������������������������������������������������������������������������������������������������������������������������������� Eupleurodon – Carapace nearly smooth or posteriorly eroded (Fig. 14.75m). Pereopods 2–5 propodi non-prehensile����������� Epialtoides 12. Rostrum acute, prominent (Fig. 14.71i). Pereopods 2–5 propodi flexor margins with tuft of setae and sometimes lobe in distal half (more or less prehensile) (Fig. 14.79e)�����������������������������������������������������������������������Huenia – Rostrum bilobed. Pereopods 2–5 propodi flexor margins with few setae����������������������������������������������������������������������������13 13. Carapace more or less rectangular, hepatic and branchial lobes squarish, similar. Rostrum visible in dorsal view (Fig. 14.77h)���������������������������������������������������������������������������������������������������������������������������������������������Sargassocarcinus – Carapace with hepatic lobe directed laterally, longer than branchial lobe, separated by deep U. Rostrum not visible in dorsal view, concealed by medial spine behind on upper surface of carapace (Fig. 14.77e)������������������������������������������������������������������������������������������������������������������������������������������������� Pteromaja 14. Carapace posterior half flat, oblique, bordered by ridge around branchial regions������������������������������������������������������������15

582

Marine Decapod Crustacea

Fig. 14.71.  Epialtidae. a, Anamathia rissoana (Roux, 1828); b, Chorilia turgida Rathbun, 1924; c, Chorinus heros (Herbst, 1790); d, Coelocerus spinosus A. Milne-Edwards, 1875; e, Crocydocinus ewok (Lee, Richer de Forges & Ng, 2019); f, Cyclocoeloma tuberculatum Miers, 1880; g, Delsolaria enriquei Garth, 1973; h, Hoplophrys oatesii Henderson, 1893; i, Huenia australis Griffin & Tranter, 1986.

– 15. – 16. – 17. – 18.

Carapace dorsally not divided by branchial ridges�������������������������������������������������������������������������������������������������������������������17 Carapace pyriform; branchial margin with 1 anterior spine; posterior margin with intestinal spine (Fig. 14.73k)������������������������������������������������������������������������������������������������������������������������������������������������������������Stilbognathus Carapace orbit and hepatic region laterally expanded; branchial margin without spine; posterior margin bilobed������������� 16 Carapace supraorbital eave with triangular preorbital tooth. Rostrum bilobed (Fig. 14.73l). Maxilliped 3 exopod basal mesial process straight, overlapping endite (Fig. 14.79l)����������������������������������Stilbomastax Carapace supraorbital eave with elongate preorbital tooth. Rostrum comprising 2 spines (Fig. 14.74b). Maxilliped 3 exopod basal mesial process curved, lying in groove on ischium (Fig. 14.79k)��������������������������������Tyche Carapace wider than long, branchial regions inflated, meeting but not fusing on midline; cardiac region obscured; hepatic margin and branchial region each with triangular tooth (Fig. 14.72f)���������� Lophorochinia Carapace branchial regions not inflated, cardiac region not obscured���������������������������������������������������������������������������������18 Carapace widest at anterolateral teeth; anterior third of carapace comprising orbits and rostrum triangular; posterior two-thirds D-shaped (Fig. 14.75r)������������������������������������������������������������������ Goniothorax

14 – Brachyura – crabs

583

Fig. 14.72.  Epialtidae. a, Lepteces ornatus Rathbun, 1893; b, Leptopisa setirostris (Stimpson, 1871); c, Leucippa pentagona H. Milne Edwards, 1834; d, Libidoclaea granaria H. Milne Edwards & Lucas, 1842; e, Libinia setosa Lockington, 1877; f, Lophorochinia parabranchia Garth, 1969; g, Macrocoeloma camptocerum (Stimpson, 1871); h, Microlissa tuberosa (Rathbun, 1898).

– 19.

Carapace typically pyriform or subcircular�������������������������������������������������������������������������������������������������������������������������������19 Intercalated spine present (rudimentary in Apiomithrax, sometimes fused in Micippa) (e.g. Figs 14.75f, 14.76j, 14.77b)���������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 20 – Intercalated spine absent (e.g. Figs 14.76q, 14.77o)������������������������������������������������������������������������������������������������������������������� 28 20. Rostrum of 2 broad, flat plates, depressed, fused for at least proximal two-thirds. Carapace ovoid, tuberculate (Fig. 14.76j, k)��������������������������������������������������������������������������������������������������������������������������������������������� Micippa – Rostrum otherwise. Carapace various����������������������������������������������������������������������������������������������������������������������������������������21 21. Carapace as wide as or wider than postrostral length. Rostrum comprising 2 short spines fused basally�������������������� 22

584

Marine Decapod Crustacea

Fig. 14.73.  Epialtidae. a, Nibilia antilocapra (Stimpson, 1871); b, Notolopas mexicanus Garth, 1940; c, Picroceroides tubularis Miers, 1886; d, Pisoides edwardsii (Bell, 1836); e, Pliosoma parvifrons Stimpson, 1860; f, Pohleus septemspinosus (Stimpson, 1871); g, Rhinocarcinus agassizi (Rathbun, 1893); h, Samadinia longispina Ng & Richer de Forges, 2013; i, Solinca aulix Colavite, Windsor & Santana, 2019; j, Sphenocarcinus corrosus A. Milne-Edwards, 1875; k, Stilbognathus tycheformis Bouvier, 1914; l, Stilbomastax margaritifera (Monod, 1939).

– Carapace narrower than postrostral length. Rostrum comprising 2 short or long spines������������������������������������������������ 23 22. Pereopods 2–5 twice carapace width. Intercalated spine rudimentary, on anterior face of postorbital spine (Fig. 14.75f). W Atlantic������������������������������������������������������������������������������������������������������������������������������ Apiomithrax – Pereopods 2–5 slender, up to 4 times carapace length. Intercalated spine as large as postorbital spine (Fig. 14.77b). Indo-West Pacific����������������������������������������������������������������������������������������������������������������������������Phalangipus

14 – Brachyura – crabs

585

Fig. 14.74.  Epialtidae. a, Stratiolibinia bellicosa (de Oliveira, 1944); b, Tyche emarginata White, 1847; c, Xenocarcinus tuberculatus White, 1847.

23. Rostral spines with dorsal accessory spine in distal half (Fig. 14.76p)��������������������������������������������������������������������� Naxioides – Rostral spines without accessory spine�������������������������������������������������������������������������������������������������������������������������������������� 24 24. Carapace densely spinose and tuberculate. Preorbital spine half as long as rostrum (Fig. 14.73a). Maximum cl. 120 mm������������������������������������������������������������������������������������������������������������������������������������������������������Nibilia – Carapace sparsely tuberculate. Preorbital angle triangular or obsolete. Maximum cl. 50 mm��������������������������������������� 25 25. Rostrum half or more carapace length, comprising 2 spines united in basal half, contiguous or diverging almost from base (Fig. 14.77u). Antennal basal article narrow, with strong anterolateral spine and broad proximal lobe on lateral margin adjacent basally to postorbital lobe (Fig. 14.78u). Indo-West Pacific������������������������������������������������������������������������������������������������������������������������������������������������ Tylocarcinus – Rostrum one-quarter to half carapace length, comprising 2 teeth or spines united in basal half. Antennal basal article separated from postorbital spine by deep V or U. Americas, Atlantic������������������������������������������������������ 26 26. Rostrum comprising 2 teeth, each as wide as long. Carapace with few small dorsal tubercles; hepatic and branchial margins aligned (Fig. 14.75u). Antennal basal article with 1 or 2 sharp lateral spines plus anterolateral spine (Fig. 14.78f)���������������������������������������������������������������������������������������������������������������������������Herbstia – Rostrum comprising 2 teeth, each longer than wide, parallel or diverging. Carapace regions without or with 1 tubercle or small spine; hepatic and branchial margins differentiated by shallow notch. Antennal basal article with or without blunt lateral tooth plus anterolateral spine����������������������������������������������������������������������� 27 27. Carapace with spines on most regions; lateral margins spinose or tuberculate (Fig. 14.77d). Antennal basal article with blunt lateral and anterolateral teeth, separated from postorbital spine by deep U (Fig. 14.78m)����������� Pisa – Carapace smooth, with cardiac, intestinal and pair of prominent branchial spines; lateral margins without spines (Fig. 14.73b). Antennal basal article broad, with proximolateral lobe next to postorbital lobe (Fig. 14.78k)������������������������������������������������������������������������������������������������������������������������������������������������������������������ Notolopas 28. Carapace as wide as or wider than postrostral length, globose or subcircular������������������������������������������������������������������� 29 – Carapace as wide as or wider than postrostral length, pyriform or ovoid����������������������������������������������������������������������������31 29. Carapace wider than postrostral length (Fig. 14.76q); pterygostomial region with strong spine exceeding front�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������Neodoclea – Carapace about as wide as postrostral length; pterygostomial region without strong spine�������������������������������������������� 30 30. Carapace with broad orbit and front between pair of anterolateral teeth; postorbital tooth small, triangular; posterolateral margin smooth (Fig. 14.77o). E Pacific����������������������������������������������������������������������������������������������Taliepus

586

Marine Decapod Crustacea

Fig. 14.75.  Epialtidae. Carapace, typical or only species: a, Acanthonyx; b, Acanthophrys; c, Alfredalcockia; d, Antilibinia; e, Apias; f, Apiomithrax; g, Austrolibinia; h, Criocarcinus; i, Cyclonyx; j, k, Cyphocarcinus (dorsal, lateral); l, Doclea; m, Epialtoides; n, Epialtus; o, Eupleurodon; p, Garthinia; q, Goniopugettia; r, Goniothorax; s, Griffinia; t, Guinotinia; u, Herbstia; v, Holoplites; w, Hyastenus.



Carapace globular, postorbital lobe prominent; lateral and posterior margins with spines (Fig. 14.75l). Indo-West Pacific��������������������������������������������������������������������������������������������������������������������������������������������������������������Doclea 31. Carapace triangular or ovoid, with large well defined plates (1 gastric, 1 cardiac, 1 intestinal, 2 hepatic, 2 branchial) divided by deep grooves����������������������������������������������������������������������������������������������������������������������������������������32 – Carapace pyriform, ovoid, rarely triangular, smooth or with tubercles not divided by deep grooves (rarely with well spaced plates)������������������������������������������������������������������������������������������������������������������������������������������������������������33 32. Carapace with pair of protogastric plates alongside mesogastric plate (Fig. 14.73j). W Atlantic�������������� Sphenocarcinus

14 – Brachyura – crabs

587

Fig. 14.76.  Epialtidae. Carapace, representative or only species: a, Lahaina; b, Laubierinia; c, Lepidonaxia; d, Leptomaia; e, Lissa; f, Loxorhynchus; g, Lyramaia; h, Menaethiops; i, Menaethius; j, k, Micippa (dorsal, anterior views); l, Micippoides; m, Micropisa; n, Minyorhyncha; o, Mocosoa; p, Naxioides; q, Neodoclea; r, Neophrys; s, Nicoya; t, Oplopisa; u, Oxypleurodon; v, w, Paranaxia (dorsal, lateral views); x, Pelia.

588

Marine Decapod Crustacea



Carapace without pair of protogastric plates alongside mesogastric plate (Fig. 14.76u). Indo-West Pacific, Australia������������������������������������������������������������������������������������������������������������������������������Oxypleurodon 33. Carapace about as wide as long, lateral margins smooth or finely tuberculate; supraorbital eave without preorbital tooth or spine��������������������������������������������������������������������������������������������������������������������������������������������������������� 34 – Carapace usually longer than wide, lateral margins usually ornamented; supraorbital eave with preorbital tooth or spine, rarely without������������������������������������������������������������������������������������������������������������������������������������������������� 38 34. Orbit oblique-transverse (Fig. 14.77p). Pereopods 2–5 flattened, prehensile (Fig. 14.79h)������������������������������ Thersandrus

Fig. 14.77.  Epialtidae. Carapace, representative or only species: a, Perinia; b, Phalangipus; c, Picrocerus; d, Pisa; e, Pteromaja; f, Pugettia; g, Rochinia; h, Sargassocarcinus; i, Scyra; j, Scyramathia; k, Siderochinia (lateral); l, Simocarcinus; m, Stegopleurodon; n Stenocionops; o, Taliepus; p, Thersandrus; q, Thusaenys; r, Tiarinia; s, Trachymaia; t, Tunepugettia; u, Tylocarcinus.

14 – Brachyura – crabs

– 35. – 36. – 37. – 38. – 39. – 40. – 41. – 42. – 43. – 44. – 45. – 46. – 47. – 48. – 49.

589

Orbit oblique or longitudinal. Pereopods 2–5 flat or cylindrical, not prehensile����������������������������������������������������������������35 Pereopods 2–5 carinate���������������������������������������������������������������������������������������������������������������������������������������������������������������� 36 Pereopods 2–5 not carinate�����������������������������������������������������������������������������������������������������������������������������������������������������������37 Antennal basal article little longer than wide, with rounded anterolateral lobe (Fig. 14.78n)���������������������������������Pisoides Antennal basal article longer than wide, with anterolateral lobe visible in dorsal view (Fig. 14.78l)�������������������������� Pelia Carapace smooth. Antennal basal article not visible in dorsal view (Fig. 14.75e)���������������������������������������������������������Apias Carapace tuberculate. Antennal basal article visible in dorsal view, lateral margin tuberculate, with blunt anterolateral tooth (Figs 14.76s, 14.78j)������������������������������������������������������������������������������������������������������Nicoya Pereopods 2–5 meri with 2 sharp teeth or lobes on extensor margin, tubercles on outer face (Fig. 14.79j). Carapace subcircular. Rostrum of 2 short teeth separated by U-shaped notch (Fig. 14.77a)�������������������������������Perinia Characters not so combined�������������������������������������������������������������������������������������������������������������������������������������������������������� 39 Rostrum single, cylindrical or flat or triangular with apical spine, sometimes with bifid apex. Postorbital spine obsolete or close to eye. Carapace smooth or with few tubercles������������������������������������������������������ 40 Rostrum double (if apically bifid postorbital spine prominent or carapace tuberculate)���������������������������������������������������41 Carapace teardrop-shaped. Rostrum cylindrical or flat, sometimes with bifid apex. Preorbital angle obsolete (Figs 14.74c, 14.78v)��������������������������������������������������������������������������������������������������Xenocarcinus Carapace with triangular branchial lobes. Rostrum triangular with apical spine. Preorbital angle triangular (Fig. 14.76i)���������������������������������������������������������������������������������������������������������������� Menaethius Rostrum comprising 2 leaf-shaped flat spines, fused in basal half, laterally expanded, with sharp apex (Fig. 14.77i)������������������������������������������������������������������������������������������������������������������������������������������������������� Scyra Rostrum otherwise������������������������������������������������������������������������������������������������������������������������������������������������������������������������ 42 Supraorbital eave and postorbital lobe fused, contiguous (separated by slit; e.g. Figs 14.75j, 14.76b–d, g, h, l, 14.77p), or widely separated by U-shaped or keyhole-shaped concavity (e.g. Figs 14.75b, c, h, p, 14.76b, m, 14.77c), antorbital angle acute or at least obtuse������������������������������������������������������������������������������������������������������������ 43 Supraorbital eave and postorbital lobe indistinctly separated by broad concavity (e.g. Figs 14.75s, v, 14.77s) or V-shaped notch (e.g. Figs 14.75q, t, 14.76.r, t, 14.77f, j, t), antorbital angle absent or nearly so��������������������������������75 Rostral spines parallel, one-third to half postrostral carapace length, with single accessory submesial or dorsal tooth (Figs 14.75k, 14.76w)������������������������������������������������������������������������������������������������������������������������������������������ 44 Rostral spines without single accessory tooth (some Tiarinia with lateral teeth)��������������������������������������������������������������� 45 Carapace narrow, ovoid. Preorbital spine small. Rostral accessory spine dorsal (Fig. 14.75j, k). Maximum cl. 21 mm������������������������������������������������������������������������������������������������������������������������������������������Cyphocarcinus Carapace pyriform, widest at branchial. Preorbital spine acute. Rostral accessory spine distomesial (Fig. 14.76v, w) Maximum cl. 116 mm�������������������������������������������������������������������������������������������������������������������� Paranaxia Orbits tubular, reaching maximum carapace width���������������������������������������������������������������������������������������������������������������� 46 Orbits tubular or not, not reaching maximum carapace width��������������������������������������������������������������������������������������������� 49 Rostrum comprising 2 diverging spines almost as long as postrostral carapace length���������������������������������������������������� 47 Rostrum comprising 2 short spines�������������������������������������������������������������������������������������������������������������������������������������������� 48 Orbit with spine-like postorbital spine, obsolete preorbital spine. Eyestalk extending well beyond orbit. Hepatic margin with 2 spines (Fig. 14.77c). Pacific���������������������������������������������������������������������������������������������� Picrocerus Orbit with triangular postorbital spine, strong preorbital spine. Eyestalk not extending beyond orbit. Hepatic margin with obsolete lobe (Fig. 14.73c). Atlantic�������������������������������������������������������������������������������Picroceroides Orbit with prolonged postorbital, antorbital and preorbital spines; preorbital angle separate from supraorbital eave. Rostrum bifid, directed laterally. Eyestalk extending well beyond orbit (Fig. 14.75h)������������������������������������������������������������������������������������������������������������������������������������������������������������� Criocarcinus Orbit with blunt postorbital, antorbital and preorbital angles; preorbital lobe upturned, obtuse, extending almost to level of rostrum apex. Rostrum triangular, bifid. Eyestalk scarcely extending beyond orbit (Fig. 14.71f)�������������������������������������������������������������������������������������������������������������������������������������Cyclocoeloma Rostrum Y-shaped, comprising 2 spines fused at base, curving and diverging abruptly. Carapace triangular, tomentose, with 7 strong spines in posterior half (Fig. 14.73f)������������������������������������������������������������ Pohleus

590

Marine Decapod Crustacea

– Rostrum bifid or comprising 2 spines, not Y-shaped. Carapace pyriform or ovoid����������������������������������������������������������� 50 50. Rostrum lyre-shaped (lateral edges convex proximally, spines parallel to divergent distally, apices curved outwards) (Fig. 14.76g)������������������������������������������������������������������������������������������������������������������������������������������������Lyramaia – Rostrum not lyre-shaped��������������������������������������������������������������������������������������������������������������������������������������������������������������51 51. Rostrum simple, tapering, apex emarginate, with ventral gutter. Carapace swollen, spinose. Postorbital lobe squarish (Fig. 14.71d)����������������������������������������������������������������������������������������������������������������������������������������Coelocerus – Rostrum double or single, without ventral gutter. Carapace and postorbital lobe various������������������������������������������������52 52. Carapace with rows of large mushroom-shaped tubercles, interspersed with cylindrical spines. Preorbital spine prominent, curved upwards. Pereopods spinose, pereopod 1 longest (Fig. 14.72a)���������������������������������� Lepteces – Carapace without rows of large mushroom-shaped tubercles. Preorbital spine usually triangular. Pereopods usually smooth�������������������������������������������������������������������������������������������������������������������������������������������������������53 53. Supraorbital eave, postorbital lobe, hepatic margin aligned, evenly merging into convex branchial margin. Eye visible only between eave and postorbital lobe. Rostrum comprising 2 flat diverging spines, one-fifth postrostral carapace length (Fig. 14.76c)�������������������������������������������������������������������������������Lepidonaxia – Supraorbital eave, postorbital lobe, hepatic margin, branchial margin variously prominent. Eye usually clearly visible lateral to eave. Rostrum various���������������������������������������������������������������������������������������������� 54 54. Carapace ovoid, inflated, regions marked, with 4 erect, curved medial blunt spines, pair on branchial regions; margins with 4 spines. Orbit tubular, directed anteriorly (Fig. 14.73e)�������������������������������������������������Pliosoma – Carapace pyriform, ovoid or triangular, not so ornamented. Orbit directed laterally�������������������������������������������������������55 55. Carapace with elevated swellings, some flat; hepatic region particularly elevated; branchial margin with 1 or 2 flat plates separated by groove from rest of carapace (Fig. 14.76b)����������������������������������������������Laubierinia – Carapace tuberculate or spinose, never with flat swellings; branchial margin smooth or tuberculate or with spine(s)������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 56 56. Carapace triangular, with prominent triangular branchial lobe. Rostrum comprising 2 long, parallel, thick, contiguous, obtuse spines, fused over one-third length. Postorbital lobe merged with hepatic margin, together separated by deep notch from branchial margin (Fig. 14.75p)���������������������������������������������� Garthinia – Carapace pyriform or ovoid. Postorbital lobe usually distinct from hepatic margin, latter more or less merging with branchial margin����������������������������������������������������������������������������������������������������������������������������������������������57 57. Rostrum triangular, as wide as long, with apical notch. Postorbital spine prominent, acute. Branchial region with 1 long spine directed laterally (Fig. 14.75c)�������������������������������������������������������������������������������� Alfredalcockia – Rostrum of 2 spines, sometimes fused in part. Postorbital spine and branchial region various�������������������������������������� 58 58. Carapace triangular, with few tubercles, cardiac region elevated. Rostrum as wide as long, with V-shaped notch over distal half (Fig. 14.71g). Antennal basal article broad proximally, with strong anterolateral tooth visible in dorsal view (Fig. 14.78c)������������������������������������������������������������������������������������������Delsolaria – Characters not so combined���������������������������������������������������������������������������������������������������������������������������������������������������������59 59. Carapace ovoid, orbits almost as wide as maximum carapace width. Supraorbital eave expanded, antorbital angle produced next to adjacent postorbital spine. Rostrum comprising 2 diverging spines one-quarter carapace length (Fig. 14.76d)������������������������������������������������������������������������������������������������������������������������������������Leptomaia – Characters not so combined; carapace usually pyriform; antorbital angle produced only if remote from postorbital spine������������������������������������������������������������������������������������������������������������������������������������������������������������� 60 60. Rostrum fused over at least basal half, distal section with apical notch or parallel or diverging spines�������������������������61 – Rostrum with 2 teeth or spines diverging from base or basal third�������������������������������������������������������������������������������������� 65 61. Antennal basal article with distolateral spine often visible in dorsal view (Fig. 14.78i). Carapace hepatic and branchial margins swollen, usually with tubercle or short spine (Fig. 14.76h)����������������������� Menaethiops – Antennal basal article distolateral angle rarely visible in dorsal view. Carapace hepatic and branchial margins swollen or not, with tubercle or spines������������������������������������������������������������������������������������������������������������������62 62. Pleonites 4–6 fused in female. Gonopod 1 simple (Fig. 14.79q). Central Pacific���������������������������������������������������������������� 63 – Pleonites free in female. Gonopod 1 bifid (Fig. 14.79r). Americas���������������������������������������������������������������������������������������� 64 63. Carapace with pair of short intestinal spines (Fig. 14.71h). Pereopod 2 less than twice carapace width�����������Hoplophrys

14 – Brachyura – crabs

591



Carapace with single prominent intestinal spine (Fig. 14.75g). Pereopod 2 up to 4 times carapace width������������������������������������������������������������������������������������������������������������������������������������������������������� Austrolibinia 64. Rostrum simple, apex emarginate or bifid. Carapace branchial margin with 3 or more small spines (Fig. 14.72e). Male sterno-abdominal cavity deep, narrow, closed anteriorly by crest on thoracic sternite 4, with lateral tubercles (Fig. 14.79m)��������������������������������������������������������������������������������������������������������������������������� Libinia – Rostrum broad, basally fused, ending in 2 widely divergent spines; branchial margin with 1 spine directed laterally (Fig. 14.74a). Male sterno-abdominal cavity deep, narrow, closed anteriorly by crest on thoracic sternite 4, without lateral tubercles (Fig. 14.79n)��������������������������������������������������������������������������������������������� Stratiolibinia

Fig. 14.78.  Epialtidae. Left antennal basal article, orbit: a, Acanthophrys; b, Chorilia; c, Delsolaria; d, e, Giranauria (dorsal, ventral views); f, Herbstia; g, Hyastenus; h, Lahaina; i, Menaethiops; j, Nicoya; k, Notolopas; l, Pelia; m, Pisa; n, Pisoides; o, Pteromaja; p, Crocyocinus; q, Scyramathia; r, Stilbomastax; s, Thusaenys; t, Tiarinia; u, Tylocarcinus; v, Xenocarcinus. Orbit, oblique view, antennal basal article, free articles: w, Samadinia. Features labelled on d, e: aba, antennal basal article; ao, antorbital spine; n, nephridiopore (antennal gland); po, preorbital spine; ps, postorbtial spine; r, rostral spine.

592

65. – 66. – 67. – 68. – 69. – 70. – 71. – 72. – 73. – 74. – 75. – 76. – 77. –

Marine Decapod Crustacea

Rostral spines triangular, deflexed, apices upcurved. Carapace with swollen triangular branchial plates (Fig. 14.76l). Pereopods and carapace covered with long soft setae�����������������������������������������������������������������Micippoides Rostral spines more or less horizontal. Carapace pyriform, usually with branchial spines��������������������������������������������� 66 Preorbital spine about half as long and parallel to diverging rostral spine (Fig. 14.76m). Maximum adult cl. 6 mm. E Atlantic�������������������������������������������������������������������������������������������������������������������������Micropisa Preorbital spine much less than rostral spine. Maximum adult cl. 15–147 mm. W Atlantic, Pacific����������������������������� 67 Carapace hepatic margin with 1 or 2 strong acute spines, branchial margin with 3 or more strong acute spines, dorsum spinose, often with median row of spines (Fig. 14.77n)�������������������������������������������������������Stenocionops Carapace hepatic margin smooth or with tubercle, branchial margin smooth or with 1 spine or tubercles, dorsum smooth, tuberculate or with short spines, usually with few tubercles�������������������������������������������������������������� 68 Supraorbital eave separated by U-shaped sinus from postorbital cup, together produced or not; antorbital angle square or produced. Antennal basal article narrow, eye with cornea and stalk exposed ventrally (Fig. 14.78a, d, s). Rostral spines diverging from near base����������������������������������������������������������������������������������������������� 69 Supraorbital eave in contact with postorbital cup, separated by slit, together more or less produced; antorbital angle obsolete. Antennal basal article broad, covering most of eye ventrally, usually only cornea visible (Fig. 14.78t). Rostral spines converging from base������������������������������������������������������������������������������������ 73 Supraorbital eave produced (preorbital face about as wide as rostral base). Antennal basal article having anterolateral tooth separated from article 2 by notch (Fig. 14.78a)�����������������������������������������������������������������������������������70 Supraorbital eave scarcely or not produced (preorbital face much less than rostral base). Antennal basal article anterolateral tooth not separated from article 2 by notch (Fig. 14.78h)�������������������������������������������������������������� 72 Postorbital lobe half cupped, anterodorsal margin with proximal flange, ventral margin without flange (Fig. 14.75b)����������������������������������������������������������������������������������������������������������������������������������������������������������� Acanthophrys Postorbital lobe fully cupped, with flanges on dorsal and ventral margins (Fig. 14.78e)���������������������������������������������������71 Mesogastric region with 3 strong tubercles or short spines (Fig. 14.78e). Carapace cardiac region with high tubercle or prominent cylindrical spine. Antennal basal article with blunt anterolateral angle (Fig. 14.78d)����������������������������������������������������������������������������������������������������������������������������������������������������������������Giranauria Mesogastric region smooth or with 1–3 low tubercles. Carapace cardiac region smooth or with 1 low tubercle (Fig. 14.77q). Antennal basal article with triangular anterolateral spine (Fig. 14.78s)�������������������� Thusaenys Antennal basal article narrow with strong anterolateral spine, lateral margin at base sometimes produced as dorsally extending lobe, separated from postorbital lobe by broad hiatus, with small suborbital tubercle (Fig. 14.78h). Pereopods 2, 3 meri with terminal spine (Fig. 14.79f)������������������������������������������������������Lahaina Antennal basal article without longitudinal groove, lateral marginal lobe, if present not extending dorsally into orbit, separated from postorbital lobe by narrow hiatus, with or without small suborbital tubercle (Fig. 14.78g). Pereopods 2, 3 meri with or without terminal spine�������������������������������������������������������������������� Hyastenus Antennal basal article with anterolateral spine visible in dorsal view. Rostrum ranging from 2 short teeth to 2 long spines (Fig. 14.72g)�����������������������������������������������������������������������������������������������������������������������������Macrocoeloma Antennal basal article with anterolateral spine not visible in dorsal view (Fig. 14.78t). Rostrum comprising 2 thin spines, separate from base��������������������������������������������������������������������������������������������������������������������������������������������74 Carapace anteriorly narrowed, interorbital width about half carapace width (Fig. 14.77r)������������������������������������ Tiarinia Carapace slightly narrowed anteriorly, interorbital width more than half carapace width (Fig. 14.72b)�����������Leptopisa Carapace branchial region or margin with 1 or 2 blunt or conical lobes; hepatic region or margin with blunt short or elongated lobe���������������������������������������������������������������������������������������������������������������������������������������������������76 Carapace branchial and hepatic regions or margins smooth, nodular or with sharp spines�������������������������������������������� 80 Pereopods 3–5 prehensile, dactylus closing against propodus (Fig. 14.79i). Carapace plates cushion-like. Rostrum comprising 2 short flat subparallel teeth (Fig. 14.75t)������������������������������������������������������������������������ Guinotinia Pereopods 3–5 simple. Carapace sculpture ridge-like or conical. Rostrum comprising 2 short or long spines or teeth��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 77 Branchial regions with 2 lateral projections������������������������������������������������������������������������������������������������������������������������������78 Branchial regions with 1 lateral projection������������������������������������������������������������������������������������������������������������������������������� 79

14 – Brachyura – crabs

593

78. Carapace triangular, hepatic and 2 branchial lobes increasing in length posteriorly. Rostral spines contiguous (Fig. 14.73g)������������������������������������������������������������������������������������������������������������������������������������� Rhinocarcinus – Carapace ovoid, first branchial lobe longest. Rostral spines separate (Fig. 14.75q)����������������������������������������Goniopugettia 79. Rostrum comprising 2 short, straight, diverging spines, one-fifth carapace length (Fig. 14.77t). Cheliped articles carinate (Fig. 14.79b)������������������������������������������������������������������������������������������������������������������������������� Tunepugettia – Rostrum comprising 2 straight, cylindrical diverging spines, one-third carapace length (Fig. 14.71e). Cheliped articles cylindrical��������������������������������������������������������������������������������������������������������������������������������Crocydocinus 80. Carapace covered with similar short spines or tubercles; often spherical. Americas only�������������������������������������������������81 – Carapace smooth or tuberculate and/or with long hepatic and branchial spines; usually pyriform������������������������������� 88 81. Rostrum bifid for more than half length����������������������������������������������������������������������������������������������������������������������������������� 82 – Rostrum bifid for less than half length�������������������������������������������������������������������������������������������������������������������������������������� 86 82. Rostrum thick, deflexed (Fig. 14.76f). Maximum cl. 200 mm���������������������������������������������������������������������������Loxorhynchus – Rostrum of 2 slender spines (e.g. Figs 14.76t, 14.77s). Maximum cl. 40 mm����������������������������������������������������������������������� 83 83. Pereopods with 2 rows of marginal spines (Fig. 14.79g)���������������������������������������������������������������������������������������������� Oplopisa – Pereopods without rows of marginal spines����������������������������������������������������������������������������������������������������������������������������� 84 84. Carapace postorbital spine with tooth on anterior-lower margin; supraorbital eave and postorbital lobe continuous, concave (Fig. 14.77s). Cornea flat�����������������������������������������������������������������������������������������������������Trachymaia – Carapace postorbital spine without tooth on anterior-lower margin; supraorbital eave and postorbital lobe separated by U- or V-shaped notch. Cornea spherical������������������������������������������������������������������������������������������������������� 85 85. Postorbital lobe triangular, separated from eave by V-shaped notch (Fig. 14.71b)��������������������������������������������������� Chorilia – Postorbital lobe acute, separated from eave by U-shaped notch (Fig. 14.73i)��������������������������������������������������������������Solinca 86. Carapace pyriform, densely covered with long spines. Supraorbital eave not covering eyestalk (Fig. 14.75v)�������� Holoplites – Carapace subcircular, covered with short spines. Supraorbital eave and postorbital cup enclosing eyestalk���������������� 87 87. Carapace branchial margin with 1 prominent spine longer than others (Fig. 14.72d)�������������������������������������� Libidoclaea – Carapace branchial margin with similar short spines (Fig. 14.76n)����������������������������������������������������������������� Minyorhyncha 88. Carapace elongate-pyriform, tomentose, with few tubercles on anterior half. Rostrum comprising 2 spines converging distally, almost half carapace length. Supraorbital eave with 2 marginal teeth. Preorbital lobe produced, stout. Hepatic margin with prominent blunt tooth (Fig. 14.71c)���������������������������������������������� Chorinus – Characters not so combined�������������������������������������������������������������������������������������������������������������������������������������������������������� 89 89. Carapace hexagonal, anterolateral margins longest, with short triangular hepatic and branchial teeth. Rostral spines short, separated by slit, its lateral margins convex, converging (Fig. 14.75d). Maximum cl. 72 mm. South Africa������������������������������������������������������������������������������������������������������������������������������������������������� Antilibinia – Carapace pyriform. Rostral spines separated by triangular sinus, its lateral margins straight, parallel, tapering, parallel or diverging. Maximum cl. 40 mm�������������������������������������������������������������������������������������������������������� 90 90. Carapace subrectangular or suboval; hepatic and branchial margins with strong lobes (Fig. 14.75a). Pereopods 2–5 subchelate, propodi broad, flexor margin with setiferous lobe (Fig. 14.79c)�����������������������Acanthonyx – Carapace pyriform; hepatic and branchial margins with teeth or spines. Pereopods 2–5 simple, propodi lower margin without lobe and dense setae��������������������������������������������������������������������������������������������������������������������������91 91. Gonopod 1 acute, with 2 sharp subapical flanges (Fig. 14.79s). Carapace epatic margin often expanded, laminar, triangular, separated from branchial margin by concavity (Fig. 14.77f)���������������������������������������������� Pugettia – Gonopod 1 acute, with subapical blade (Fig. 14.79p), or with truncate apex (Fig. 14.79t). Carapace hepatic margin sometimes expanded, or with spine, branchial margin with or without tubercle or spine��������������������������� 92 92. Carapace branchial margin convex, without spine or coarse tubercles (Fig. 14.75s); regions not indicated. Male cheliped merus, carpus and propodus carinate (Fig. 14.79a). Female pleonites 5, 6 fused���������������������� Griffinia – Carapace branchial margin usually with spine, often long, if absent, with coarse tubercles; regions indicated. Male cheliped merus, carpus and propodus with or without carinae. Female pleonites free������������������� 93 93. Carapace preorbital angle obsolete or blunt������������������������������������������������������������������������������������������������������������������������������ 94 – Carapace preorbital spine pointed (Fig. 14.73h, 14.78w)��������������������������������������������������������������������������������������������������������� 95

594

Marine Decapod Crustacea

Fig. 14.79.  Epialtidae. Left cheliped: a, Griffinia; b, Tunepugettia. Pereopod 2: c, Acanthonyx; d, Eupleurodon; e, Huenia; f, Lahaina; g, Oplopisa; h, Thersandrus. Pereopod 5: i, Guinotinia; j, Perinia. Maxilliped 3: k, Tyche; l, Stilbomastax. Male thoracic sternum, pleon: m, Libinia (tubercles arrowed) n, Stratiolibinia. Gonopod 1 apex: o, Anamathia; p, Griffinia; q, Hoplophrys; r, Libinia; s, Pugettia; t, Samadinia.

94. Rostral spines diverging at 60°, as long as carapace. Hepatic margin with 1 oblique spine, almost as long as rostral spine, longer than branchial marginal spine (Fig. 14.76r)������������������������������������������������������������������������ Neophrys – Rostral spines diverging at 30°, as long as carapace. Hepatic margin with 1 oblique spine, shorter than rostral spine, about as long as 3 branchial marginal spines (Fig. 14.71a)���������������������������������������������������������Anamathia 95. Carapace hepatic spine or lobe directed laterally (Fig. 14.77g); dorsal surface without spines or large tubercles�������������������������������������������������������������������������������������������������������������������������������������������������������������������������� Rochinia – Carapace hepatic spine or lobe directed inclined dorsally to fully upright; dorsal surface with large tubercles and/or spines������������������������������������������������������������������������������������������������������������������������������������������������������������ 96 96. Carapace cardiac region produced as tall, thick, blunt, peg-like dorsal projection (Fig. 14.77k)��������������������Siderochinia – Carapace cardiac region variously ornamented or not, but forming tall peg-like dorsal projection������������������������������� 97 97. Carapace with dorsal tubercles becoming large, flat-topped, leaf-like plates, wider than stem in adults (Fig. 14.77j)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Scyramathia – Carapace with prominent dorsal spines and/or large nodules and tubercles (Fig. 14.73h, 14.78w)�������������������Samadinia Acanthonyx Latreille, 1828 Diagnosis. Carapace subrectangular or ovoid, tomentose. Rostrum comprising 2 spines or lobes. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; preorbital spine triangular; postorbital spine absent, or spine acute. Hepatic margin expanded; branchial margin with 1 or more large triangular expansions. Pereopods 2–5 subchelate, propodi broad, flexor margin with setiferous lobe. Maximum cl. 28 mm. Intertidal–slope (0–290 m) (associated with macroalgae). Temperate Northern Atlantic, including Mediterranean, Tropical Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate Southern Africa. 17 species (Barnard 1950: key to 4 South African species as Dehaanius; Emparanza et al. 2007:

species redescriptions, distribution confirmed; Griffin and Tranter 1986: key to 9 Indo-West Pacific species; Manning and Holthuis 1981: key to 4 E Atlantic species; Wu et al. 1999: species listed with distributions).

Acanthophrys A. Milne-Edwards, 1865 Diagnosis. Carapace surface with many small tubercles and few short spines. Rostrum comprising 2 spines about one-third postrostral carapace length, divergent from near base. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; supraorbital eave strongly expanded, sometimes bilobed; preorbital spine triangular; postorbital lobe narrow, apex blunt, anterodorsal margin with proximal flange, ventral

14 – Brachyura – crabs

margin without flange. Antennal basal article having anterolateral angle separated from article 2 by notch. Maximum cl. 15 mm. Subtidal (2–3 m). Western and Central Indo-Pacific. 4 species (Griffin and Tranter 1986: key to 3 species).

Alfredalcockia Števčić, 2011 Diagnosis. Carapace branchial region with 1 long spine directed laterally. Rostrum triangular, as wide as long, with apical notch. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital spine triangular. Eyestalk extending well beyond eave. Maximum cl. 4 mm. Shelf (50–60 m). Western Indo-Pacific (India). 1 species (Alcock 1895: figure as Collodes malabaricus Alcock, 1895; Števčić 2005: diagnosis as Alcockia).

Anamathia Smith, 1885 Diagnosis. Carapace with strong spines: 3 medial; lateral branchial spines pointed outwards and downwards. Rostrum comprising 2 straight, spines diverging at 60°, as long as carapace. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; preorbital tooth blunt; postorbital lobe small, rounded. Pterygostomial region granular; hepatic margin with prominent oblique spine; branchial margin with 3 long spines. Gonopod 1 with oblique apex. Maximum cl. 27 mm. Shelf, slope (20–730 m) Temperate Northern Atlantic, including Mediterranean. 1 species (Lee et al. 2020: rediagnosis, species description; Tavares and Santana 2018: rediagnosis).

Antilibinia MacLeay, 1838 Diagnosis. Carapace hexagonal, anterolateral margins longest, flat. Rostrum comprising 2 short spines, separated by slit, lateral margins convex, converging. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; preorbital spine triangular; postorbital spine small. Hepatic margin with anteriorly directed spine; branchial margin with 1 tooth. Maximum cl. 72 mm. Intertidal, subtidal. Temperate Southern Africa. 1 species (Barnard 1950: figures; Richer de Forges 1994: figures, comparison with Griffinia).

Apias Rathbun, 1897 Diagnosis. Carapace tomentose. Rostrum comprising 2 leaf-shaped lobes. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine absent. Hepatic margin smooth; branchial margin smooth, slightly convex. Pereopods 2–5 with broad articles, setose. Maximum cl. 6 mm. Subtidal. Central Indo-Pacific (Fiji or Tonga). 1 species (Dana 1852: figure in Atlas). Apias pubescens (Dana, 1852) has not been recorded since its discovery.

Apiomithrax Rathbun, 1897 Diagnosis. Carapace subcircular, with tubercles and spines. Rostrum fused over basal half, 2 parallel or diverging spines distally.

595

Supraorbital eave and postorbital spine separated by intercalated spine; preorbital tooth rounded. Hepatic margin with small spines; branchial margin with 3 or mores spines. Eyestalk extending well beyond eave. Maximum cl. 50 mm. Intertidal–shelf (0–180 m) (Manning and Holthuis 1981: commentary; Monod 1956: descriptions, figures).

Austrolibinia Griffin, 1966 Diagnosis. Carapace with few slender spines, with median acute intestinal spine. Rostrum fused over basal half, 2 parallel or diverging spines distally. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital spine triangular. Antennal basal article lateral margin with 2 laterally flattened tubercles, or proximal margin produced ventrally as long compressed spine. Pereopods 2–5 slender, up to 4 times carapace length. Pleonites 4–6 fused in female. Gonopod 1 simple. Maximum cl. 27 mm. Subtidal, shelf (14–100 m). Central Indo-Pacific. 4 species (­Griffin and Tranter 1986: key to 3 species; Wagner 1992: description of fourth species).

Chorilia Dana, 1851 Diagnosis. Carapace surface with few tubercles and short spines, with epibranchial and intestinal spines. Rostrum comprising 2 spines about half postrostral carapace length. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine acute; postorbital lobe often flattened laterally, sometimes plate-like and/or confluent with hepatic spine or subacute anteriorly in lateral view; ventral margin in plane dorsal to that of antennal basal article. Maximum cl. 71 mm. Shelf, slope (35–1300 m). Temperate Northern Pacific, Eastern Indo-Pacific. 3 species (Garth 1958: rediagnosis, subspecies now recognised as species).

Chorinus Latreille, 1825 Diagnosis. Carapace pyriform, almost twice as long as wide, pubescent, with tubercles on anterior half. Rostrum comprising 2 spines converging distally, almost half carapace length. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; supraorbital eave with 2 marginal teeth; preorbital spine produced, stout. Antennal basal article lateral margin with tooth and anterolateral spine. Hepatic margin with prominent tooth; branchial margin convex. Maximum cl. 64 mm (Pl. 49a). Subtidal, shelf (1–50 m). Tropical W Atlantic. 1 species (Rathbun 1925: rediagnosis).

Coelocerus A. Milne-Edwards, 1875 Diagnosis. Carapace globular, with regions well defined, with strong conical spines. Rostrum tapering, apex emarginate, with ventral gutter. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; preorbital spine acute. Antennal basal article with anterolateral tooth visible in dorsal view. Hepatic margin

596

Marine Decapod Crustacea

with prominent tooth; branchial margin with 4 teeth; carapace posterior margin with intestinal spine. Maximum cl. 98 mm. Subtidal, shelf (1–27 m). Tropical Atlantic (Gulf of Mexico). 1 species (Rathbun 1925: rediagnosis).

Criocarcinus H. Milne Edwards, 1834 Diagnosis. Carapace broadly pyriform, longer than wide, with few dorsal tubercles. Rostrum Y-shaped, comprising 2 spines fused at base, curving and diverging abruptly. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle produced; postorbital spine slender, blunt. Hepatic margin without spine; branchial margin with 2 spines; carapace posterior margin with 1 cylindrical spine. Eyestalk extending well beyond eave. Maximum cl. 38 mm (Pl. 49b). Intertidal–shelf (0–50 m). Temperate Northern W Pacific, Central Indo-Pacific. 1 species (Sakai 1976: figure).

Crocydocinus Lee, Richer de Forges & Ng, 2019 Diagnosis. Carapace regions well defined, dorsum and legs covered in thick layer of setae, hepatic and branchial regions with conical lobes projected laterally. Rostrum comprising 2 straight, cylindrical diverging spines, one-third carapace length. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; postorbital spine slender, blunt. Maximum cl. 32 mm (Pl. 49c). Slope (350–870 m). Western and Central Indo-Pacific. 8 species (Lee et al. 2019).

Cyclocoeloma Miers, 1880 Diagnosis. Carapace oblong, with prominent transverse ridges anterior to branchial regions meeting at median tubercle. Rostrum comprising 2 short teeth fused basally. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; orbit projecting, tubular; preorbital spine blunt, with upturned lobe reaching almost level of rostrum. Antennal basal article wider than long, lateral margin rounded. Hepatic margin obtusely rounded; branchial margin smooth. Eyestalk extending well beyond eave. Maximum cl. 44 mm (Pl. 49d, e). Subtidal (1–12 m; may carry sea anemones). Central IndoPacific. 1 species (Sakai 1967: figure).

depressed spines with upcurved apices, diverging or converging, with subapical dorsal accessory spine. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; prominent, almost as wide as branchial width; preorbital tooth blunt; postorbital lobe small, rounded. Antennal basal article with lateral and anterolateral lobes or teeth. Hepatic margin smooth; branchial margin convex. Maximum cl. 21 mm. Intertidal, shelf (0–144 m). Western Indo-Pacific. 6 species (Griffin and Tranter 1986: key to 5 species; Kazmi and Tirmizi 1995).

Delsolaria Garth, 1973 Diagnosis. Carapace triangular, with few tubercles, cardiac region elevated. Rostrum as wide as long, with V-shaped notch over distal half. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital spine acute. Antennal basal article wider proximally, with strong anterolateral tooth visible in dorsal view. Branchial margin with few small teeth, 1 more prominent at posterolateral angle; carapace posterior margin convex, laminate. Eyestalk not visible in dorsal view. Maximum cl. 38 mm. Shelf (35 m). Tropical Eastern Pacific. 1 species (Garth 1973: diagnosis; Garth and Méndez 1983: male).

Doclea Leach, 1815 Diagnosis. Carapace globular, with pair of submedian tubercles behind rostrum, median row of tubercles and/or spines, diverging rows of hepatic-branchial tubercles/spines, with short dense pile. Rostrum broadly angular, or bifid. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital spine small, not expanded beyond base of rostrum, or spine absent; postorbital lobe with flanges on dorsal and ventral margins. Maxilliped 3 merus distolaterally expanded. Gonopod 1 simple. Maximum cl. 62 mm (Pl. 49f). Intertidal–shelf (0–60 m). Western and Central Indo-Pacific. 11 species (Chen and Ng 2004: review, new species; Griffin and Tranter 1986: key to 11 species, not all now accepted; Wagner 1986: rediagnosis, key to 11 species).

Epialtoides Garth, 1958

Diagnosis. Carapace subrectangular, anterolateral margin broad, convex, entire. Rostrum strongly produced laterally, concealing antennae. Eyestalk not visible in dorsal view. Pereopods 2–5 subchelate, propodi flexor margin lobate. Range uncertain, probably West Pacific. 1 species (Adams and White 1849: figure; Griffin and Tranter 1986).

Diagnosis. Carapace broad, subpentagonal or oblong, almost smooth. Rostrum oblong or tapering, with apical notch. Supraorbital eave and postorbital spine indistinguishable; supraorbital eave forming continuous rim, without or with small preorbital and postorbital teeth. Hepatic margin expanded, laminar, triangular; branchial margin expanded, laminar, triangular. Pleonites 3–5 fused in both sexes. Sexes dissimilar. Maximum cl. 17 mm. Intertidal, subtidal (0–25 m). Tropical W Atlantic, Tropical Eastern Pacific. 5 species (Garth 1958: rediagnosis, key to 3 E Pacific species).

Cyphocarcinus A. Milne-Edwards, 1868

Epialtus H. Milne Edwards, 1834

Diagnosis. Carapace ovoid, uneven, mesogastric and frontal regions usually prominently elevated. Rostrum comprising 2

Diagnosis. Carapace broad, subpentagonal or oblong, almost smooth. Rostrum broad, triangular or oblong, entire or with apical

Cyclonyx Miers, 1879

14 – Brachyura – crabs

notch. Supraorbital eave and postorbital spine indistinguishable; supraorbital eave forming continuous rim, without or with small preorbital and postorbital teeth. Hepatic margin expanded, laminar, triangular; branchial margin expanded, laminar, triangular. Pleonites 4, 5 fused in male, pleonites 3–5 fused in female. Sexes dissimilar. Maximum cl. 12 mm (Pl. 49g). Intertidal–shelf (0–54 m). Tropical W Atlantic, Tropical Eastern Pacific. 10 species (Garth 1958: rediagnosis, key to 3 E Pacific species; Rathbun 1925: key to species, includes other genera).

Eupleurodon Stimpson, 1871 Diagnosis. Carapace almost smooth. Rostrum longer than wide, with apical notch. Supraorbital eave and postorbital spine indistinguishable; supraorbital eave forming continuous rim, without preorbital and postorbital teeth. Hepatic margin with long tapering expansion; branchial margin tuberculate. Pereopods 2–5 propodi with proximal thumb opposing dactylus. Pleonites 3–5 fused in both sexes. Maximum cl. 9.5 mm. Intertidal. Tropical Eastern Pacific. 3 species (Garth 1958: rediagnosis, key to species).

Garthinia Richer de Forges & Ng, 2009 Diagnosis. Carapace triangular, flat, with tomentum of coneshaped setae giving spiny appearance extending to legs. Rostrum comprising 2 long, parallel, thick, contiguous, obtuse spines. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital tooth rounded; postorbital spine fused with hepatic margin. Hepatic margin separated from branchial margin by deep notch; branchial margin with posterolateral lobe. Maximum cl. 33 mm. Slope (314–410 m). Central Indo-Pacific. 1 species (Richer de Forges and Ng 2009a).

Giranauria Griffin & Tranter, 1986 Diagnosis. Carapace mesogastric region with 3 strong tubercles or short spines; cardiac region with high tubercle or prominent cylindrical spine. Rostrum comprising 2 spines about one-third postrostral carapace length, divergent from near base. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; supraorbital eave strongly expanded, sometimes bilobed; preorbital spine triangular; postorbital lobe truncate, with flanges on dorsal and ventral margins. Antennal basal article having anterolateral angle separated from article 2 by notch. Maximum cl. 15 mm. Subtidal, shelf (11–90 m). Western and Central Indo-Pacific. 3 species (Griffin and Tranter 1986: diagnosis, key to species).

Goniopugettia Ng, Guinot & Davie, 2008 Diagnosis. Carapace with 3 gastric, 1 cardiac bosses, 1 hepatic, 2 branchial strong transverse ridges protruding over lateral margins. Rostrum fused over basal half, 2 parallel or diverging spines distally. Supraorbital eave and postorbital spine separated by

597

U-shaped notch, antorbital angle obsolete; preorbital spine absent; postorbital spine slender, blunt. Maximum cl. 60 mm. Shelf. Temperate Northern W Pacific. 1 species (Ng et al. 2017: rediagnosis, figures).

Goniothorax A. Milne-Edwards, 1879 Diagnosis. Carapace subpentagonal, smooth. Rostrum triangular, with shallow median notch. Supraorbital eave and postorbital spine indistinguishable; preorbital spine triangular; postorbital spine absent. Hepatic margin with blunt tubercle, anteriorly directed spine; branchial margin parallel, smooth; carapace posterior margin convex. Maximum cl. 5.5 mm. Intertidal (uncertain). Western Indo-Pacific (Madagascar). 1 species (Milne-Edwards 1879: type figured on MNHN collections website).

Griffinia Richer de Forges, 1994 Diagnosis. Carapace smooth, regions not indicated covered with long upright setae; branchial margin without spine; hepatic margin forming angular lobe or anterolaterally directed spine. Rostrum comprising 2 spines separated by triangular notch. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete, or separated by U-shaped notch, antorbital angle obsolete; preorbital spine triangular; postorbital tubercle small. Male cheliped merus, carpus and propodus carinate. Gonopod 1 thin, acute, subdistally flattened. Female pleonites 5, 6 fused. Maximum cl. 20 mm. Slope (345–670 m). Temperate Northern W Pacific, Central Indo-Pacific, Temperate Australasia. 4 species (Richer de Forges 1994: key to species; Richer de Forges and Ng 2012).

Guinotinia Richer de Forges & Ng, 2009 Diagnosis. Carapace with tomentum of cone-shaped setae giving spiny appearance extending to legs, with raised plates with rounded surfaces. Rostrum bifid, comprising 2 short flat subparallel teeth. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine triangular. Hepatic margin with rounded lobe; branchial margin with 2 blunt lobes. Pereopods 3–5 pseudochelate. Maximum cl. 23 mm. Slope (500–614 m). Central Indo-Pacific. 2 species (Richer de Forges and Ng 2009a).

Herbstia H. Milne Edwards, 1834 Diagnosis. Carapace pyriform or triangular, with tubercles and spines. Rostrum comprising 2 short triangular teeth. Supraorbital eave and postorbital spine separated by intercalated spine; preorbital spine triangular, or spine absent. Antennal basal article with 1 or 2 acute lateral spines and 1 acute anterolateral spine. Maximum cl. 43 mm (Pl. 49h). Subtidal, shelf (0–92 m). Temperate Northern and Tropical Atlantic, Tropical Eastern Pacific. 11 species (Garth 1958: key to 6 E Pacific species; Manning and Holthuis 1981: key to 3 E Atlantic species; Rathbun 1925: rediagnosis, key to 7 species).

598

Marine Decapod Crustacea

Holoplites Rathbun, 1894 Diagnosis. Carapace spinose. Rostrum comprising 2 diverging spines one-third postrostral carapace length. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine half as long as rostrum. Antennal basal article expanded, with 1 long basal spine, 1 lateral tooth and 1 long anterolateral spine. Hepatic margin spinose; branchial margin spinose; carapace posterior margin with intestinal spine. Eyestalk extending well beyond eave. Maximum cl. 24 mm. Shelf, slope (160–780 m). Tropical W Atlantic. 1 species (Rathbun 1925: rediagnosis).

Hoplophrys Henderson, 1893 Diagnosis. Carapace with 15 short conical tubercles. Rostrum as wide as long, with V-shaped notch over distal half. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; preorbital spine triangular; postorbital lobe squarish. Antennal basal article narrow, with anterolateral spine, separated from postorbital lobe by shallow U-shaped hiatus. Pleonites 4–6 fused in female. Maximum cl. 17 mm (Pl. 49i). Subtidal, shelf (1–90 m; associated with alcyonarian corals including Dendronephthya). Western and Central Indo-Pacific. 1 species (Griffin and Tranter 1986: discussion; Henderson 1893: figure).

Huenia De Haan, 1837 Diagnosis. Carapace flat, smooth, sometimes with gastric and cardiac tubercles. Rostrum oblong or tapering, with apical notch, or triangular, tapering to apical tooth. Supraorbital eave and postorbital spine indistinguishable; preorbital spine triangular; postorbital spine absent. Hepatic margin expanded or not in some males; branchial margin expanded, laminar. Pereopods 2–5 flexor margin with tuft of setae and sometimes lobe in distal half; pereopod 2 propodus flexor margin with tuft of setae and sometimes lobe in distal half. Sexes dissimilar. Maximum cl. 36 mm (Pl. 49j, k). Intertidal–shelf (0–304 m; associated with macroalgae). Temperate Northern Pacific (Japan), Western and Central IndoPacific, Temperate South America (Salas y Gomez I.), Temperate Australasia (Australia). 10 species (Griffin and Tranter 1986: key to 8 species; Takeda and Marumura 2010: review, 2 species described).

Hyastenus White, 1847 Diagnosis. Carapace surface with few tubercles and short spines, with epibranchial and intestinal spines. Rostrum comprising 2 spines one-fifth to more than carapace length, rarely united in basal third. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital spine triangular, or tooth rounded; postorbital lobe with flanges on dorsal and ventral margins. Antennal basal article without longitudinal groove, lateral marginal lobe, if present not extending dorsally into orbit, separated from postorbital lobe by narrow hiatus, with or without small suborbital tubercle Maximum cl. 53 mm (Pl. 49l).

Intertidal–slope (0–450 m) Indo-West Pacific, Temperate ­ ustralia. 37 species (Griffin and Tranter 1986: rediagnosis, key to A 30 species, descriptions of 25 species; Lee and Ng 2020; Windsor and Ahyong 2013: key to 31 species).

Lahaina Dana, 1851 Diagnosis. Carapace surface with few tubercles and short spines, with epibranchial and intestinal spines. Rostrum comprising 2 sinuous spines as long as carapace. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital spine acute; postorbital lobe small, laterally flattened, sometimes with suborbital tubercle. Antennal basal article narrow; with distinct longitudunal groove; with strong anterolateral spine, lateral margin at base sometimes produced as dorsally extending lobe, separated from postorbital lobe by broad hiatus, with small suborbital tubercle. Pereopods 2, 3 meri with terminal spine. Maximum cl. 27 mm. Subtidal, shelf (1–150 m). Indo-West Pacific. 5 species (Griffin and Tranter 1986: rediagnosis, key and descriptions of 4 species).

Laubierinia Richer de Forges & Ng, 2009 Diagnosis. Carapace ovoid, with several strong elevated swellings, some flat; hepatic region particularly elevated; covered with tomentum. Rostrum comprising 2 short triangular teeth. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital spine triangular. Hepatic margin with round flat plate; branchial margin with 1 or 2 flat plates separated by groove from rest of carapace. Maximum cl. 27 mm. Shelf, slope (173–439 m). Central Indo-Pacific. 2 species (Richer de Forges and Ng 2009a).

Lepidonaxia Targioni-Tozzetti, 1877 Diagnosis. Carapace ovoid, evenly tapering anteriorly, sparsely tuberculate. Rostrum comprising 2 flat diverging spines, one-fifth postrostral carapace length. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; supraorbital eave aligned with lateral margin; preorbital tooth blunt. Antennal basal article with blunt lateral and anterolateral teeth. Hepatic margin smooth; branchial margin with 1 tooth. Eyestalk not visible in dorsal view. Maximum cl. 22 mm. Subtidal (?). Central Indo-Pacific (Java). 1 species (Targioni Tozzetti 1877). Lepidonaxia defilippi Targioni Tozzetti, 1877 has not been recorded since its discovery.

Lepteces Rathbun, 1893 Diagnosis. Carapace with rows of large mushroom-shaped tubercles, interspersed with cylindrical spines. Rostrum fused over basal half, 2 parallel or diverging spines distally. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital spine curved upwards. Pereopods 2–5 spinose; pereopod 1 longest. Maximum cl. 17 mm. Slope (237–475 m). Tropical Atlantic (Gulf of Mexico). 1 species (Rathbun 1925: rediagnosis).

14 – Brachyura – crabs

Leptomaia Griffin & Tranter, 1986 Diagnosis. Carapace ovoid, orbits almost as wide as maximum carapace width, with many low tubercles. Rostrum comprising 2 diverging spines one-quarter carapace length. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; preorbital spine triangular. Antennal basal article narrow, lateral margin with broad lobe, with strong anterolateral spine. Hepatic margin with 2 tubercles. Maximum cl. 15 mm. Subtidal, shelf (14–85 m). Temperate Australasia (E Australia, New Zealand). 1 species (Griffin and Tranter 1986).

Leptopisa Stimpson, 1871 Diagnosis. Carapace uneven, with pair of epibranchial spines. Rostrum comprising 2 long spines, contiguous at midpoint, apices slightly diverging. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; orbit projecting, tubular; preorbital spine triangular. Antennal basal article expanded, with anterolateral spine not visible in dorsal view, separated from postorbital spine by slit. Maximum cl. 22 mm (Pl. 50a). Subtidal, shelf (1–80 m). Tropical W Atlantic, Central IndoPacific. 3 species (Griffin and Tranter 1986: key to 3 species; Rathbun 1925: rediagnosis).

Leucippa H. Milne Edwards, 1833

599

antorbital angle squarish; preorbital spine acute; postorbital lobe squarish. Hepatic margin with 1 or 2 spines; branchial margin with 3 or mores spines; carapace posterior margin with intestinal spine. Male sterno-abdominal cavity deep, narrow, closed anteriorly by crest on thoracic sternite 4, with lateral tubercles. Gonopod 1 bifid. Maximum cl. 107 mm (Pl. 50b). Subtidal, shelf (1–124 m). Temperate Northern and Tropical Atlantic, Temperate South America (Atlantic coast), Eastern Indo-Pacific. 10 species (Tamburus and Mantelatto 2021: rediagnosis, key to species).

Lissa Leach, 1815 Diagnosis. Carapace diamond-shaped, widest posteriorly, with regions well defined, with 10 large bosses. Rostrum comprising 2 contiguous square plates, depressed, with concave apex. Supraorbital eave and postorbital spine indistinguishable; preorbital spine triangular; postorbital tooth merged with hepatic flange. Hepatic margin with concave flange; branchial margin with 2 rounded lobes, second larger. Pereopods 2–5 tuberculate. Pleonites free in both sexes. Maximum cl. 51 mm. Shelf (20–200 m). Mediterranean Sea only. 1 species (Pretzmann 1961: rediagnosis; Zariquiey Alvarez 1968: rediagnosis, figure).

Lophorochinia Garth, 1969

Diagnosis. Carapace subpentagonal, smooth. Rostrum lamellate, lateral margins convex, with median slit. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine small, not expanded beyond base of rostrum. Hepatic margin smooth; branchial margin expanded as triangular lamina. Pereopods 2–5 meri carinate. Pleonites free in both sexes. Maximum cl. 23 mm. Subtidal, shelf (13–100 m). Tropical W Atlantic, Tropical Eastern Pacific, Temperate South America (Pacific coast). 1 species (Garth 1958: rediagnosis; Milne Edwards 1834: figure).

Diagnosis. Carapace subcircular, branchial regions inflated, meeting but not fusing on midline; cardiac region obscured. Rostrum comprising 2 short spines fused basally. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine triangular. Hepatic margin with prominent tooth; branchial margin with lateral tooth. Pleonites free in both sexes. Maximum cl. 66 mm. Slope (282–509 m). Temperate South America (Chile). 1 species (Garth 1969; Garth and Haig 1971: figure).

Libidoclaea H. Milne Edwards & Lucas, 1842

Loxorhynchus Stimpson, 1857

Diagnosis. Carapace broadly pyriform, as long as wide, with tubercles and spines. Rostrum tapering, with 2 diverging apical spines very short or almost as long as basal part. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine acute. Antennal basal article lateral margin with tooth and anterolateral spine. Hepatic margin with prominent spine or tooth; branchial margin with 3 spines, second longest; carapace posterior margin with intestinal spine. Gonopod 1 bifid. Maximum cl. 67 mm. Subtidal–slope (18–1921 m). Temperate South America (Chile). 2 species (Rathbun 1925: rediagnosis, key to species).

Diagnosis. Carapace broadly pyriform, longer than wide, with tubercles and spines. Rostrum fused over basal half, 2 parallel or diverging spines distally. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; preorbital spine acute. Antennal basal article with anterolateral tooth visible in dorsal view, lateral margin with 2 teeth. Hepatic margin with 1 or 2 spines; branchial margin spinose. Maximum cl. 200 mm. Subtidal, shelf (7–135 m). Tropical Eastern Pacific. 3 species (Hendrickx and Cervantes 2003: comparison of species; Rathbun 1925: rediagnosis, key to 2 species).

Libinia Leach, 1815 Diagnosis. Carapace globular or ovoid (rare), with tubercles and spines. Rostrum simple, apex emarginate or bifid. Supraorbital eave and postorbital spine separated by U-shaped notch,

Lyramaia Griffin & Tranter, 1986 Diagnosis. Carapace surface with many small tubercles and few short spines. Rostrum lyre-shaped (lateral edges convex proximally, spines parallel to divergent distally, apices curved outwards), basally fused, one-third-half carapace length. Supraorbital

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eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital tooth rounded; postorbital lobe with flanges on dorsal and ventral margins. Maximum cl. 14 mm. Shelf (75–79 m). Temperate E Australia. 1 species (Griffin and Tranter 1986).

margin tuberculate; branchial margin tuberculate. Maximum cl. 36 mm (Pl. 50e). Intertidal–shelf (0–100 m). Indo-West Pacific, Temperate ­Australia, M. thalia (Herbst, 1803) exotic in Mediterranean. 10 species (Griffin and Tranter 1986: key to 10 species).

Macrocoeloma Miers, 1879

Micippoides A. Milne-Edwards, 1873

Diagnosis. Carapace pyriform or triangular, with tubercles and spines or smooth. Rostrum comprising 2 flat diverging spines, one-fifth postrostral carapace length. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; orbit projecting, tubular; preorbital spine triangular, or tooth blunt. Antennal basal article expanded, with anterolateral spine visible in dorsal view, separated from postorbital spine by slit. Pterygostomial region with short spines not visible in dorsal view; hepatic margin smooth; branchial margin with 1 or more prominent spines, or smooth. Maximum cl. 43 mm (Pl. 50c). Intertidal–slope (0–298 m). Tropical W Atlantic, Tropical Eastern Pacific. 12 species (Garth 1940: species description, figure; Rathbun 1925: rediagnosis, key to 11 species).

Diagnosis. Carapace with swollen triangular branchial plates; pereopods and carapace covered with long soft setae. Rostrum comprising 2 deflexed triangular spines, apices upcurved, onesixth carapace length. Supraorbital eave and postorbital spine separated by suture, antorbital angle square. Antennal basal article broad, tapering. Maximum cl. 12 mm. Intertidal, subtidal (0–5 m). Indo-West Pacific. 1 species (Griffin and Tranter 1986: discussion; Rathbun 1911: figure).

Menaethiops Alcock, 1895 Diagnosis. Carapace sparsely tuberculate. Rostrum comprising 2 spines, contiguous or diverging, or single spine. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; preorbital spine absent, or spine acute; postorbital lobe fused to supraorbital eave, sometimes expanded and distally in contact with hepatic margin. Antennal basal article with distolateral spine often visible in dorsal view. Hepatic margin tuberculate, or with small spines; branchial margin tuberculate, or with small spines. Eyestalk extending well beyond eave. Maximum cl. 14 mm. Intertidal–shelf (0–69 m). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate Southern E Africa. 16 species (Griffin and Tranter 1986: key to 12 Indo-West Pacific species; Naderloo 2015: species description, figure).

Menaethius A. Milne-Edwards, 1834 Diagnosis. Carapace pyriform or ovoid, with few tubercles, cardiac region elevated. Rostrum triangular, tapering to apical tooth. Supraorbital eave and postorbital spine indistinguishable; preorbital spine triangular; postorbital spine absent. Hepatic margin with small lateral lobe just behind eye; branchial margin with 2 lobes, sometimes obsolete. Eyestalk extending well beyond eave. Maximum cl. 26 mm (Pl. 50d). Intertidal–shelf (0–72 m). Indo-West Pacific. 2 species (Griffin and Tranter 1986: discussion).

Micippa Leach, 1817 Diagnosis. Carapace oblong-ovate, with tubercles and spines. Rostrum of 2 broad, flat plates, depressed, fused for at least proximal two-thirds. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; preorbital tooth blunt. Hepatic

Microlissa Pretzmann, 1961 Diagnosis. Carapace triangular, with regions well defined, with 10 large bosses. Rostrum rectangular, truncate. Supraorbital eave and postorbital spine indistinguishable; preorbital tooth blunt; postorbital lobe small, rounded. Hepatic margin smooth; branchial margin with posterolateral lobe. Eyestalk not visible in dorsal view. Pereopods 2–5 meri carinate. Pleonites 3–5 fused in both sexes. Maximum cl. 25 mm (Pl. 50f). Subtidal, shelf (4–61 m). Tropical Atlantic, Tropical Eastern Pacific. 5 species (Pretzmann 1961: key to species with Lissa).

Micropisa Stimpson, 1858 Diagnosis. Carapace with tubercles and spines. Rostrum comprising 2 short spines fused basally. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital spine half as long as rostrum; postorbital lobe with flanges on dorsal and ventral margins. Hepatic margin with 1 spine; branchial margin with lateral tooth. Maximum cl. 6 mm. Intertidal–shelf (0–110 m). Tropical E Atlantic. 1 species (Monod 1956: description, figures).

Minyorhyncha Tavares & Santana, 2018 Diagnosis. Carapace subcircular, vaulted, with 6 mesial spines, with short branchial spines. Rostrum comprising 2 short parallel spines as adult, diverging as juvenile. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine acute. Hepatic margin with 1 spine; branchial margin spinose. Maximum cl. 86 mm (Pl. 50h). Shelf, slope (66–1216 m). Temperate Northern and Tropical W Atlantic. 1 species (Tavares and Santana 2018).

Mocosoa Stimpson, 1871 Diagnosis. Carapace subpentagonal, tumid, regions with 8 prominent bosses. Rostrum as long as wide, with apical notch. Supraorbital eave and postorbital spine indistinguishable; supraorbital

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eave forming continuous rim, without preorbital and postorbital teeth. Hepatic margin obtusely rounded; branchial margin obtusely rounded. Eyestalk short, immovable. Pleonites 3–5 fused in both sexes. Maximum cl. 7 mm. Shelf (20–131 m). Tropical W Atlantic. 1 species (Rathbun 1925: rediagnosis).

Naxioides A. Milne-Edwards, 1865 Diagnosis. Carapace spinose. Rostrum comprising 2 spines with dorsal accessory spine in distal half. Supraorbital eave and postorbital spine separated by intercalated spine. Maximum cl. 73 mm (Pl. 50g). Intertidal–slope (0–400 m). Indo-West Pacific, Temperate Australasia. 10 species (Griffin and Tranter 1986: key to 6 species; Poupin 1995: key to 3 species of N. robillardi group; Takeda et al. 2022: species comparisons).

Neodoclea Buitendijk, 1950 Diagnosis. Carapace globular, tomentose, with few scarcely visible gastric and branchial tubercles or spines. Rostrum simple, not projected beyond orbit. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine absent. Pterygostomial region with strong spine exceeding front; branchial margin with 3 spines, first shortest; carapace posterior margin with intestinal spine. Maxilliped 3 merus narrower than ischium, tapering. Gonopod 1 bifid. Maximum cl. 37 mm. Subtidal (9–20 m). Tropical Eastern Pacific. 1 species (­Buitendijk 1950; Holthuis 1966: redescription).

Neophrys Lee, Richer de Forges & Ng, 2019 Diagnosis. Carapace regions defined by long spines, covered with thin layer of setae. Rostrum comprising 2 straight, spines diverging at 30°, as long as carapace. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine obsolete; postorbital tooth merged with hepatic flange. Hepatic margin with prominent oblique spine; branchial margin with 1 spine directed laterally, as long as hepatic spine. Maximum cl. 27 mm (Pl. 50i). Slope (434–590 m). Central Indo-Pacific. 1 species (Lee et al. 2019).

Nibilia A. Milne-Edwards, 1878 Diagnosis. Carapace with tubercles and spines. Rostrum comprising 2 spines about one-third postrostral carapace length, divergent from near base. Supraorbital eave and postorbital spine separated by intercalated spine; preorbital spine acute. Antennal basal article expanded, with 1 lateral and 1 similar long anterolateral spine. Hepatic margin with 1 or 2 spines; branchial margin spinose. Maximum cl. 120 mm. Shelf, slope (55–342 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Colavite et al. 2019: rediagnosis, second species).

601

Nicoya Wicksten, 1987 Diagnosis. Carapace evenly tuberculate, without spines. Rostrum comprising 2 short teeth fused basally. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine absent. Antennal basal article with blunt anterolateral tooth visible in dorsal view, lateral margin tuberculate. Hepatic margin tuberculate; branchial margin tuberculate. Maximum cl. 7.4 mm. Shelf (48 m). Tropical Eastern Pacific. 1 species (Wicksten 1987).

Notolopas Stimpson, 1871 Diagnosis. Carapace smooth, with cardiac, intestinal and pair of prominent branchial spines. Rostrum fused over basal half, 2 parallel or diverging spines distally. Supraorbital eave and postorbital spine separated by intercalated spine; preorbital spine triangular. Antennal basal article wider proximally, with tubercle next to postorbital lobe. Hepatic margin without spine; branchial margin convex. Maximum cl. 17 mm. Intertidal–shelf (0–101 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 3 species (Garth 1958: figures; Rathbun 1925: rediagnosis, key to 2 species).

Oplopisa A. Milne-Edwards, 1879 Diagnosis. Carapace with few dorsal tubercles. Rostrum comprising 2 diverging spines one-third postrostral carapace length. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; preorbital spine triangular. Antennal basal article with lateral and anterolateral lobes or teeth. Hepatic margin expanded, laminar, triangular; branchial margin regularly tuberculate. Eyestalk extending well beyond eave. Pereopods 2–5 spinose; pereopod 1 longest. Maximum cl. 12.5 mm. Shelf (185 m). Tropical W Atlantic. 1 species (Rathbun 1925: rediagnosis).

Oxypleurodon Miers, 1885 Diagnosis. Carapace triangular or ovoid, convex, with prominent raised convex plates divided by deep grooves (1 gastric, 1 cardiac, 1 intestinal, pairs of hepatic and branchial - all sometimes subdivided). Rostrum comprising 2 contiguous spines. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine absent, or tooth blunt. Hepatic margin flat; branchial margin with plate, sometimes acute, swollen. Maximum cl. 25 mm (Pl. 50j). Shelf, slope (70–1076 m). Indo-West Pacific. Temperate SW Australia. 33 species (Lee et  al. 2019; Richer de Forges and Ng 2009b: rediagnosis, 23 species listed, plus 4 in Nasutocarcinus Tavares, 1991, now a junior synonym; Tavares 1991).

Paranaxia Rathbun, 1924 Diagnosis. Carapace pyriform or triangular, tomentose, with hepatic and larger epibranchial spine. Rostrum comprising 2 parallel spines, one-third postrostral carapace length, apices with accessory

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Marine Decapod Crustacea

mesial tooth. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital spine acute. Antennal basal article broad, without teeth. Hepatic margin obtusely rounded; branchial margin convex. Maximum cl. 116 mm. Subtidal, shelf (0–30 m). Central Indo-Pacific, Temperate SW Australia. 2 species (Hosie and Hara 2016). Paranaxia is unique among spider crabs in exhibiting direct development, where the young hatch as first stage crabs and are retained under the female’s pleon for at least one moult cycle (Morgan 1987).

Pelia Bell, 1836 Diagnosis. Carapace smooth, sparsely setose. Rostrum comprising 2 short teeth fused basally. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine absent. Antennal basal article narrow, with anterolateral tooth visible in dorsal view. Hepatic margin smooth; branchial margin convex. Pereopods 2–5 meri carinate. Maximum cl. 19 mm (Pl. 50k). Subtidal, shelf (1–100 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 6 species (Garth 1958: key to 3 E Pacific species; Milne-Edwards 1873–1881: figure; Rathbun 1925: rediagnosis, key to 5 species).

Perinia Dana, 1851 Diagnosis. Carapace subcircular, with few large tubercles. Rostrum comprising 2 small lobes separated by U-shaped notch. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; supraorbital eave forming continuous rim around eye without preorbital or postorbital spines; eye retractile against rim. Pereopods 2–5 meri with 2 sharp teeth or lobes on extensor margin, tubercles on outer face; pereopods 2, 3 meri with 2 blade-like lobes or high tubercles on extensor margin, tubercles on outer face. Maximum cl. 7.7 mm. Intertidal–shelf (0–40 m). Indo-West Pacific. 2 species (Dai et al. 1994: comparison of 2 species; Sakai 1938: figure).

Phalangipus Latreille, 1828 Diagnosis. Carapace subcircular, with tubercles and spines. Rostrum comprising 2 short spines fused basally. Supraorbital eave and postorbital spine separated by intercalated spine; preorbital spine absent. Pereopods 2–5 slender, up to 4 times carapace length. Maximum cl. 35 mm. Subtidal–slope (5–353 m). Western and Central Indo-Pacific, Temperate Australia. 9 species (Griffin 1973: revision; Griffin and Tranter 1986: key to 9 species).

Picroceroides Miers, 1886 Diagnosis. Carapace ovoid, with 1 gastric, 1 cardiac and 1 intestinal spine. Rostrum comprising 2 widely divergent spines, more than half postrostral length. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; orbit tubular, exceeding carapace width; preorbital spine angled dorsally, half as long as rostrum; postorbital spine acute. Antennal basal article

expanded forming floor of orbit. Hepatic margin obtusely rounded; branchial margin convex. Pereopods 2–5 slender, up to 4 times carapace length. Maximum cl. 23 mm. Subtidal, shelf (1–110 m). Tropical W Atlantic. 1 species (Rathbun 1925: rediagnosis).

Picrocerus A. Milne-Edwards, 1865 Diagnosis. Carapace with few dorsal spines. Rostrum comprising 2 long spines, contiguous at midpoint, apices slightly diverging. Supraorbital eave and postorbital spine fused; orbit tubular, exceeding carapace width; preorbital spine absent; postorbital spine attenuated. Hepatic margin with 2 long diverging spines; branchial margin with 2 spines; carapace posterior margin with 2 short teeth. Eyestalk extending well beyond eave. Maximum cl. 62 mm (Pl. 50l). Subtidal, shelf (1–50 m). Temperate Northern W Pacific, Central Indo-Pacific. 1 species (Sakai 1976: figure).

Pisa Leach, 1814 Diagnosis. Carapace pyriform or triangular, with regions well defined, each with single spine. Rostrum comprising 2 spines onefifth to more than carapace length, rarely united in basal third. Supraorbital eave and postorbital spine separated by intercalated spine; preorbital spine acute. Antennal basal article with blunt lateral and anterolateral teeth, separated from postorbital spine by deep U. Hepatic margin with 1 or more tubercles; branchial margin tuberculate, or spinose. Maximum cl. 53 mm. Intertidal–shelf (0–162 m). Temperate Northern and Tropical E Atlantic, including Mediterranean. 9 species (Forest and Guinot 1966: descriptions of 3 species; Manning and Holthuis 1981: discussion of 5 species; Zariquiey Alvarez 1968: rediagnosis, key to 6 species).

Pisoides H. Milne Edwards & Lucas, 1843 Diagnosis. Carapace smooth. Rostrum comprising 2 teeth fused over basal half, one-fifth carapace length. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; preorbital spine absent. Antennal basal article little longer than wide, with rounded anterolateral lobe. Hepatic margin smooth; branchial margin smooth, slightly convex. Pereopods 2–5 meri carinate. Maximum cl. 24 mm. Shelf (0–70 m). Tropical Eastern Pacific, Temperate South America (Chile). 1 species. The two species from Japan, P.  ortmanni (Balss, 1924) and P. bidentatus (A. Milne Edwards, 1873), included by Sakai (1976) have a tuberculate carapace and a strong preorbital spine. Griffin and Tranter (1986) noted that their gonopod 1 differed from that of the type species. They are not encompassed by this generic diagnosis.

Pliosoma Stimpson, 1860 Diagnosis. Carapace ovoid, inflated, regions marked, with 4 erect, curved medial blunt spines, pair on branchial regions. Supraorbital eave and postorbital spine separated by suture, antorbital angle

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603

square; tubular, directed anteriorly. Hepatic margin with 3 blunt teeth; branchial margin with 2 teeth; carapace posterior margin convex. Pereopods 2–5 diverse, pereopod 2 slender, longest; pereopods 4, 5 short, setose, with flat distal articles. Maximum cl. 20 mm. Subtidal (11–22 m). Tropical Eastern Pacific. 1 species (Rathbun 1923).

U-shaped notch, antorbital angle obsolete; preorbital spine triangular. Hepatic margin obtusely rounded; branchial margin with 2 triangular lobes, second longer. Eyestalk extending well beyond eave. Maximum cl. 39 mm. Subtidal, shelf (1–170 m). Tropical Eastern Pacific. 1 species (Richer de Forges and Ng 2009b).

Pohleus Colavite, Windsor & Santana, 2020

Rochinia A. Milne-Edwards, 1875

Diagnosis. Carapace triangular (wider than long), tomentose, with 7 strong spines in posterior half. Rostrum Y-shaped, comprising 2 spines fused at base, curving and diverging abruptly. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; orbit projecting, tubular; preorbital spine acute; postorbital spine acute. Antennal basal article with 3 ventrally directed spines, lateral one longest, 1 tubercle, not visible in dorsal view. Pterygostomial region with strong spine visible in dorsal view. Maximum cl. 35 mm. Subtidal, shelf (9–117 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Colavite et al. 2020).

Diagnosis. Carapace without long dorsal spines or covering of large nodules, at most with low protuberances; rostrum comprising 2 spines or lobes; supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; preorbital spine acute; postorbital spine angular in lateral view; pterygostomial region with low teeth; hepatic margin with prominent laterally directed spine or tooth; branchial margin with 1 or more prominent spines. Male pleon acutely triangular to T-shaped, pleonites 3–6 trapezoidal. Gonopod 1 with oblique apex. Maximum cl. 24 mm. Shelf (21–101 m). Tropical W Atlantic, 1 species (Griffin and Tranter 1986: rediagnosis, key to 29 Indo-West Pacific species, all except one now in other genera; Lee et  al. 2017: diagnoses of related genera; 2019: discussion; Ng and Richer de Forges 2013: problems with diagnosis, species groups; Tavares and Santana 2018: description of only species. After considerable revision, most species of this once large genus (Griffin and Tranter 1986) have been removed to other genera (Lee et al. 2017, 2019, 2021; Ng and Richer de Forges 2013; Tavares and Santana 2018). Four Atlantic and Eastern Pacific species, not congeneric, remain to be placed [R. cornuta (Rathbun, 1898); R. hystrix (Stimpson, 1871); R. occidentalis (Faxon, 1893); R. tanneri (Smith, 1883)] (Lee et al. 2021). They will key to the Indo-Pacific genus Samadinina and are excluded from Rochinia. Rochinia is restricted here to its type species, R. gracilipes A. Milne-Edwards, 1875.

Pteromaja Ng & Anker, 2014 Diagnosis. Carapace with prominent hepatic and branchial wings, much wider than long, with medial vertical spine behind eyes. Rostrum broadly bilobed, short, deflexed, not visible in dorsal view. Supraorbital eave and postorbital spine indistinguishable; preorbital spine absent; postorbital spine absent. Maximum cl. 8 mm (Pl. 51a). Subtidal, shelf (6–50 m; associated with red algae, coral). Central Indo-Pacific (Papua New Guinea). 1 species (Ng and Anker 2014).

Pugettia Dana, 1851 Diagnosis. Carapace pyriform or ovoid, smooth, sparsely setose. Rostrum comprising 2 spines separated by triangular notch, lateral margins almost parallel. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; preorbital spine triangular; postorbital tooth commonly formed by the anterior angle of the hepatic expansion. Hepatic margin expanded; branchial margin laterally produced, sometimes with spine, separated from hepatic lobe by concavity. Pleonites free in both sexes. Gonopod 1 acute, with 2 subapical sharp flanges. Maximum cl. 29 mm. Subtidal–slope (0–600 m; associated with macroalgae). Temperate Northern W Pacific, Central Indo-Pacific. 24 species (Garth 1958: rediagnosis, key to 6 E Pacific species; Griffin and Tranter 1986: key to 12 Indo-West Pacific species; Ohtsuchi et al. 2020: key to 7 E Asian shallow-water species; Sakai 1976: key to 10 Japanese species).

Rhinocarcinus Richer de Forges & Ng, 2009 Diagnosis. Carapace triangular, with median ridge, few tubercles. Rostrum comprising 2 long, parallel, thick, contiguous, obtuse spines. Supraorbital eave and postorbital spine separated by

Samadinia Ng & Richer de Forges, 2013 Diagnosis. Carapace regions indicated, smooth, prominently granular or nodular, setose or glabrous; rostrum comprising 2 widely divergent spines, more than half postrostral length; supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; preorbital spine acute; postorbital spine merged with hepatic flange, anteriorly rounded in lateral view; hepatic margin with prominent oblique spine or lobe, upright or inclined dorsally; branchial margin with spines, large tubercles or both. Male pleon broadly triangular, pleonites 3, 4 trapezoidal, pleonites 5, 6 subrectangular. Gonopod 1 with oblique apex. Maximum cl. 25 mm (Pl. 51b). Slope–bathyal (165–3109 m). Central and Eastern Indo-Pacific. 29 species (Ng and Richer de Forges 2013); Lee et al. 2021). Four Atlantic–East Pacific species formerly included in Rochinia will key here (see Rochinia, above).

Sargassocarcinus Ward, 1936 Diagnosis. Carapace hexagonal, wider than long, flat, flat. Rostrum comprising 2 short triangular teeth. Supraorbital eave and

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Marine Decapod Crustacea

postorbital spine indistinguishable; preorbital spine prominent, as long as rostrum; postorbital spine absent. Hepatic margin laterally produced, expanded in both sexes, separated from similar expanded branchial lobe by shallow concavity or fissure; branchial margin expanded, laminar. Pereopods 2–5 meri carinate. Maximum cl. 10.5 mm. Subtidal, shelf (15–30 m; associated with macroalgae). Temperate Northern W Pacific, Central Indo-Pacific. 3 species (Sakai 1965: discussion, figures).

Scyra Dana, 1851 Diagnosis. Carapace with prominent transverse ridges anterior to branchial regions meeting at median tubercle. Rostrum comprising 2 leaf-shaped flat spines, fused in basal half, laterally expanded, with sharp apex. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine acute. Antennal basal article broad, tapering. Maximum cl. 46 mm (Pl. 51c). Intertidal–slope (0–304 m). Temperate Northern Pacific, Tropical Eastern Pacific. 3 species (Garth 1958: rediagnosis; Sakai 1976: Japanese species).

Scyramathia A. Milne-Edwards, 1880 Diagnosis. Carapace with dorsal tubercles becoming large, flattopped, leaf-like plates, wider than stem in adults; rostrum comprising 2 widely divergent spines, more than half postrostral length; supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; preorbital spine acute; postorbital spine merged with hepatic flange, anteriorly rounded in lateral view; pterygostomial region with large, flattened granules on outer margin; hepatic margin with prominent upright spine or tooth; branchial margin with lateral tooth. Gonopod 1 with oblique apex. Maximum cl. 41 mm. Shelf–slope (60–1340 m). Temperate Northern Atlantic, including Mediterranean, Tropical W Atlantic, Temperate Southern Africa. 4 species (Lee et al. 2020: rediagnosis, key to species, species descriptions; Tavares and Santana 2018).

Siderochinia Lee, Richer de Forges & Ng, 2021 Diagnosis. Carapace cardiac region produced as tall, thick, blunt, peg-like dorsal projection. Carapace margins with long hepatic, branchial and intestinal spines, inclined dorsad; surface smooth, cardiac region produced as tall, thick, blunt, peglike dorsal projection; rostrum comprising 2 widely divergent spines, about half postrostral length; supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; preorbital angle acute; postorbital spine merged with hepatic flange, anteriorly rounded in lateral view. Male pleon broadly triangular, pleonites 3, 4 trapezoidal, pleonites 5, 6 subrectangular. Gonopod 1 with oblique apex. Maximum cl. 25 mm. Shelf, slope (100–220 m). Central Indo-Pacific. 2 species (Lee et al. 2021: comparison of species)

Simocarcinus Miers, 1879 Diagnosis. Carapace triangular, smooth. Rostrum simple, onefifth to as long as postorbital carapace length, subrectangular to triangular. Supraorbital eave and postorbital spine indistinguishable; preorbital spine absent; postorbital spine absent. Hepatic margin with lobe in some females; branchial margin expanded, laminar. Eyestalk short, immovable. Maximum cl. 25 mm (Pl. 51d). Intertidal. Western and Central Indo-Pacific. 9 species (Griffin and Tranter 1986: key to 3 species; Tirmizi 1978: figure).

Solinca Colavite, Windsor & Santana, 2019 Diagnosis. Carapace subcircular, spinose. Rostrum comprising 2 teeth fused over basal third, one-fifth carapace length. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; preorbital spine triangular; postorbital spine long, curved. Hepatic margin with small spines; branchial margin with small spines. Maximum cl. 40 mm. Subtidal–slope (13–311 m). Tropical Eastern Pacific. 1 species (Colavite et al. 2019).

Sphenocarcinus A. Milne-Edwards, 1875 Diagnosis. Carapace triangular, convex, regions with prominent raised flat plates divided by deep grooves (1 gastric, 1 cardiac, 1 intestinal, pairs of protogastric, hepatic and branchial). Rostrum comprising 2 long spines, contiguous or slightly diverging. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine absent. Branchial margin with triangular posterolateral angle. Maximum cl. 16 mm. Shelf, slope (170–300 m). Temperate Northern and Tropical W Atlantic. 1 species (Richer de Forges and Ng 2009b: rediagnosis; Tavares 1991).

Stegopleurodon Richer de Forges & Ng, 2009 Diagnosis. Carapace with large plate arching beyond lateral margins, roof-like, surface corrugated, with median ridge. Rostrum short, comprising 2 flat, apically round teeth separated by narrow fissure. Supraorbital eave and postorbital spine separated by V-shaped notch, antorbital angle obsolete; preorbital spine absent; postorbital tooth merged with hepatic flange. Maximum cl. 19 mm (Pl. 51e). Slope (260–669 m). Central Indo-Pacific. 2 species (Lee et al. 2019; Richer de Forges and Ng 2009b).

Stenocionops Desmarest, 1823 Diagnosis. Carapace with tubercles and spines. Rostrum comprising 2 diverging spines one-quarter or more postrostral carapace length. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; preorbital spine acute. Hepatic margin with 1 or 2 spines; branchial margin with 3 or mores spines. Maximum cl. 147 mm.

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Subtidal–slope (2–508 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 8 species (Colavite et  al. 2016; Rathbun 1925: rediagnosis, key to 8 species).

Stilbognathus von Martens, 1866 Diagnosis. Carapace broadly pyriform, longer than wide, with prominent transverse ridges anterior to branchial regions meeting at median tubercle. Rostrum comprising 2 spines or lobes. Supraorbital eave and postorbital spine indistinguishable; preorbital spine prominent, as long as rostrum; postorbital spine absent. Hepatic margin with small spine; branchial margin with 1 anterior spine; carapace posterior margin with broad-based medial lobe or spine. Maximum cl. 32 mm. Subtidal, shelf (1–63 m). Indo-West Pacific. 7 species (Griffin and Tranter 1974: figure; 1986: key to species).

Stilbomastax Williams, Shaw & Hopkins, 1977 Diagnosis. Carapace oblong-ovate, with prominent transverse ridges anterior to branchial regions meeting at median tubercle. Rostrum comprising 2 spines or lobes. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; preorbital spine triangular. Branchial margin convex; carapace posterior margin with 2 broad lobes. Maxilliped 3 exopod with basal oblique process overlapping sternite. Maximum cl. 28 mm. Subtidal, shelf (15–38 m). Tropical W Atlantic. 1 species (Garth 1952: figure of Stilbomastax margaritifera (Monod, 1939) as Stilbognathus burryi Garth, 1952; Williams et al. 1977).

Stratiolibinia Tavares & Santana, 2011 Diagnosis. Carapace globular, with tubercles and spines. Rostrum broad, basally fused, ending in 2 widely divergent spines. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; preorbital spine acute; postorbital lobe squarish. Hepatic margin with 1 or 2 spines; branchial margin with 1 spine directed laterally; carapace posterior margin with intestinal spine. Male sterno-abdominal cavity deep, narrow, closed anteriorly by crest on thoracic sternite 4, without lateral tubercles. Gonopod 1 bifid. Maximum cl. 74 mm (Pl. 51f). Subtidal, shelf (4–100 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Tavares and Santana 2011).

Taliepus A. Milne-Edwards, 1878 Diagnosis. Carapace subcircular, smooth. Rostrum triangular, with shallow median notch. Supraorbital eave and postorbital spine indistinguishable; preorbital spine small, not expanded beyond base of rostrum, or spine absent; postorbital spine acute. Hepatic margin with or without tooth medial to anterolateral spine; branchial margin convex. Pleonites free in both sexes. Maximum cl. 108 mm. Subtidal, shelf (0–100 m). Eastern Indo-Pacific, Temperate South America (Pacific coast). 3 species (Garth 1958: rediagnosis, key to 3 species).

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Thersandrus Rathbun, 1897 Diagnosis. Carapace ovoid, flat, covered with long setae extending on to pereopods. Rostrum comprising 2 short triangular teeth. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; orbit oblique-transverse; preorbital spine absent. Antennal basal article broad, without teeth. Hepatic margin obtusely rounded; branchial margin convex. Pereopods 2–5 flattened, prehensile. Maximum cl. 23 mm. Subtidal (1–5 m). Tropical W Atlantic. 1 species (Colavite et al. 2020: systematic position; Rathbun 1925: rediagnosis).

Thusaenys Griffin & Tranter, 1986 Diagnosis. Carapace mesogastric region smooth or with 1–3 low tubercles; cardiac region smooth or with 1 low tubercle. Rostrum comprising 2 spines about one-third postrostral carapace length, divergent from near base. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle squarish; supraorbital eave strongly expanded, sometimes bilobed; preorbital spine triangular; postorbital lobe with flanges on dorsal and ventral margins. Antennal basal article having anterolateral spine separated from article 2 by notch. Maximum cl. 17.5 mm. Shelf (25–50 m). Western and Central Indo-Pacific. 6 species (Chen et  al. 2015: redescription of sixth species; Griffin and Tranter 1986: diagnosis, key to 5 species).

Tiarinia Dana, 1851 Diagnosis. Carapace with few dorsal tubercles. Rostrum comprising 2 long spines, parallel, fused or contiguous, with or without lateral spines. Supraorbital eave and postorbital spine separated by suture, antorbital angle square; orbit projecting, tubular; preorbital spine triangular, or spine acute. Antennal basal article expanded, with anterolateral spine not visible in dorsal view, separated from postorbital spine by slit. Maximum cl. 40 mm (Pl. 51g). Intertidal–shelf (0–22 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate SW Australia. 14 species (Griffin and Tranter 1986: key to 11 species; Sakai 1976: key to 4 Japanese species).

Trachymaia A. Milne-Edwards, 1880 Diagnosis. Carapace broadly pyriform, longer than wide, spinose. Rostrum comprising 2 short spines fused basally. Supraorbital eave and postorbital spine continuous, antorbital angle obsolete; preorbital spine acute; postorbital spine with tooth on anteriorlower margin. Hepatic margin spinose; branchial margin spinose. Eyestalk with large flat cornea. Pereopods 2–5 slender, up to 4 times carapace length. Maximum cl. 16 mm. Shelf, slope (150–615 m). Tropical W Atlantic. 1 species (Rathbun 1925: rediagnosis).

Tunepugettia Ng, Komai & Sato, 2017 Diagnosis. Carapace regions well defined, dorsum and legs covered in thick layer of setae, hepatic and branchial regions with conical

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lobes projected laterally. Rostrum comprising 2 short, straight, diverging spines, one-fifth carapace length. Supraorbital eave and postorbital spine separated by U-shaped notch, antorbital angle obsolete; postorbital spine slender, blunt. Cheliped articles carinate. Gonopod 1 with oblique apex. Maximum cl. 37 mm (Pl. 51h). Slope (200–1085 m). Central Indo-Pacific. 2 species (Lee et al. 2019: rediagnosis, figures; Ng et al. 2017: diagnosis).

Tyche Bell, 1836 Diagnosis. Carapace oblong-ovate, with prominent transverse ridges anterior to branchial regions meeting at median tubercle. Rostrum comprising 2 spines or lobes. Supraorbital eave and postorbital spine indistinguishable; preorbital spine acute. Branchial margin lobed, or convex; carapace posterior margin with 2 broad lobes. Maxilliped 3 exopod with basal curved process lying in groove on ischium. Maximum cl. 28 mm. Intertidal, shelf (0–60 m). Tropical W Atlantic, Tropical Eastern Pacific. 6 species (Garth 1958: rediagnosis, key to 3 E Pacific species; Williams et  al. 1977: redescriptions, key to 6 species of Tyche and Stilbomastax).

Tylocarcinus Miers, 1879 Diagnosis. Carapace ovoid, with many low tubercles. Rostrum comprising 2 long spines, contiguous or slightly diverging, or fused over basal half, 2 parallel or diverging spines distally (half or more carapace length). Supraorbital eave and postorbital spine separated by intercalated spine; preorbital spine acute. Antennal basal article narrow, lateral margin with broad lobe, with strong anterolateral spine. Maximum cl. 23 mm (Pl. 51i). Intertidal–shelf (0–278 m). Western and Central Indo-Pacific. 5 species (Dai et  al. 1994: descriptions of 2 species; Griffin and Tranter 1986: key to 3 species).

Xenocarcinus White, 1847 Diagnosis. Carapace elongate, teardrop-shaped, dorsally convex. Rostrum stout, cylindrical, with simple or bifid tip (short, with obtuse tip in X. truncatifrons). Supraorbital eave and postorbital spine indistinguishable, or separated by V-shaped notch, antorbital angle obsolete; preorbital spine absent, or spine obsolete. Hepatic margin obtusely rounded; branchial margin convex. Maximum cl. 40 mm (Pl. 51j, k). Subtidal, shelf (1–180 m; commensal with antipatharian corals). Indo-West Pacific. 7 species (Griffin and Tranter 1986: key to 5 species). References Adams A, White A (1849) Crustacea. In The Zoology of the voyage of H.M.S. Samarang; under the command of Captain Sir Edward Belcher, C.B., F.R.A.S., F.G.S., during the years 1843–1846. (Ed. Adams A) pp.  i–viii, 1–66, pls 1–13. Reeve, Bentham, and Reeve, London. Alcock A (1895) Materials for a carcinological fauna of India. No. 1. The Brachyura Oxyrhyncha. Journal of the Asiatic Society of Bengal 64, 157–291, pls 3–5.

Barnard KH (1950) Descriptive catalogue of South African decapod Crustacea (crabs and shrimps). Annals of the South African Museum 38, 1–837. Buitendijk AM (1950) Note on a collection of Decapoda Brachyura from the coasts of Mexico, including the description of a new genus and species. Zoologische Mededelingen 30, 269–282, pl. 10. Chen H-L, Ng PKL (2004) On two species of spider crabs of the genus Doclea (Crustacea: Decapoda: Brachyura: Majidae: Pisinae) from China, one of which is new. Raffles Bulletin of Zoology 52, 201–208. Chen W, Lo L, Tseng C (2015) Redescription of one epialtid crab of the genus Thusaenys from the Siaoliouciou Island, Taiwan (Crustacea: Decapoda: Brachyura: Majoidea). International Journal of Ecology 4, 1–5. doi:10.12677/IJE.2015.41001 Colavite J, Santana W, Tavares M (2016) Morphological differences between Stenocionops furcatus (Olivier, 1791) and S. coelatus (A. MilneEdwards, 1878) (Crustacea, Decapoda, Brachyura, Majoidea). Zootaxa 4184, 517–528. doi:10.11646/zootaxa.4184.3.6 Colavite J, Windsor A, Santana W (2019) Three new species and a new genus of majoid crabs from the eastern Pacific (Decapoda, Brachyura). ZooKeys 825, 1–24. doi:10.3897/zookeys.825.32271 Colavite J, Windsor AM, Santana W (2020) A new genus for Pericera septemspinosa Stimpson, 1871 and Pericera heptacantha Bell, 1836 (Crustacea, Brachyura, Majoidea), based on morphology and molecular data. Zoosystematics and Evolution 96, 205–216. doi:10.3897/zse.96.50360 Dai A, Cai Y, Yang S (1994) Studies on the crabs (Crustacea: Brachyura) in the northern part of Nansha Islands. Memoirs of the Beijing Natural History Museum 54, 1–18. Dana JD (1852) Crustacea. Part I. United States Exploring Expedition during the years 1838, 1839, 1840, 1841, 1842 under the command of Charles Wilkes, U.S.N. 13, 1–685, with a folio atlas of 96 plates (published 1855). Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. Emparanza EJM, Guzmán GL, Ng PKL (2007) A redescription of Acanthonyx petiverii H. Milne Edwards, 1834, and designation of a neotype for Acanthonyx simplex Dana, 1852 (Brachyura, Majidae). Crustaceana 80, 533–543. doi:10.1163/156854007780765542 Forest J, Guinot D (1966) Crustacés Décapodes: Brachyoures. Résultats Scientifiques des Campagnes de la Calypso 7, 23–124. Garth JS (1940) Some new species of brachyuran crabs from Mexico and the Central and South American mainland. Allan Hancock Pacific Expeditions 5, 53–95, pls 11–26. Garth JS (1952) A review of the crustacean genus Stilbognathus von Martens (Decapoda, Maiidae) with description of a new species from the east coast of Florida. Bulletin of Marine Science of the Gulf and Caribbean 1, 249–256. Garth JS (1958) Brachyura of the Pacific coast of America. Oxyrhyncha. Allan Hancock Pacific Expeditions 21, 1–854. Garth JS (1969) A new genus and species of oxyrhynchous crab from the west coast of South America (Crustacea, Decapoda, Brachyura). Boletín de la Sociedad de Biología de Concepción 41, 5–7. Garth JS (1973) New taxa of brachyuran crabs from deep water off western Peru and Costa Rica. Bulletin of the Southern California Academy of Sciences 72, 1–12. Garth JS, Haig J (1971) Decapod crustacea (Anomura and Brachyura) of the Peru-Chile Trench. Anton Bruun Report 6, 3–20. Garth JS, Méndez MG (1983) A new species of spider crab of the genus Libinia from Perú, and first known male of Delsolaria enriquei Garth,

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1973 (Crustacea, Brachyura, Majidae). Bulletin of the Southern California Academy of Sciences 82, 125–130. Griffin DJG (1973) A revision of the spider crabs of the genus Phalangipus (Crustacea, Brachyura, Majidae). Journal of Natural History 7, 165–207. doi:10.1080/00222937300770151 Griffin DJG, Tranter HA (1974) Spider crabs of the family Majidae (Crustacea: Decapoda: Brachyura) from the Red Sea. Israel Journal of Zoology 23, 162–198. Griffin DJG, Tranter HA (1986) The Decapoda Brachyura of the Siboga Expedition. Part VIII Majidae. Siboga-Expéditie 39 c4, 1–335. Guinot D (1979) Données nouvelles sur la morphologie, la phylogenèse et la taxonomie des Crustacés Décapodes Brachyoures. Mémoires du Muséum National d’Histoire Naturelle, Paris, ser. Zoologie 112, 1–354. Henderson JR (1893) A contribution to Indian carcinology. Transactions of the Linnean Society of London, Series 2, Zoology 5, 325–458, pls 36–40. Hendrickx ME, Cervantes JL (2003) A new species of Loxorhynchus Stimpson (Decapoda, Majoidea, Pisidae) from the Pacific Coast of Mexico. Crustaceana 76, 103–113. doi:10.1163/156854003321672863 Holthuis LB (1966) Notes on Neodoclea boneti Buitendijk, a crab from the Eastern Pacific (Crustacea: Decapoda, Brachyura). Bulletin of Marine Science 16, 376–381. Hosie AM, Hara A (2016) Description of a new species of brooding spider crab in the genus Paranaxia Rathbun, 1924 (Brachyura: Majoidea), from northern Australia and Indonesia. Zootaxa 4127, 121–134. doi:10.11646/zootaxa.4127.1.6 Kazmi QB, Tirmizi NM (1995) A new species of Cyphocarcinus and a new record of Macropodia formosa Rathbun (Crustacea, Decapoda, Majidae) from Pakistan. Pakistan Journal of Marine Sciences 4, 59–65. Lee BY, Ng PKL (2020) The identity of Hyastenus pleione (Herbst, 1803) and description of a new species from China (Decapoda, Brachyura, Majoidea, Epialtidae). Crustaceana 93, 1343–1360. doi:10.1163/15685403-bja10064 Lee BY, Richer de Forges B, Ng PKL (2017) Deep-sea spider crabs of the families Epialtidae MacLeay, 1838 and Inachidae MacLeay, 1838, from the South China Sea, with descriptions of two new species (Decapoda, Brachyura, Majoidea). European Journal of Taxonomy 358, 1–37. doi:10.5852/ejt.2017.358 Lee BY, Richer de Forges B, Ng PKL (2019) Deep-sea spider crabs of the family Epialtidae MacLeay, 1838, from Papua New Guinea, with a redefinition of Tunepugettia Ng, Komai & Sato, 2017, and descriptions of two new genera (Crustacea: Decapoda: Brachyura: Majoidea). Zootaxa 4619, 1–44. doi:10.11646/zootaxa.4619.1.1 Lee BY, Richer De Forges B, Ng PKL (2020) Revision of the deep-water spider crab genus, Scyramathia A. Milne-Edwards, 1880, with the description of a new species from the Mediterranean and notes on Rochinia A. Milne-Edwards, 1875, and Anamathia Smith, 1885 (Crustacea, Decapoda, Brachyura, Epialtidae). Zoosystematics and Evolution 96, 537–569. doi:10.3897/zse.96.48041 Lee BY, Richer de Forges B, Ng PKL (2021) The generic affinities of the Indo-West Pacific species assigned to Rochinia A. Milne-Edwards, 1875 (Crustacea: Brachyura: Majoidea: Epialtidae). Raffles Bulletin of Zoology 69, 19–44. Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Milne Edwards H (1834) Histoire naturelle des Crustacés, comprenant l’anatomie, la physiologie et la classification de ces animaux. Vol. 1. Librairie Encyclopédique de Roret, Paris. Milne-Edwards A (1879) Description de quelques Crustacés nouveaux. Bulletin de la Société Philomathique de Paris, 7e série 7, 103–110. Milne-Edwards A (1873–1881) Études sur les xiphosures et les crustacés de la région mexicaine. In Mission scientifique au Mexique et dans l’Amérique Centrale. Recherches zoologiques pour servie a l’histoire de

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la faune de l’Amerique Centrale et du Mexique. 5ème partie. pp. 1–368, pls 1–29. Imprimerie Nationale, Paris. Monod T (1956) Hippidea et Brachyura ouest-africains. Mémoires de I’Institut Français d’Afrique Noire 45, 1–674. Morgan GJ (1987) Brooding of juveniles and observations on dispersal of young in the spider crab Paranaxia serpulifera (Guérin)(Decapoda, Brachyura, Majidae) from Western Australia. Records of the Western Australian Museum 13, 337–343. Naderloo R (2015) A new species of decorator crabs, genus Menaethiops Alcock, 1895 (Crustacea: Decapoda: Brachyura: Majoidea: Epialthidae), from Abu-Musa Island, Persian Gulf, Iran. Zootaxa 3925, 431–437. doi:10.11646/zootaxa.3925.3.7 Ng PKL, Anker A (2014) Pteromaja maklayi gen. et sp. nov., a remarkable new spider crab (Crustacea: Brachyura: Epialtidae) from Papua New Guinea. Marine Biology Research 10, 816–823. doi:10.1080/17451000.2 013.853127 Ng PKL, Richer de Forges B (2013) Samadinia longispina, a new genus and species of deep-sea spider crab from the western Pacific, and a new species of Rochinia A. Milne-Edwards, 1875, from Papua New Guinea (Crustacea: Brachyura: Majoidea: Epialtidae). Zootaxa 3718, 357–366. doi:10.11646/zootaxa.3718.4.5 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Ng PKL, Komai T, Sato T (2017) On the trail of a Japanese “ghost species”—the identity of Goniopugettia tanakae Sakai, 1986, and the establishment of a new genus for Pugettia sagamiensis Gordon, 1930 (Decapoda, Brachyura, Epialtidae). Crustacean Research 46, 133–152. doi:10.18353/crustacea.46.0_133 Ohtsuchi N, Komatsu H, Li X (2020) A new kelp crab species of the genus Pugettia (Crustacea: Decapoda: Brachyura: Epialtidae) from Shandong Peninsula, northeast China. Species Diversity 25, 237–250. doi:10.12782/specdiv.25.237 Poupin J (1995) Etude des Naxioides du groupe robillardi (Miers, 1882) (Brachyura: Majidae; Pisinae). Journal of Natural History 29, 85–109. doi:10.1080/00222939500770051 Pretzmann G (1961) Eine neue amerikanische Krabbe der Gattung Lissa. Annalen des Naturhistorischen Museums in Wien 64, 173–177, pl. 4. Rathbun MJ (1911) Marine Brachyura. In: The Percy Sladen Trust expedition to the Indian Ocean in 1905, under the leadership of Mr. J. Stanley Gardiner. Volume III. No. XI. Transactions of the Linnean Society of London, Series 2, Zoology 14, 191–261, pls 15–20. Rathbun MJ (1923) The brachyuran crabs collected by the U.S. Fisheries steamer ‘Albatross’ in 1911, chiefly on the west coast of Mexico. Bulletin of the American Museum of Natural History 48, 619–637. Rathbun MJ (1925) The spider crabs of America. Bulletin of the United States National Museum 129, 1–613. Richer de Forges B (1994) A new genus of deep-sea majid crab: Griffinia gen. nov. (Crustacea, Decapoda, Brachyura). The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 11, 65–72. Richer de Forges B, Ng PKL (2009a) New genera, new species and new records of Indo-West Pacific spider crabs (Crustacea: Brachyura: Epialtidae: Majoidea). Zootaxa 2025, 1–20. doi:10.11646/zootaxa.2025.1.1 Richer de Forges B, Ng PKL (2009b) On the majoid genera Oxypleurodon Miers, 1886, and Sphenocarcinus A. Milne-Edwards, 1875 (Crustacea: Brachyura: Epialtidae), with descriptions of two new genera and five new species. Raffles Bulletin of Zoology Supplement 20, 247–266. Richer de Forges B, Ng PKL (2012) Griffinia takedai, a new species of deep sea majoid crab (Decapoda, Brachyura, Epialtidae) from the Philippines.

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In: Komatsu H, Okuno J, Fukuoka K (Eds) Studies on Eumalacostraca: a homage to Masatsune Takeda. Crustaceana Monographs 17, 275–284. doi:10.1163/9789004202894_023 Sakai T (1938) Studies on the crabs of Japan. III. Brachygnatha, Oxyrhyncha. Yokendo C., Tokyo. Sakai T (1965) The crabs of Sagami Bay collected by His Majesty the Emperor of Japan. Maruzen Co. Ltd, Tokyo. Sakai T (1967) Notes from the carcinological fauna of Japan (III). Crustacean Research 3, 68–83[frontispiece.]. doi:10.18353/rcustacea.3.0_68 Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo. Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Takeda M, Marumura M (2010) Spider crabs of the genus Huenia De Haan, 1837 (Crustacea, Decapoda, Brachyura) from Japan, with descriptions of two new species. Bulletin of the National Museum of Nature and Science. Series A, Zoology 36, 39–48. Takeda M, Ahyong ST, Ohtsuchi N, Komatsu H (2022) Crabs (Crustacea, Decapoda) from the sea off East and Southeast Asia collected by the RV Hakuhō Maru (KH-72-1 Cruise) 2. Timor Sea. Bulletin of the National Museum of Nature and Science. Series A, Zoology 48, 5–24. doi:10.50826/bnmnszool.48.1_5 Tamburus AF, Mantelatto FL (2021) Revision of the spider crab genus Libinia Leach, 1825 (Decapoda: Brachyura: Epialtidae: Pisinae). Journal of Crustacean Biology 41, ruab035. doi:10.1093/jcbiol/ruab035 Targioni Tozzetti A (1877) Zoologia del viaggio intorno al Globo della R. Pirocorvetta Magenta durante gli anni 1865–1868. Crostacei Brachyuri e Anomuri. Pubblicazioni del R. Istituto di Studi Superiori Pratici e di Perfezionamento in Firenze, Sezione di Scienze Fisiche e Naturali 1, i–xxix, 1–257, pls I–XII. Tavares MS (1991) Redéfinition des genres Rochinia A. Milne Edwards, Sphenocarcinus A. Milne Edwards et Oxypleurodon Miers, et établissement du genre Nasutocarcinus gen. nov. (Crustacea, Brachyura, Majidae). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 13, 159–179. Tavares M, Santana W (2011) A new genus for Libinia rostrata Bell, 1835, with comments on the validity of Libinia bellicosa Oliveira, 1944 (Crustacea, Brachyura, Majoidea, Epialtidae). Zootaxa 3057, 61–68. doi:10.11646/zootaxa.3057.1.3 Tavares M, Santana W (2018) Refining the genus Rochinia A. MilneEdwards, 1875: reinstatement of Scyramathia A. Milne-Edwards, 1880 and Anamathia Smith, 1885, and a new genus for Amathia crassa A. Milne-Edwards, 1879, with notes on its ontogeny (Crustacea: Brachyura: Epialtidae). Zootaxa 4418, 201–227. doi:10.11646/zootaxa.4418.3.1 Tirmizi NM (1978) Notes on Simocarcinus simplex (Dana) from the Arabian Sea (Decapoda, Brachyura). Crustaceana 35, 311–312. doi:10.1163/156854078X00475 Wagner HP (1986) A revision of the genus Doclea Leach, 1815 (Crustacea, Brachyura, Majidae). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série 8, 893–953. Wagner HP (1992) A new species of Austrolibinia from the South China Sea and Indonesia (Crustacea: Brachyura: Majidae). Raffles Bulletin of Zoology 40, 69–73. Wicksten MK (1987) Nicoya tuberculata, a new genus and species of spider crab from Pacific Costa Rica (Majidae: Pisinae). Proceedings of the Biological Society of Washington 100, 691–693. Williams AB, Shaw JK, Hopkins TS (1977) Stilbomastax, a new genus of spider crab (Majidae: Tychinae) from the West Indies region, with notes on American relatives. Proceedings of the Biological Society of Washington 90, 884–893. Windsor AM, Ahyong ST (2013) Hyastenus baru, a new species of spider crab from Indonesia (Brachyura, Majoidea, Epialtidae) with a key to

the species of Hyastenus. Crustaceana 86, 718–727. doi:10.1163/​ 15685403-00003202 Windsor AM, Felder DL (2014) Molecular phylogenetics and taxonomic reanalysis of the family Mithracidae MacLeay (Decapoda : Brachyura : Majoidea). Invertebrate Systematics 28, 145–173. doi:10.1071/ IS13011 Wu S-H, Yu H-P, Ng PKL (1999) Acanthonyx formosa, a new species of spider crab (Decapoda, Brachyura, Majidae) from seaweed beds in Taiwan. Crustaceana 72, 193–202. doi:10.1163/156854099503285 Zariquiey Alvarez R (1968) Crustáceos decápodos ibéricos. Investigaciones Pesqueras 32, 1–510.

Inachidae Macleay, 1838 spider crabs Figure 14.80–14.82, Plate 52a–e The absence of an orbit and the exposed eyestalks are the defining features of members of this family. They are distinguished from similar inachoidids by not having the thoracic pleura visible, and from oregoniids in usually not having the male telson embedded in pleonite 6 and lacking a longitudinal groove on gonopod 1. The differences from Oregoniidae are subtle. Oregoniidae were recently expanded to include several genera previously included in Inachidae on evidence from larval studies (Clark and Webber 1991) and molecular support (Marco-Herrero et  al. 2013). Inachids now include mostly species of usually 30 mm carapace length or less, whereas oregoniids usually exceed 40 mm carapace length. Earlier divisions of Inachidae into subfamilies (Guinot 2012) or into 17 tribes (Števčić 2005) are largely now irrelevant. Diagnosis. Orbit absent, eyes unprotected though orbital margin usually with several small spines and postorbital spine; basal antennal article usually extremely slender, more than twice as long as wide, and sometimes free distally. Thoracic pleurites 5–8 not visible dorsally, covered by posterolateral margin of carapace. Eyestalk long, non-retractile or retractile against acute postorbital spine affording no concealment. Male pleonite 6 and telson usually articulating, if telson recessed, pleonites 3–5 fused (Paratymolus, Dumea). Pleonite 1 free from carapace. Maxilliped 3 merus often not as wide as ischium. Gonopod 1 usually slender, straight or weakly curved, rarely twisted, often protected by flap, apex simple or with lobe. Implicit generic attributes. Carapace pyriform. Interantennular spine absent. Supraorbital eave smooth, without preorbital spine, without antorbital lobe, without postorbital spine, without intercalated spine. Basal antennal article distal margin at least level with, usually extending forward beyond, anterior margin of eyestalk, free or fused to front. Pereopod 2 little longer than other pereopods; 5 dactylus not reflexed.

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609

Key to genera of Inachidae 1.

Rostrum of 2 broad, blunt or acute lobes. Carapace without dorsal spines (Fig. 14.81p). Interantennular septum weakly developed (Fig. 14.82e). Pereopods 2, 3 propodi flexor margins expanded, dactyli simple; pereopods 4, 5 prehensile (Fig. 14.82o)�������������������������������������������������������������������������������������������������������������������Oncinopus – Rostrum of 2 teeth , lobes or spines, or single. Carapace often with dorsal spines. Interantennular septum usually present. Pereopods 2–5 without expanded propodi, sometimes becoming more prehensile posteriorly������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 2. Rostrum comprising pair or teeth or spines, sometimes contiguous or fused basally; with or without interantennular spine�����������������������������������������������������������������������������������������������������������������������������������������������������������������3 – Rostrum comprising single lobe, tooth or spine, or absent; without or with interantennular tooth not visible in dorsal view�����������������������������������������������������������������������������������������������������������������������������������������������������������������19 3. Postorbital spine or lobe present (Figs 14.80i, 14.81a, h, j–l, 14.82h)����������������������������������������������������������������������������������������4 – Postorbital spine or lobe absent (e.g. Fig. 14.81f, n, s)���������������������������������������������������������������������������������������������������������������10 4. Supraorbital eave with preorbital tooth (Fig. 14.81a, h)�������������������������������������������������������������������������������������������������������������5 – Supraorbital eave without preorbital tooth (e.g. Fig. 14.81j–l)���������������������������������������������������������������������������������������������������8 5. Rostrum comprising pair of long, tapering, largely contiguous spines, curving downwards, with lateral spiniform setae. Carapace dorsum smooth (Fig. 14.80d)�������������������������������������������������������������������������������� Anomalothir – Rostral spines separate, not contiguous. Carapace with few middorsal tubercles or spines (Fig. 14.81n, o)���������������������6 6. Rostrum comprising 2 blunt lobes, interantennular spine with lateral spinules, visible in dorsal view. Postorbital spine with accessory spine (Fig. 14.82h)���������������������������������������������������������������������������������������� Rhinospinosa – Rostrum comprising 2 sharp spines, interantennular spine if present, without lateral spinules. Postorbital spine without accessory spine���������������������������������������������������������������������������������������������������������������������������������������������������7 7. Rostrum of pair of diverging slender acute spines, smooth laterally (Fig. 14.81a). Pereopods 4, 5 dactyli curved, bifid, flexor margins dentate (Fig. 14.82q)���������������������������������������������������������������������������������������������� Achaeopsis – Rostrum of 2 diverging or converging spines, with lateral spinules (Fig. 14.81h). Pereopods 2–5 dactyli gently curved, styliform����������������������������������������������������������������������������������������������������������������������������������������� Dorhynchus 8. Carapace with 1 mesogastric and 1 cardiac spine, plus lateral pairs of spines (Fig. 14.81l). Postorbital spine broad in lateral view, inner surface cupped����������������������������������������������������������������������������������������������������������������Inachus – Carapace smooth or with tubercles or spines not so arranged. Postorbital spine sharp������������������������������������������������������9 9. Carapace intestinal region with large blunt medial lobe on posterior margin. Pereopod 2 1.5 times length and more robust than other pereopods (Fig. 14.80i). Australian endemic�������������������������������������������������������� Ephippias – Carapace intestinal region without lobe on posterior margin. Pereopod 2 sometimes longer but not more robust than other pereopods. Americas��������������������������������������������������������������������������������������������������������������������������������10 10. Rostrum of 2 spines, contiguous over at least half length (Fig. 14.81k). Pereopods 2–5 twice carapace length, propodi flexor margins expanded (Fig. 14.82n)�������������������������������������������������������������������������������������Eurypodius – Rostrum of 2 rounded-triangular teeth, together shorter than basal width (Fig. 14.81j). Pereopods 2–5 about carapace length, propodi cylindrical (Fig. 14.82m)�����������������������������������������������������������������������������������Eucinetops 11. Rostrum comprising pair of spines basally fused or contiguous������������������������������������������������������������������������������������������� 12 – Rostrum comprising pair of short spines or teeth free from base������������������������������������������������������������������������������������������13 12. Rostral spines fused for about basal half. Carapace intestinal region with slender medial spine or low tubercle on posterior margin (Fig. 14.81u). Pereopods 2–5 about as long as carapace�����������������������������������������Sunipea – Rostral spines contiguous or fused over whole length. Carapace intestinal region without medial spine on posterior margin. Pereopods 2–5 longer than carapace����������������������������������������������������������������������������������� Macropodia 13. Rostrum comprising pair of sharp spines with spinulose margins; interantennular process cupped, with distal spine, visible in dorsal view. Supraorbital margin spinose (Fig. 14.81b). E Atlantic only�������������������������������������������������������������������������������������������������������������������������������������� Calypsachaeus – Rostrum comprising pair of blunt lobes or spines with smooth margins; interantennular process absent. Supraorbital margin smooth or spinulose�����������������������������������������������������������������������������������������������������������������������������14

610

Marine Decapod Crustacea

Fig. 14.80.  Inachidae. a, Achaeus curvirostris (A. Milne-Edwards, 1873); b, c, Anisonotus curvirostris A. Milne-Edwards, 1879 (with lateral orbit, rostrum); d, Anomalothir hoodensis Garth, 1939; e, f, Dumea latipes (Haswell, 1880) (with lateral anterior carapace); g, Capartiella longipes (Capart, 1951); h, Encephaloides armstrongi Wood-Mason & Alcock, 1891; i, Ephippias endeavouri Rathbun, 1918; j, Metoporhaphis calcarata (Say, 1818).

14. – 15. – 16.

Eyestalk small, almost immovable, almost unpigmented (Fig. 14.81r)��������������������������������������������������������������� Physachaeus Eyestalk movable, cornea pigmented������������������������������������������������������������������������������������������������������������������������������������������15 Cheliped carpus with tooth on upper margin (Fig. 14.82j–l)��������������������������������������������������������������������������������������������������16 Cheliped carpus without tooth on upper margin���������������������������������������������������������������������������������������������������������������������18 Carapace and posterolateral margin tuberculate. Eyestalk as long as half width between orbits (Fig. 14.81q). Cheliped carpal spine long, slender (Fig. 14.82l). Male pleonites 3–5 fused, lateral margins straight (Fig. 14.82v)������������������������������������������������������������������������������������������������������������������������������������������������������������� Paratymolus

14 – Brachyura – crabs

611

Fig. 14.81.  Inachidae. Carapace, representative or only species: a, Achaeopsis; b, Calypsachaeus; c, Camposcia; d, Chalaroachaeus; e, Chorinachus; f, g, Coryrhynchus (with lateral view of rostrum); h, Dorhynchus (anterior only); i, Ericerodes; j, Eucinetops; k, Eurypodius; l, Inachus; m, Litosus; n, o, Macropodia (with lateral view of rostrum); p, Oncinopus (anterior only); q, Paratymolus; r, Physachaeus; s, Podochela; t, Prosphorachaeus; u, Sunipea.

– 17. – 18. – 19.

Carapace and posterolateral margin smooth. Eyestalk short (Figs 14.80e, f, 14.81m). Cheliped carpal spine short, stout (Fig. 14.82j, k). Male pleonites 3–5 fused, lateral margins concave (Fig. 14.82u)���������������������������������������17 Carapace anterolateral margin with 3 large spines and 2 intermediate spines. Subhepatic region spinose (Fig. 14.80e, f). Cheliped outer surface with acute granules (Fig. 14.82j)����������������������������������������������������������������Dumea Carapace anterolateral margin with 1 broad lobe and 1 anteriorly directed spine (Fig. 14.81m). Subhepatic region smooth. Cheliped outer surface smooth (Fig. 14.82k)����������������������������������������������������������������������������������� Litosus Pereopods 2–5 slender, 3–5 times carapace length (Fig. 14.80a). Gonopod 1 simple, with subterminal aperture, without lobes (Fig. 14.82y). Male pleonite 3 not longer than pleonites 4 and 5, smooth or sometimes with spinules on inflated lateral surfaces, without transverse ridge (Fig. 14.82t)���������������������������Achaeus Pereopods 2–5 less than twice carapace length. Gonopod 1 complex, aperture surrounded by lobes (Fig. 14.82z). Male pleonite 3 longer than pleonites 4 and 5, arched, with transverse ridge (Fig. 14.82w, x)�������������������������������������������������������������������������������������������������������������������������������������������������Prosphorachaeus Rostrum absent. Supraorbital eave rudimentary (Fig. 14.81d). Interantennular septum absent (Fig. 14.82b)������������������������������������������������������������������������������������������������������������������������������������������������������� Chalaroachaeus

612

Marine Decapod Crustacea

– 20. – 21.

Rostrum present. Supraorbital eave narrow. Interantennular septum present������������������������������������������������������������������� 20 Rostrum almost as long as or longer than postocular length��������������������������������������������������������������������������������������������������21 Rostrum much shorter than postocular length������������������������������������������������������������������������������������������������������������������������ 22 Carapace with few gastric and cardiac tubercles. Rostrum laterally smooth. Pereopods 2–5 meri with strong distal spine (Fig. 14.80j). W Atlantic��������������������������������������������������������������������������������������������������� Metoporhaphis – Carapace with few branchial tubercles, with intestinal tooth. Rostrum with lateral spinules (Fig. 14.81e). Pereopods 2–5 meri without strong distal spine. Indo-West Pacific������������������������������������������������������������� Chorinachus 22. Rostrum narrow curved downwards, with dorsal tubercles. Postocular process lobe-like. Supraorbital eave with 2 long spines (Fig. 14.80b, c)���������������������������������������������������������������������������������������������������������������������������Anisonotus – Rostrum not curved downwards, without dorsal tubercles. Postocular process absent or a simple granule. Supraorbital eave smooth�������������������������������������������������������������������������������������������������������������������������������������������������������� 23 23. Rostrum semicircular or almost circular, excavate below, with dorsal ridge surrounded by hooked setae (Fig. 14.81f, g)��������������������������������������������������������������������������������������������������������������������������������������������������������Coryrhynchus

Fig. 14.82.  Inachidae. Antennule, antennal basal article, orbit: a, Camposcia; b, Chalaroachaeus; c, Eucinetops; d, Macropodia; e, Oncinopus; f, Podochela; g, Prosphorachaeus; h, Rhinospinosa. Rostrum, orbit, oblique-anterior view: i, Capartiella. Cheliped: j, Dumea; k, Litosus; l, Paratymolus. Pereopods 2, 4: m, Eucinetops; n, Eurypodius; o, Oncinopus; p, Podochela. Pereopod 5 dactylus: q, Achaeopsis; r, s, Achaeus. Male pleon: t, Achaeus; u, Dumea; v, Paratymolus; w, x, Prosphorachaeus (dorsal and lateral views). Gonopod 1: y, Achaeus; z, Prosphorachaeus. Features labelled on g: aba, antennal basal article; n, nephropore (antennal gland); r, rostral spine

14 – Brachyura – crabs

613

– Rostrum blunt or triangular, with or without terminal spine����������������������������������������������������������������������������������������������� 24 24. Rostrum blunt, with apical notch. Carapace densely setose. Eyestalk long, curved (Fig. 14.81c). Interantennular septum absent (Fig. 14.82a)�������������������������������������������������������������������������������������������������������� Camposcia – Rostrum triangular, with or without terminal spine. Carapace not setose. Eyestalk straight. Interantennular septum present (Fig. 14.82f)���������������������������������������������������������������������������������������������������������������������� 25 25. Rostrum triangular, with terminal spine (Fig. 14.81i)����������������������������������������������������������������������������������������������� Ericerodes – Rostrum triangular, without terminal spine���������������������������������������������������������������������������������������������������������������������������� 26 26. Carapace branchial regions extremely elevated, approximated in midline. Rostrum broad, beak-like, longer than wide at base (Fig. 14.80h). India���������������������������������������������������������������������������������Encephaloides – Carapace branchial regions not greatly elevated or approximated in midline. Rostrum triangular, wider than or as wide as long. Americas, Atlantic�������������������������������������������������������������������������������������������������������������� 27 27. Pereopods 2–5 setose, 3 times carapace length. Rostrum broadly triangular; interantennular spine visible in antero-oblique view (Figs 14.80g, 14.82i)����������������������������������������������������������������������������������������������������������Capartiella – Pereopods 2–5 not setose, about twice times carapace length. Rostrum triangular, with rounded tip; interantennular spine absent (Figs 14.81s, 14.82f)�������������������������������������������������������������������������������������������������Podochela Achaeopsis Stimpson, 1857

Anomalothir Miers, 1879

Diagnosis. Carapace with dorsal spines. Rostrum comprising 2 short diverging spines. Interantennular spine present. Supraorbital eave with preorbital spine, with postorbital spine. Eyestalk short, straight. Pereopods 2–5 slender, 4, 5 dactyli curved, bifid, flexor margins dentate. Maximum cl. 12 mm. Shelf (40–200 m). Temperate Southern Africa. 1 species (Barnard 1950).

Diagnosis. Carapace without dorsal spines. Rostrum comprising 2 tapering, largely contiguous spines, longer than to about half postorbital length, curving downwards, with lateral spiniform setae. Supraorbital eave with small preorbital spine, with triangular postorbital tooth. Eyestalk short. Pereopods 2–5 longer than carapace length, 2 and 3 much longer than 3 and 4. Maximum cl. 20 mm. Shelf, slope (50–293 m). Temperate W Northern and Tropical W Atlantic, Tropical Eastern Pacific. 3 species (Garth 1939: figures; 1958; Rathbun 1925: rediagnosis, key to 2 species).

Achaeus Leach, 1817 Diagnosis. Carapace with or without dorsal spines. Rostrum comprising 2 blunt spines or lobes. Supraorbital eave spinulose or smooth. Eyestalk elongate. Pereopods 2–5 slender, 3–5 times carapace length; 3–5, 4 and 5, or 5 only dactyli arcuate, flexor margins dentate. Male pleonites 3 not longer than pleonites 4 and 5, smooth or sometimes with spinules on inflated lateral surfaces, without transverse ridge. Gonopod 1 simple, with subterminal aperture. Maximum cl. 12 mm. Intertidal–slope (0–540 m). Temperate Northern and Tropical E Atlantic, Temperate Northern W Pacific, Indo-West Pacific, Temperate Southern Africa, Temperate Australasia. 38 species (Griffin 1970: redagnosis, key to 8 Australian species; Griffin and Tranter 1986: key to 27 species; Manning and Holthuis 1981: key to 7 E Atlantic species).

Anisonotus A. Milne-Edwards, 1879 Diagnosis. Carapace without dorsal spines. Rostrum single, arched, with dorsal tubercles, shorter than postorbital carapace length. Interantennular septum produced, with small tooth, visible in lateral view. Supraorbital eave with 2 long spines, with lobate postorbital process. Eyestalk elongate. Pereopods 2–5 about twice carapace length, first longest; dactyli more curved posteriorly. Maximum cl. 11 mm. Shelf, slope (134–448 m). Tropical W Atlantic. 2 species (Coelho 2006: diagnosis; Rathbun 1925: species descriptions as Podochela).

Calypsachaeus Manning & Holthuis, 1981 Diagnosis. Carapace without dorsal spines. Rostrum comprising 2 short spines with spinulose margins. Interantennular process cupped, with distal spine. Supraorbital eave spinulose. Eyestalk elongate. Pereopods 2–5 slender, setose, 3 times carapace length; 4, 5 dactyli strongly arcuate, flexor margins dentate. Maximum cl. 9.3 mm. Shelf (30–70 m). Tropical E Atlantic. 1 species (Forest and Guinot 1966).

Camposcia Latreille, 1829 Diagnosis. Carapace setose. Rostrum broad, short, with apical notch. Interantennular septum absent. Supraorbital eave with small postorbital spine. Eyestalk long, curved. Maximum cl. 28 mm (Pl. 52a). Intertidal–slope (0–220 m; always densely covered in sponge). Temperate Northern W Pacific, Western and Central IndoPacific, Temperate Australasia. 1 species (Barnard 1950: figure).

Capartiella Manning & Holthuis, 1981 Diagnosis. Carapace without dorsal spines. Rostrum broadly triangular, depressed. Interantennular spine broad, not visible in dorsal view. Eyestalk elongate. Pereopods 2–5 slender, setose, 3

614

Marine Decapod Crustacea

times carapace length; all dactyli slender, gently curved, flexor margin dentate. Maximum cl. 14 mm. Shelf (33–82 m). Tropical E Atlantic. 1 species (Manning and Holthuis 1981).

margin with few teeth. Male pleonites 3–5 fused, lateral margins concave. Maximum cl. 10 mm (Pl. 52c). Intertidal, subtidal (0–102 m). Central Indo-Pacific, Temperate Australasia. 2 species (Loh and Ng 1999: key to species).

Chalaroachaeus De Man, 1902

Encephaloides Wood-Mason & Alcock, 1891

Diagnosis. Carapace with dorsal spines. Rostrum absent. Interantennular septum weakly developed. Eyestalk elongate. Pereopods 2–5 slender, 3–5 times carapace length; 3–5 dactyli with bifid apex. Maximum cl. 4 mm. Shelf (to 36 m). Temperate Northern W Pacific, Central IndoPacific. 1 species (Griffin and Tranter 1986; De Man 1902: diagnosis, figures).

Diagnosis. Carapace cordiform; branchial regions extremely elevated, approximated in midline. Rostrum broad, beak-like, longer than wide at base. Eyestalk short. Pereopods 2–5 slender, 4 times carapace length; all dactyli slender, gently curved, styliform. Maximum cl. 42 mm. Shelf, slope (110–1230 m). Western Indo-Pacific (India). 2 species (Alcock and Anderson 1896: figure).

Chorinachus Griffin & Tranter, 1986 Diagnosis. Carapace with dorsal spines. Rostrum long, spine-like with lateral spinules, almost as long as postorbital carapace length. Supraorbital eave with small preorbital spine, with postorbital spine. Eyestalk short. Maximum cl. 17 mm. Shelf, slope (60–300 m). Western Indo-Pacific. 1 species (Alcock 1895: figure; Griffin and Tranter 1986: diagnosis).

Coryrhynchus Kingsley, 1880 Diagnosis. Carapace with cardiac tubercle. Rostrum semicircular or almost circular, excavate below, with dorsal ridge surrounded by hooked setae. Supraorbital eave with postorbital granule. Eyestalk short. Pereopods 2–5 about twice carapace length, first longest; dactyli more curved posteriorly. Maximum cl. 21 mm. Subtidal, shelf (1–229 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 7 species (Coelho 2006: diagnosis; Garth 1958: as Podochela; Rathbun 1925: species descriptions as Podochela).

Dorhynchus Thomson, 1873 Diagnosis. Carapace with dorsal spines. Rostrum comprising 2 spines diverging or converging anteriorly, with lateral spinules. Interantennular spine present. Supraorbital eave with preorbital spine, with postorbital spine. Eyestalk short, straight. Pereopods 2–5 dactyli slender, gently curved, styliform. Maximum cl. 25 mm. Shelf–bathyal (106–2080 m). Temperate Northern and Tropical Atlantic, Temperate Northern W Pacific, Temperate Southern Africa, Temperate Australasia. 5 species (Griffin and Tranter 1986: description as Achaeus; Macpherson 1983; Manning and Holthuis 1981: diagnosis; Sakai 1932: figure).

Dumea Loh & Ng, 1999 Diagnosis. Carapace pentagonal; anterolateral margin with 3 large spines and 2 intermediate spines, subhepatic region spinose. Rostrum of 2 blunt lobes. Eyestalk short, straight. Cheliped carpus with short stout tooth on upper margin. Pereopods 2–5 little longer than carapace length; dactyli almost straight, flexor

Ephippias Rathbun, 1918 Diagnosis. Carapace intestinal region with large blunt medial lobe on posterior margin. Rostrum comprising 2 slender spines, separate from base. Supraorbital eave with postorbital spine (remote). Eyestalk elongate. Pereopods 2–5 about twice carapace length; pereopod 2 1.5 times length of other pereopods, more robust. Maximum cl. 120 mm. Subtidal, shelf (18–196 m). Central Indo-Pacific, Temperate Australia. 1 species (Poore 2004).

Ericerodes Rathbun, 1897 Diagnosis. Carapace with cardiac tubercle. Rostrum with triangular base and prolonged spine. Supraorbital eave with postorbital granule. Eyestalk short. Pereopods 2–5 about twice carapace length, first longest; dactyli more curved posteriorly. Maximum cl. 26 mm. Subtidal, shelf (1–120 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 9 species (Coelho 2006: diagnosis; Garth 1958: species descriptions as Podochela; Rathbun 1925: species descriptions as Podochela).

Eucinetops Stimpson, 1860 Diagnosis. Carapace without dorsal spines. Rostrum of 2 roundedtriangular teeth, together shorter than basal width. Supraorbital eave with postorbital spine. Pereopods 2–5 about carapace length; propodi cylindrical. Maximum cl. 12.5 mm. Intertidal, subtidal (0–6 m). Temperate E Northern Pacific, Tropical Atlantic, Tropical Eastern Pacific. 4 species (Garth 1958: rediagnosis, key to 3 Pacifc species).

Eurypodius Guérin, 1828 Diagnosis. Carapace with dorsal tubercles. Rostrum comprising 2 spines, contiguous over at least half length. Interantennular spine absent, or spine present. Supraorbital eave with postorbital spine. Pereopods 2–5 twice carapace length; propodi flexor margins expanded. Maximum cl. 69 mm. Subtidal–slope (7–320 m). Temperate South America. 2 species (Rathbun 1925: rediagnosis, key to species).

14 – Brachyura – crabs

Inachus Weber, 1795 Diagnosis. Carapace with 1 mesogastric and 1 cardiac spine, plus lateral pairs of spines. Rostrum comprising 2 blunt spines, or of 2 blunt lobes. Interantennular spine present. Supraorbital eave with postobital broad, cupped process. Eyestalk short, straight. Pereopods 2–5 about twice carapace length; pereopod 2 1.5 times length of other pereopods, more robust. Maximum cl. 30 mm. Subtidal–slope (10–548 m). Temperate Northern and Tropical Atlantic, Temperate Southern Africa. 13 species (Manning and Holthuis 1981: rediagnosis, key to 8 W Atlantic species).

Litosus Loh & Ng, 1999 Diagnosis. Carapace pentagonal; anterolateral margin with 1 broad lobe and 1 anteriorly directed spine, subhepatic region smooth. Rostrum of 2 blunt lobes. Eyestalk short, straight. Cheliped carpus with short stout tooth on upper margin. Pereopods 2–5 little longer than carapace length; dactyli almost straight, flexor margin with few teeth. Male pleonites 3–5 fused, lateral margins concave. Maximum cl. 8 mm. Intertidal, subtidal (0–18 m). Western and Central Indo-Pacific, Temperate Australia. 2 species (Loh and Ng 1999: key to species).

Macropodia Leach, 1814 Diagnosis. Carapace with dorsal spines. Rostrum comprising 2 tapering fused spines. Interantennular septum produced, with small tooth, visible in lateral view. Supraorbital eave with antorbital lobe. Eyestalk elongate. Pereopods 2–5 slender, 3–5 times carapace length; 4, 5 dactyli strongly arcuate, flexor margins dentate. Maximum cl. 19 mm. Intertidal–slope (0–748 m). Temperate Northern and Tropical E Atlantic, Western Indo-Pacific, Temperate Australasia. 17 species (Forest 1978: rediagnosis, key to 5 species from E Atlantic; Griffin 1974: figures; Manning and Holthuis 1981: key to 8  species from E Atlantic; Spiridonov et  al. 2020: species of Black Sea).

615

propodi flexor margins expanded, pereopods 4, 5 subchelate. Maximum cl. 13 mm (Pl. 52b). Subtidal–slope (1–460 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia. (­Australia). 5 species (Davie 2011: rediagnosis, key to 5 species).

Paratymolus Miers, 1879 Diagnosis. Carapace and posterolateral margin tuberculate. Rostrum of 2 blunt lobes. Eyestalk as long as half interorbital width. Cheliped carpus with long, slender spine on upper margin. Pereopods 2–5 little longer than carapace length; dactyli almost straight, flexor margin with few teeth. Male pleonites 3–5 fused, lateral margins straight. Maximum cl. 6 mm (Pl. 52d). Intertidal–shelf (0–134 m). Temperate Northern W Pacific, Central Indo-Pacific. 10 species (Loh and Ng 1999: key to 9 species).

Physachaeus Alcock, 1895 Diagnosis. Carapace with dorsal tubercles. Rostrum comprising 2 equilateral triangular teeth. Eyestalk about as long as interorbital width, almost immovable. Pereopods 2–5 slender, 3–5 times carapace length; all dactyli slender, gently curved, flexor margin dentate. Maximum cl. 11 mm. Slope (350–694 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 2 species (Griffin and Tranter 1986).

Podochela Stimpson, 1860 Diagnosis. Carapace without dorsal spines. Rostrum triangular, thick, with rounded tip. Eyestalk elongate. Pereopods 2–5 about twice carapace length, first longest; dactyli more curved posteriorly. Maximum cl. 22 mm. Subtidal–slope (1–448 m). Tropical W Atlantic, Tropical Eastern Pacific. 4 species (Coelho 2006: rediagnosis, key to 3 species; Rathbun 1925).

Prosphorachaeus Takeda & Miyake, 1969 Metoporhaphis Stimpson, 1860 Diagnosis. Carapace with few gastric and cardiac tubercles. Rostrum spiniform, tapering, as long as or longer than postocular length. Supraorbital eave with postorbital granule. Eyestalk elongate. Pereopods 2–5 slender, 3 times carapace length; meri with distal spine. Maximum cl. 21 mm. Subtidal, shelf (1–90 m). Temperate Northern and Tropical W Atlantic. 1 species (Coelho 2006: in key to genera; Milne-Edwards 1873–1881: figure; Rathbun 1925).

Oncinopus De Haan, 1839 Diagnosis. Carapace without dorsal spines. Rostrum comprising 2 spines or teeth, or comprising 2 blunt spines separated by narrow slit. Interantennular septum weakly developed. Basal antennal article free, distal margin not overlapping eyestalk. Pereopods 2, 3

Diagnosis. Carapace with dorsal tubercles. Rostrum of 2 blunt lobes. Supraorbital eave spinulose, with short acute antorbital spine. Eyestalk elongate. Pereopods 2–5 little longer than carapace length; dactyli of posterior pereopods curved. Male pleonites 3 longer than pleonites 4 and 5, arched, with transverse ridge. Gonopod 1 complex, aperture surrounded by lobes. Maximum cl. 5 mm. Subtidal, shelf (10–91 m). Temperate Northern W Pacific, Central Indo-Pacific. 4 species (Griffin and Tranter 1986: key to 4 species; Takeda and Miyake 1969: diagnosis).

Rhinospinosa Griffin & Tranter, 1986 Diagnosis. Carapace with dorsal spines. Rostrum of 2 blunt lobes. Interantennular spine short with lateral basal spinules. Supraorbital eave with preorbital spine and postorbital spine having

616

Marine Decapod Crustacea

accessory spine. Eyestalk short, straight. Pereopods 2–5 about twice carapace length; all dactyli slender, gently curved, styliform. Maximum cl. 11 mm (Pl. 52e). Shelf, slope (115–350 m). Temperate Northern W Pacific, Central Indo-Pacific. 1 species (Griffin and Tranter 1986).

Sunipea Griffin & Tranter, 1986 Diagnosis. Carapace intestinal region with slender medial spine or low tubercle on posterior margin. Rostrum comprising 2 spines, contiguous over at least half length. Supraorbital eave with preorbital spine, with antorbital lobe (small). Pereopods 2–5 about carapace length; propodi cylindrical. Maximum cl. 14 mm. Shelf (40–155 m). Western and Central Indo-Pacific. 2 species (Griffin 1972: description as Aepinus; Griffin and Tranter 1986: diagnosis; Santana 2015: rediagnosis, second species). References Alcock A (1895) Materials for a carcinological fauna of India. No. 1. The Brachyura Oxyrhyncha. Journal of the Asiatic Society of Bengal 64, 157–291, pls 3–5. Alcock A, Anderson ARS (1896) Illustrations of the zoology of the Royal Indian Marine Surveying Steamer “Investigator”. Crustacea, Part IV. Pls XVI–XXVII. Trustees of the Indian Museum, Calcutta. Barnard KH (1950) Descriptive catalogue of South African decapod Crustacea (crabs and shrimps). Annals of the South African Museum 38, 1–837. Clark PF, Webber WR (1991) A redescription of Macrocheira kaempferi (Temminck, 1836) zoeas with a discussion of the classification of the Majoidea Samouelle, 1819 (Crustacea: Brachyura). Journal of Natural History 25, 1259–1279. doi:10.1080/00222939100770781 Coelho PA (2006) Revisão de Podochela Stimpson e gêneros afins nas costas caribenha e atlântica da América do Sul (Crustacea, Decapoda, Inachidae). Revista Brasileira de Zoologia 23, 678–691. doi:10.1590/ S0101-81752006000300010 Davie PJF (2011) A new species and new records of Oncinopus (Crustacea: Brachyura: Inachidae) from northern Australia. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 27, 123–133. De Man JG (1902) Die von herrn Professor Kükenthal im Indischen Arrchipel gesammelten Dekapoden und Stomatopoden. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 25, 465–929. Forest J (1978) Le genre Macropodia Leach dans les eaux atlantiques européennes (Crustacea Brachyura Majidae). Cahiers de Biologie Marine 19, 323–342. Forest J, Guinot D (1966) Crustacés Décapodes: Brachyoures. Résultats Scientifiques des Campagnes de la Calypso 7, 23–124. Garth JS (1939) New brachyuran crabs from the Galapagos Islands. Allan Hancock Pacific Expeditions 5, 9–29, pls 1–10. Garth JS (1958) Brachyura of the Pacific coast of America. Oxyrhyncha. Allan Hancock Pacific Expeditions 21, 1–854. Griffin DJG (1970) The Australian majid spider crabs of the genus Achaeus (Crustacea, Brachyura). Journal of the Royal Society of Western ­Australia 53, 97–119. Griffin DJG (1972) Brachyura collected by Danish expeditions in southeastern Australia (Crustacea, Decapoda). Steenstrupia 2, 49–90. Griffin DJG (1974) Spider crabs (Crustacea: Brachyura: Majidae) from the International Indian Ocean Expedition, 1963–1964. Smithsonian Contributions to Zoology 182, 1–35. doi:10.5479/si.00810282.182

Griffin DJG, Tranter HA (1986) The Decapoda Brachyura of the Siboga Expedition. Part VIII Majidae. Siboga-Expéditie 39 c4, 1–335. Guinot D (2012) Remarks on Inachoididae Dana, 1851, with the description of a new genus and the resurrection of Stenorhynchinae Dana, 1851, and recognition of the inachid subfamily Podochelinae Neumann, 1878 (Crustacea, Decapoda, Brachyura, Majoidea). Zootaxa 3416, 22–40. doi:10.11646/zootaxa.3416.1.2 Loh LW, Ng PKL (1999) A revision of the spider crabs of the genus Paratymolus Miers, 1879, with descriptions of two new genera and six new species (Crustacea: Decapoda: Brachyura: Majidae). Raffles Bulletin of Zoology 47, 365–407. Macpherson E (1983) Crustáceos Decápodos capturados en las costas de Namibia. Resultados Expediciones Cientificas del Buque Oceanografico Cornide de Saavedra. Supplement Investigación Pesquera 11, 3–80. Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Marco-Herrero E, Torres AP, Cuesta JA, Guerao G, Palero F, et al. (2013) The systematic position of Ergasticus (Decapoda, Brachyura) and allied genera, a molecular and morphological approach. Zoologica Scripta 42, 427–439. doi:10.1111/zsc.12012 Milne-Edwards A (1873–1881) Études sur les xiphosures et les crustacés de la région mexicaine. In Mission scientifique au Mexique et dans l’Amérique Centrale. Recherches zoologiques pour servie a l’histoire de la faune de l’Amerique Centrale et du Mexique. 5ème partie. pp. 1–368, pls 1–29. Imprimerie Nationale, Paris. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Rathbun MJ (1925) The spider crabs of America. Bulletin of the United States National Museum 129, 1–613. Sakai T (1932) Notes on some rare materials of Japanese Oxyrhyncha. Science Reports of the Tokyo Bunrika Daigaku. Section B 4, 41–59. Santana W (2015) A new species of Sunipea Griffin & Tranter, 1986 (Crustacea: Brachyura: Inachidae) and a redescription of Sunipea indicus (Alcock, 1895). Zootaxa 4052, 383–393. doi:10.11646/zootaxa.4052.3.8 Spiridonov VA, Simakova UV, Anosov SE, Zalota AK, Timofeev VA (2020) Review of Macropodia in the Black Sea supported by molecular barcoding data; with the redescription of the type material, observations on ecology and epibiosis of Macropodia czernjawskii (Brandt, 1880) and notes on other Atlanto-Mediterranean species of Macropodia Leach, 1814 (Crustacea, Decapoda, Inachidae). Zoosystematics and Evolution 96, 609–635. Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Takeda M, Miyake S (1969) Crabs from the East China Sea. III Brachygnatha Oxyrhyncha. Journal of the Faculty of Agriculture, Kyushu University 15, 469–521. doi:10.5109/22797

Inachoididae Dana, 1851 Figures 14.83, 14.84, Plate 52f, g A group of 14 genera from the New World, one ranging across the Atlantic, comprise the Inachoididae. Drach and Guinot (1983) advocated family status because of the unique relationship between the carapace and the thoracic sternum in which posterior thoracic pleurites are exposed dorsally and the edge of the carapace sits on them in a gutter. Lemaitre et  al. (2001) supported distinction of inachoidids from other majoids. Guinot and Richer de Forges (1997) went so far as to suggest that the family’s affinities lay not within

14 – Brachyura – crabs

Majoidea but with Hymenosomatidae. This was not the view of Pohle and Marques (2000) whose work with larval stages lead them to doubt their monophyly and placed the genera back as a subfamily of Inachidae. The family status of Inachoididae is now widely recognised. The family and its genera were examined in detail by Santana (2008) but two genera have been added since then (Guinot 2012). Santana’s (2008) phylogeny, based on morphological characters, found the largest genera, Collodes and Euprognatha, to be paraphyletic. Guinot and Van Bakel (2020a, 2020b) recognised seven subfamilies, five of one genus each. They did not allocate the remaining eight genera among the other two subfamilies but discussed potential similarities between them. The arrangement of genera here does not rely on use of subfamilies. Inachoidid spider crabs possess no or few hooked setae and rarely decorate (Guinot and Wicksten 2015).

617

Diagnosis. Orbit absent, eyes unprotected though orbital margin usually with several small spines and postorbital spine; basal antennal article usually extremely slender, more than twice as long as wide, and sometimes free distally. Thoracic pleurites 5–8 visible dorsally, not covered by posterolateral margin of carapace. Eyestalk long, non-retractile or retractile against acute postorbital spine affording no concealment. Male pleotelson formed by fused pleonite 6 and telson in males, fused pleonites 5–6 and telson in females. Pleonite 1 fused to carapace. Maxilliped 3 merus often not as wide as ischium. Implicit generic attributes. Carapace and exposed thoracic pleurites 5–8 granular or tuberculate; subhepatic region ventrolateral margin without anterior spine or tubercle. Postorbital spine triangular; mesial face without row of spines. Antennal basal article ventrolateral margin entire. Pereopods 2–5 dactyli with irregular row of spinules; pereopod 2 about as long as pereopod 3.

Key to genera of Inachoididae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. – 9. –

Rostrum single, longer than carapace. Postorbital spine small, distant from eyestalk (Figs 14.83e, 14.84m). Carapace and exposed thoracic pleurites 5–8 smooth�������������������������������������������Stenorhynchus Rostrum single, shorter than carapace. Postorbital process triangular or cup-shaped, close to eyestalk. Carapace and exposed thoracic pleurites 5–8 granular or tuberculate (Fig. 14.84o)�������������������������������������������������������2 Rostrum wide, blunt, strongly folded downwards as protruding, narrow, beak-shaped extension between antennules, joining anterior epistome process (Figs 14.83b, 14.84n)����������������������������������������������������������������������� Esopus Rostrum triangular or bifid, not expanded ventrally�����������������������������������������������������������������������������������������������������������������3 Pereopod 2 shorter than pereopod 3 (Fig. 14.83c)�����������������������������������������������������������������������������������������������������������������������4 Pereopod 2 as long or longer than pereopod 3 (Fig. 14.83a, d, e)����������������������������������������������������������������������������������������������5 Carapace broadly triangular, regions convex, tuberculate, divided by deep transverse parallel grooves (Fig. 14.83c)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������Paulita Carapace narrower anteriorly, regions indistinct, granular, not divided by parallel grooves (Fig. 14.84l)����������������������������������������������������������������������������������������������������������������������������������������������������������� Paradasygyius Antennal basal article with carina or row of tubercles subparallel to lateral margin of antennular fossa (Fig. 14.84p, r–v)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������6 Antennal basal article without carina subparallel to lateral margin of antennular fossa, sometimes with few tubercles, sometimes with tooth (Fig. 14.84q)��������������������������������������������������������������������������������������������������������������������� 12 Antennal basal article lateral margin with distinct notch in proximal third (Fig. 14.84p). Carapace with 7 prominent knobbed spines (Fig. 14.84a, b)���������������������������������������������������������������������������������������������������������������� Aepinus Antennal basal article lateral margin entire. Carapace with 3 or fewer spines����������������������������������������������������������������������7 Rostrum longer than wide at base, tapering or with distal spine (Figs 14.83a, 14.84h, j)�����������������������������������������������������8 Rostrum not longer than wide, truncate or triangular, sometimes bifid or with medial notch (Fig. 14.84e–g, i)���������������� 9 Carapace without dorsal spines (Fig. 14.84j); subhepatic region ventrolateral margin with anterior spine or tubercle (Fig. 14.84v)��������������������������������������������������������������������������������������������������������������������������������������������������Inachoides Carapace with 3 gastric, 2 branchial, 1 cardiac spines (Figs 14.83a, 14.84h); subhepatic region ventrolateral margin without anterior spine or tubercle����������������������������������������������������������������������������������������������������������������Erileptus Carapace about as wide as long, with 1 gastric, 1 cardiac, 1 intestinal spines: postorbital spine triangular, shorter than hepatic angle (Fig. 14.84f)����������������������������������������������������������������������������������������������������������� Batrachonotus Carapace pyriform, longer than wide, spines or tubercles; Postorbital spine triangular, as long as hepatic angle, or elongate, remote from hepatic margin������������������������������������������������������������������������������������������������������������������10

618

Marine Decapod Crustacea

Fig. 14.83.  Inachoididae. a, Erileptus spinosus Rathbun, 1894; b, Esopus crassus A. Milne-Edwards, 1875; c, Paulita tuberculata (Lemos de Castro, 1949); d, Pyromaia tuberculata (Lockington, 1877); e, Stenorhynchus seticornis (Herbst, 1788).

Carapace postorbital spine elongate, oblique, mesial face with row of small spines (Fig. 14.84e)������������������� Arachnopsis Carapace postorbital spine triangular, mesial face without spines (Fig. 14.84g, i)��������������������������������������������������������������11 Antennal basal article with prominent smooth or crenellate carina subparallel to lateral margin of antennular fossa (Fig. 14.84s). Rostrum triangular, truncate or bifid (Fig. 14.84g)��������������������������������������������Collodes – Antennal basal article with longitudinal row of tubercles subparallel to lateral margin of antennular fossa (Fig. 14.84u). Rostrum bifid, short (Fig. 14.84i)������������������������������������������������������������������������������������������������Euprognatha 12. Carapace subcircular, wider than long; postorbital lobe, cup-shaped, enclosing eyestalk (Fig. 14.84k)��������� Leurocyclus – Carapace pyriform; postorbital spine acute, remote from eyestalk (Figs 14.83d, 14.84c, d)����������������������������������������������13 13. Antennal basal article with strong tooth near midpoint (Fig. 14.84q). Pereopods 2–5 dactyli with 2 rows of setae, one on mesial margin, other lateral��������������������������������������������������������������������������������������������������������������� Anasimus – Antennal basal article without or with few tubercles subparallel to lateral margin of antennular fossa, usually with subterminal marginal tooth (Fig. 14.84w). Pereopod 2–5 dactyli without setae or with setae all around��������������������������������������������������������������������������������������������������������������������������������������������������������������������� Pyromaia 10. – 11.

Aepinus Rathbun, 1897 Diagnosis. Carapace pyriform, with 7 prominent knobbed spines, 2 supraorbital, 1 gastric, 1 cardiac, 2 branchial, and 1 on pleonite 1. Rostrum short, truncate, with medial notch. Postorbital spine

blunt. Antennal basal article with prominent smooth or crenellate carina subparallel to lateral margin of antennular fossa, with anterolateral tooth; lateral margin with distinct notch in proximal third. Maximum cl. 10 mm.

14 – Brachyura – crabs

619

Fig. 14.84.  Inachoididae. Carapace, representative or only species: a, b, Aepinus; c, d, Anasimus; e, Arachnopsis; f, Batrachonotus; g, Collodes; h, Erileptus; i, Euprognatha; j, Inachoides; k, Leurocyclus; l, Paradasygyius; m, Stenorhynchus. n, rostrum, eye, lateral view, Esopus. o, posterior carapace, thoracic pleurites 6–8 (pl6–pl8), pleonites 1–3 (a1–a3), gutter (s.g.), Esopus crassus. Rostrum, epistome, antennal basal articles, eyestalks: p, Aepinus; q, Anasimus; r, Arachnopsis; s, Collodes; t, Erileptus; u, Euprognatha; v, Inachoides (with subhepatic regions); w, Pyromaia.

Shelf (24–175 m). Tropical W Atlantic. 1 species (Santana 2008: species description).

Anasimus A. Milne-Edwards, 1880 Diagnosis. Carapace pyriform or subcircular, with spines and tubercles. Rostrum triangular, as long or much longer than

wide. Antennal basal article with few tubercles, with ventrallydirected tooth near mesial margin. Pereopods 2–5 dactyli with 2 rows of setae, one on mesial margin, other lateral. Maximum cl. 26 mm. Shelf–slope (48–214 m). Tropical W Atlantic. 2 species (Santana 2008: species descriptions).

620

Marine Decapod Crustacea

Arachnopsis Stimpson, 1871

Euprognatha Stimpson, 1871

Diagnosis. Carapace pyriform, with 3 prominent spines, 1 gastric, 1 cardiac, and 1 on pleonite 1. Rostrum short, bifid. Postorbital spine elongate, oblique; mesial face with row of small spines. Antennal basal article with longitudinal row of tubercles subparallel to lateral margin of antennular fossa, with anterolateral tooth. Pereopods 2–5 slender. Maximum cl. 7 mm. Shelf–slope (27–238 m). Tropical W Atlantic. 1 species (Santana 2008: species description).

Diagnosis. Carapace pyriform, with tubercles, sometimes spines. Rostrum short, bifid. Antennal basal article with longitudinal row of tubercles subparallel to lateral margin of antennular fossa, with anterolateral tooth. Pereopod 2 longer than pereopod 3. Maximum cl. 15 mm. Subtidal–slope (17–730 m). Tropical W Atlantic, Tropical Eastern Pacific. 6 species (Santana 2008: species descriptions, key to species).

Batrachonotus Stimpson, 1871

Inachoides H. Milne Edwards & Lucas, 1842

Diagnosis. Carapace pyriform, about as wide as long, with 3 short spines, 1 gastric, 1 cardiac, 1 intestinal. Rostrum triangular, with medial notch. Postorbital spine triangular, shorter than hepatic angle. Antennal basal article with longitudinal row of tubercles subparallel to lateral margin of antennular fossa, with anterolateral tooth. Pereopod 2 longer than pereopod 3 (in male). Maximum cl. 9 mm. Subtidal–slope (12–247 m). Tropical W Atlantic. 1 species (Santana 2008: species description).

Diagnosis. Carapace pyriform, without dorsal spines; subhepatic region ventrolateral margin with anterior spine or tubercle. Rostrum triangular, tapering to distal spinulose spine. Postorbital spine cup-like, almost enclosing eyestalk. Antennal basal article with longitudinal row of tubercles subparallel to lateral margin of antennular fossa, with anterolateral tooth. Pereopods 2–5 almost prehensile, dactyli curved. Maximum cl. 20 mm. Subtidal, shelf (1–102 m). Tropical W Atlantic, Tropical Eastern Pacific, Temperate South America (Chile). 3 species (Garth 1958: key to Pacific species; Guinot and Van Bakel 2020b: rediagnosis as Inachoidinae; Santana 2008: descriptions of 2 species).

Collodes Stimpson, 1860 Diagnosis. Carapace pyriform or subcircular, with tubercles, sometimes spines. Rostrum truncate, triangular or bifid. Antennal basal article with prominent smooth or crenellate carina subparallel to lateral margin of antennular fossa, with anterolateral tooth. Maximum cl. 30 mm. Subtidal–slope (7–683 m). Tropical W Atlantic, Tropical Eastern Pacific. 16 species (Colavite et  al. 2019: new species; Garth 1958: key to 5 Pacific species; Guinot and Van Bakel 2020b: diagnosis as Collodinae; Rathbun 1925: key to 14 species; Santana 2008: species descriptions, key to 14 species).

Erileptus Rathbun, 1894 Diagnosis. Carapace pyriform, with 3 gastric, 2 branchial, 1 cardiac spines. Rostrum triangular, tapering to distal spinulose spine. Postorbital spine elongate, oblique. Antennal basal article with longitudinal row of tubercles subparallel to lateral margin of antennular fossa, with anterolateral tooth. Pereopods 2–5 slender. Maximum cl. 11.5 mm. Subtidal–slope (4–549 m). Tropical Eastern Pacific. 1 species (Garth 1958; Santana 2008: species description).

Esopus A. Milne-Edwards, 1875 Diagnosis. Carapace suboval, longer than wide; regions inflated, tuberculate. Rostrum wide, blunt, strongly folded downwards as protruding, narrow, beak-shaped extension between antennules, joining anterior epistome process. Postorbital spine thick, cupshaped, fused with rounded subhepatic region. Antennal basal article without carina. Maximum cl. 13 mm. Slope (201–359 m). Tropical W Atlantic. 1 species (Guinot and Van Bakel 2020b).

Leurocyclus Rathbun, 1897 Diagnosis. Carapace subcircular, wider than long. Rostrum triangular, small. Postorbital spine cup-like, curving around eyestalk. Antennal basal article without carina. Maximum cl. 60 mm. Subtidal, shelf (18–166 m). Tropical W Atlantic, Temperate South America (Atlantic coast). 2 species (Guinot and Van Bakel 2020b: diagnosis as Salaciinae; Santana 2008: species descriptions).

Paradasygyius Garth, 1958 Diagnosis. Carapace subpyriform, flat, granular. Rostrum triangular, small. Postorbital spine cup-like, curving around eyestalk. Antennal basal article with longitudinal tuberculate carina subparallel to lateral margin of antennular fossa, with anterolateral tooth. Pereopod 2 shorter than pereopod 3. Maximum cl. 27 mm. Subtidal, shelf (1–200 m). Tropical W Atlantic, Tropical Eastern Pacific. 1 species (Guinot and van Bakel 2020a: rediagnosis as Paradasygyiinae; Santana 2008: species description).

Paulita Guinot, 2012 Diagnosis. Carapace subpyriform, regions convex, tuberculate, divided by deep transverse parallel grooves. Rostrum broadly triangular. Postorbital spine cup-like, curving around eyestalk. Antennal basal article with longitudinal row of tubercles subparallel to lateral margin of antennular fossa, with anterolateral tooth. Pereopod 2 shorter than pereopod 3. Maximum cl. 21 mm. Subtidal, shelf (10–50 m). Tropical W Atlantic. 1 species (­Guinot 2012: diagnosis; Guinot and Van Bakel 2020b: rediagnosis as Paulitinae).

14 – Brachyura – crabs

Pyromaia Stimpson, 1871 Diagnosis. Carapace pyriform, with tubercles, sometimes spines. Rostrum triangular, as long as or much longer than wide, or triangular, tapering to distal spinulose spine. Postorbital spine cuplike, curving around eyestalk. Antennal basal article without or with few tubercles subparallel to lateral margin of antennular fossa, usually with subterminal marginal tooth. Pereopods 2–5 dactyli without setae or with setae all around; pereopod 2 longer than pereopod 3. Maximum cl. 41 mm (pl. 52f). Subtidal–slope (1–412 m). Temperate Northern and Tropical Atlantic, Tropical Eastern Pacific, Temperate South America, Temperate Australasia. 7 species (Lemaitre et al. 2001). Pyromaia tuberculata (Lockington, 1877) is native to the Tropical E Pacific but has been introduced and become locally abundant in Japan (Furota 1990; Furota and Furose 1988), Australia (Poore 2004), New Zealand (Webber and Wear 1981), Brazil and Argentina (Schejter et al. 2002).

Stenorhynchus Lamarck, 1818 arrow crabs Diagnosis. Carapace triangular, longer than wide; and exposed thoracic pleurites 5–8 smooth. Rostrum slender, flat, longer than postorbital carapace. Postorbital spine small, distant from orbit. Pereopods 2–5 more than 3 times length of carapace including rostrum. Maximum cl. 38 mm (Pl. 52g). Subtidal–slope (1–1465). Temperate Northern and Tropical Atlantic, Tropical Eastern Pacific, Temperate South America (Atlantic coast). 4 species (Goeke 1989: species description; Guinot 2012: distribution of 4 species; Guinot and Van Bakel 2020b: rediagnosis as Stenorhynchinae; Monod 1956: figures). Species of Stenorhynchus, known as arrow crabs, are popular in the ornamental aquarium trade (Monteforte-Sánchez et al. 2018). References Colavite J, Windsor A, Santana W (2019) Three new species and a new genus of majoid crabs from the eastern Pacific (Decapoda, Brachyura). ZooKeys 825, 1–24. doi:10.3897/zookeys.825.32271 Drach P, Guinot D (1983) Les Inachoididae Dana, famille de Majoidea caractérisée par des connexions morphologiques d’un type nouveau entre carapace, pleurites, sternites, pléon. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 297, 37–42. Furota T (1990) Population structure of the introduced spider crab Pyromaia tuberculata in the uppermost region of Tokyo Bay. Benthos Research 1990, 1–7. doi:10.5179/benthos1990.1990.39_1 Furota T, Furose K (1988) Distribution of the introduced spider crab, Pyromaia tuberculata, along the coast of Japan. Benthos Research 1988, 75–78. doi:10.5179/benthos1981.1988.75 Garth JS (1958) Brachyura of the Pacific coast of America. Oxyrhyncha. Allan Hancock Pacific Expeditions 21, 1–854. Goeke GD (1989) Stenorhynchus yangi, a new Western Atlantic species of arrow crab (Crustacea, Brachyura, Majidae) and a redescription of S. seticornis (Herbst, 1788). Proceedings of the Biological Society of Washington 102, 620–636. Guinot D (2012) Remarks on Inachoididae Dana, 1851, with the description of a new genus and the resurrection of Stenorhynchinae Dana,

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1851, and recognition of the inachid subfamily Podochelinae Neumann, 1878 (Crustacea, Decapoda, Brachyura, Majoidea). Zootaxa 3416, 22–40. doi:10.11646/zootaxa.3416.1.2 Guinot D, Richer de Forges B (1997) Affinités entre les Hymenosomatidae MacLeay, 1838 et les Inachoididae Dana, 1851 (Crustacea, Decapoda, Brachyura). Zoosystema 19, 453–502. Guinot D, Van Bakel B (2020a) Diagnosis of the inachoidid subfamily Paradasygyiinae subfam. nov. (Crustacea, Decapoda, Brachyura, Majoidea). Zootaxa 4801, 198–200. doi:10.11646/zootaxa.4801.1.13 Guinot D, Van Bakel B (2020b) Extraordinary majoid crabs: the genus Esopus A. Milne-Edwards, 1875 in the new subfamily Esopinae subfam. nov., and erection of Paulitinae subfam. nov. (Crustacea, Decapoda, Brachyura, Majoidea, Inachoididae Dana, 1851). Zootaxa 4766, 101–127. doi:10.11646/zootaxa.4766.1.5 Guinot D, Wicksten MK (2015) Chapter 71–11. Camouflage: carrying behaviour, decoration behaviour, and other modalities of concealment in Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 583–638. Brill, Leiden. Lemaitre R, Campos NH, Bermúdez A (2001) A new species of Pyromaia from the Caribbean Sea, with a rediscription of P. propinqua Chace, 1940 (Decapoda: Brachyura: Majoidea: Inachoididae). Journal of Crustacean Biology 21, 760–773. doi:10.1163/20021975-99990172 Monod T (1956) Hippidea et Brachyura ouest-africains. Mémoires de I’Institut Français d’Afrique Noire 45, 1–674. Monteforte-Sánchez M, Monsalvo-Spencer P, Robles-Villegas G, Reynoso-Granados T (2018) Profile analysis on the ornamental marine yellow-arrow spider crab Stenorhynchus debilis (Smith 1871) into experimental studies for live management and cultivation in laboratory. Thalassas: An International Journal of Marine Sciences 34, 113–129. doi:10.1007/s41208-017-0044-z Pohle G, Marques FPL (2000) Larval stages of Paradasygyius depressus (Bell, 1835) (Crustacea: Decapoda: Brachyura: Majidae) and a phylogenetic analysis for 21 genera of Majidae. Proceedings of the Biological Society of Washington 113, 739–760. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Rathbun MJ (1925) The spider crabs of America. Bulletin of the United States National Museum 129, 1–613. Santana WRA (2008) Revisão taxonômica e relações filogenéticas em Inachoididae Dana, 1851 (Crustacea, Brachyura, Majoidea). Thesis. University of São Paulo, São Paulo. Schejter L, Spivak ED, Luppi TA (2002) Presence of Pyromaia tuberculata (Lockington, 1877) adults and larvae in the Argentine continental shelf (Crustacea: Decapoda: Majoidea). Proceedings of the Biological Society of Washington 115, 605–610. Webber WR, Wear RG (1981) Life history studies on New Zealand Brachyura 5. Larvae of the family Majidae. New Zealand Journal of Marine and Freshwater Research 15, 331–383. doi:10.1080/00288330.1981.9515929

Majidae Samouelle, 1819 decorator crabs Figures 14.85–14.88, Plates 53, 54a–c Majidae were formerly a massive and morphologically diverse family of multiple subfamilies (Griffin and Tranter 1986a), most of which are now treated as separate families (Ng et  al. 2008). As a result, many of the crabs classified

622

Marine Decapod Crustacea

under Majidae in various subfamilies in earlier works can be found in this work under the corresponding upraised families. Majidae are now restricted to the majoid crabs having well developed orbits, with the eyes protected from above, but not below. The well developed orbits of majid decorator crabs surround the deeply set eyestalks, differing from the closely related majoid family, Mithracidae, in having the eye not protected below by an expanded basal antennal article. Unlike mithracids which are largely confined to tropical and subtropical Americas down to ~100 m depth, majids occur globally. Most live at subtidal to upper shelf depth but some species extend on to the upper slope. One deepwater species, Teratomaia richardsoni (Dell, 1960), possibly reaches hadal depths (Griffin and Tranter 1986b). Most, but not all, majid species are classic decorators, collecting algae, sponges, bryozoans etc. to attach to the hooked setae on the carapace and limbs (Guinot and Wicksten 2015; Hultgren and Stachowicz 2011). Identification to genus and species often involves cleaning algae and epifauna away from the dorsal surface of the carapace, rostral spines and orbit to see details of ornamentation. Our key is based essentially on that in Griffin and Tranter (1986a) who treated the family as subfamily

Majinae. Their key to Indo-West Pacific genera has been updated to include the subdivision of Maja (Ng and Richer de Forges 2015) and to include American genera. Diagnosis. Carapace with orbits present, formed dorsally by supraorbital eave, adjacent spines and postorbital spine or lobe.; basal antennal article not expanded laterally to form floor of orbit (at most twice as long as wide), usually with anterior angles spine-like. Thoracic pleurites 5–8 not visible dorsally, covered by posterolateral margin of carapace. Male pleon almost uniform in width, pleonite 3 only slightly wider than pleonite 6, telson not inserted in pleonite 6. Pleonite 1 free from carapace. Maxilliped 3 merus usually about as wide as ischium. Gonopod 1 slender, weakly curved, aperture usually subterminal, usually protected by lobes, apex usually slender. Implicit generic characters. Carapace pyriform. Orbit open ventrally; antennal basal article widely separated from postorbital lobe; intercalated spine present; postorbital lobe cupped; cornea of retracted eye protected on dorsal and ventral surfaces or dorsal surface only. Eyestalk cylindrical, short, less than half length exposed in dorsal view. Antennal peduncular articles 4, 5 slender, article 4 longer than wide. Cheliped visible in dorsal view.

Key to genera of Majidae 1. – 2. – 3.

– 4. – 5. – 6. – 7.

Carapace longitudinally suboval, front flat (Fig. 14.85k), or subtriangular, wider than long (Fig. 14.85c). Margins expanded, lamellate. Pereopods 2–5 carinate. Antennal basal article with distal lobe, or more distal articles lamellate, visible in dorsal view (Fig. 14.87d)�������������������������������������������������������������������������������������������������2 Carapace pyriform. Margins not expanded or lamellate. Pereopods 2–5 rarely carinate. Antennal basal article rarely visible in dorsal view, more distal articles cylindrical�����������������������������������������������������������������������������������3 Carapace longitudinally suboval, front flat. Antennal articles 4, 5 lamellate, visible in dorsal view (Fig. 14.85k). Chelipeds not visible in dorsal view�������������������������������������������������������������������������������������������� Planotergum Carapace subtriangular, wider than long (Fig. 14.85c). Antennal basal article with distal lobe visible in dorsal view (Fig. 14.87d). Chelipeds massive������������������������������������������������������������������������������������������������ Eurynolambrus Eyestalk dorsally flattened or lobed, not retractile against postorbital lobe, with lobe on anterior margin. Supraorbital eave weakly expanded, barely covering base of eyestalk. Carapace with numerous blunt compound tubercles (Fig. 14.86m). Antennal peduncular article 4 flat, broad; article 5 laterally directed (Fig. 14.87t)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Pippacirama Eyestalk not dorsally flattened or lobed. Supraorbital eave longer than basal width of eyestalk, not curving over it. Carapace smooth or spinose or with tubercles or with plates. Antennal peduncular article 5 in line with article 4�������������������������������������������������������������������������������������������������������������������������������������������������������������������������4 Eyestalks long, more than two-thirds exposed in dorsal view (Fig. 14.85a, f, l)���������������������������������������������������������������������5 Eyestalks short, less than half length exposed in dorsal view�������������������������������������������������������������������������������������������������17 Carapace intercalated spine absent (Fig. 14.87v)�������������������������������������������������������������������������������������������������Pseudomicippe Carapace intercalated spine present (Fig. 14.87b, m)������������������������������������������������������������������������������������������������������������������6 Rostral spines with lateral tooth. Supraorbital eave with preorbital spine and adjacent spine on supraorbital eave (Fig. 14.85f)������������������������������������������������������������������������������������������������������������������������������������������������������������Maiopsis Rostral teeth without lateral tooth. Preorbital spine absent. Supraorbital eave smooth (Fig. 14.85a, i)�����������������������������7 Carapace smooth, branchial margins unarmed������������������������������������������������������������������������������������������������������ Anacinetops

14 – Brachyura – crabs

623

Fig. 14.85.  Majidae. a, Anacinetops stimpsoni Miers, 1879; b, Cyclax suborbicularis (Stimpson, 1858); c, Eurynolambrus australis H. Milne Edwards & Lucas, 1841; d, Jacquinotia edwardsii (Jacquinot, 1853); e, Leptomithrax gaimardii (H. Milne Edwards, 1834); f, Maiopsis panamensis Faxon, 1893; g, Majella brevipes Ortmann, 1893; h, Naxia aurita (Latreille, 1825); i, Ovimaja compressipes (Miers, 1879); j, Planaja plana Ng & Richer de Forges, 2015; k, Planotergum mirabile Balss, 1935; l, Pseudomicippe varians Miers, 1879.

624

– 8. – 9. – 10. – 11.

Marine Decapod Crustacea

Carapace tuberculate, branchial margins spinous����������������������������������������������������������������������������������������������������������������������8 Antennal flagellum just outside orbit, near rim (Fig. 14.88a)��������������������������������������������������������������������������������Rathbunaja Antennal flagellum inside orbit (Fig. 14.88b)������������������������������������������������������������������������������������������������������������������������������9 Pereopods 2–5 meri upper margins with subdistal spine (Figs 14.88e)���������������������������������������������������������������������������������10 Pereopods 2–5 meri upper margins unarmed (Fig. 14.88d)����������������������������������������������������������������������������������������������������11 Antennal basal article as long as wide. Epistome as long as wide (Fig. 14.87p)��������������������������������������������������������Neomaja Antennal basal article longer than wide. Epistome longer than wide (Fig. 14.87r)�����������������������������������������������Paramaya Suborbital margin confluent with margin of postorbital tooth, not demarcated by fissure or cleft (Fig. 14.87q)������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 12 – Suborbital margin separated from margin of postorbital tooth by deep fissure (Fig. 14.87n)�������������������������������������������14 12. Carapace covered with small uniform granules (Fig. 14.86o). Gonopod 1 sinuous, with numerous long setae on distal surface������������������������������������������������������������������������������������������������������������������������������������������������������Sakaija – Carapace covered with numerous heterogeneous granules. Gonopod 1 straight or curved, with scattered setae on distal surface (Fig. 14.88h, m)����������������������������������������������������������������������������������������������������������������������������������13 13. Rostral spines short, stout (Fig. 14.87h). Branchial regions of adults swollen laterally and dorsally; cardiac region swollen; regions poorly demarcated. Lateral carapace margin of adults without spines. Gonopod 1 long, mostly straight, distal part conical (Fig. 14.88m)�������������������������������������������������������������������������������������� Paramaja – Rostral spines long, slender (Fig. 14.87n). Branchial regions moderately swollen; cardiac region gently convex; regions clearly demarcated. Lateral carapace margin of adults with spines. Gonopod 1 short, curved, with ventral projection (Fig. 14.88h)����������������������������������������������������������������������������������������������������������Alcomaja 14. Antennal basal article as wide as long (Fig. 14.87n). Gonopod 1 long, with distal pad-like folds (Fig. 14.88k)�������������������������������������������������������������������������������������������������������������������������������������������������������������������������� Maja – Antennal basal article longer than wide (Fig. 14.87s). Gonopod otherwise��������������������������������������������������������������������������15 15. Carapace ovoid, longer than wide (Fig. 14.85i). Male thoracic sternites 1–4 not elongated, lateral junction between sternites 2 and 3 deep, forming waist-like structure. Male sterno-abdominal cavity margin almost flat, without rim. Gonopod 1 strongly bent, distal part dilated with 2 folds (Fig. 14.88l)������������������� Ovimaja – Carapace pyriform (Figs 14.85j, 14.86g). Male thoracic sternites 1–4 elongate, lateral junction between sternites 2 and 3 shallow. Male sterno-abdominal cavity margin with rim on thoracic somite 4 (Fig. 14.88c). Gonopod 1 otherwise����������������������������������������������������������������������������������������������������������������������������������������16 16. Carapace lateral margin with 3 spines (Fig. 14.86g). Supraocular eave prominent, anterior part rounded (Fig. 14.87f). Gonopod 1 stout, straight or slightly bent, distal part simple, with spinules and short setae (Fig. 14.88j)��������������������������������������������������������������������������������������������������������������������������������������������������������������������Holthuija – Carapace lateral margin with 6 spines (Fig. 14.85j). Supraocular eave narrow (Fig. 14.87i). Gonopod 1 slender, curved, distal part sharply tapering, margins chitinised, with numerous short spinules (Fig. 14.88n)���������������������������������������������������������������������������������������������������������������������������������������������������������������������� Planaja 17. Antennal flagellum arising within orbit or just outside orbit, near rim (Fig. 14.88a, b)�����������������������������������������������������18 – Antennal flagellum excluded from orbit by antennal basal article process (Fig. 14.87j–x)����������������������������������������������� 22 18. Carapace intercalated spine absent, short (nearly obsolete) or close to eave and postorbital lobe; branchial margin with simple blunt or mushroom-shaped tubercles or large plates (Fig. 14.86d, h, 14.87e)������������������������������19 – Carapace intercalated spine well developed, clearly demarcated from eave and postorbital lobe; branchial margin lined with spines (Fig. 14.85g)������������������������������������������������������������������������������������������������������������������������� Majella 19. Chelipeds of male ~3 times carapace length. Carapace intercalated spine absent (Fig. 14.86h)������������������������������Kasagia – Chelipeds of male about twice carapace length. Carapace intercalated spine present, even if small (Fig. 14.86a, d, f)����������������������������������������������������������������������������������������������������������������������������������������������������������������������� 20 20. Carapace tubercles fused into 10–12 large tuberculate or spinulate plates, small tubercles over surface. Supraorbital eave margin denticulate (Fig. 14.86a)�����������������������������������������������������������������������������������������������Seiitaoides – Carapace tubercles numerous, distinct, few fused to form larger plates. Supraorbital eave margin smooth or with few tubercles����������������������������������������������������������������������������������������������������������������������������������������������������������������������21

14 – Brachyura – crabs

625

Fig. 14.86.  Majidae. a, Seiitaoides orientalis (Sakai, 1961); b, Temnonotus granulosus A. Milne-Edwards, 1875. Carapace: c, Ageitomaia; d, Choniognathus; e, Entomonyx; f, Eurynome; g, Holthuija; h, Kasagia; i, Kimbla; j, Microhalimus; k, Notomithrax; l, Paraentomonyx; m, Pippacirama; n, Prismatopus; o, Sakaija; p, Schizophroida; q, Schizophrys; r, Teratomaia; s, Tumulosternum.

21.

Carapace tubercles mushroom-shaped with crenulate edges, sometimes fused into plates. Hepatic margin with flat triangular plate or with large scutellate or mushroom-shaped tubercle (Fig. 14.86f). Gonopod 1 with 1 or 2 recurved subdistal lobes and long setae��������������������������������������������������������������������������������������������� Eurynome – Carapace tubercles blunt or pointed but always simple, neither flat nor mushroom-shaped. Hepatic margin with broad blunt tuberculate lobe or 1 or 2 sharp processes (Fig. 14.86d). Gonopod 1 without recurved lobes or long setae, aperture simple and subterminal (Fig. 14.88i)��������������������������������������������������������������Choniognathus 22. Carapacce cardiac region surrounded except anteriorly by deep trench (Fig. 14.86b). Caribbean�����������������Temnonotus

626

Marine Decapod Crustacea

Fig. 14.87.  Majidae. Anterior dorsal carapace, representative or only species: a, Alcomaja; b, Anacinetops; c, Cyclax; d, Eurynolambrus; e, Eurynome; f, Holthuija; g, Jacquinotia; h, Paramaja; i, Planaja. Antennal basal article, distal articles, eyestalk, epistome (antennal gland shaded): j, Cyclax; k, Eurynome; l, Jacquinotia; m, Maiopsis; n, Maja; o, Naxia; p, Neomaja; q, Paramaja; r, Paramaya; s, Paramithrax; t, Pippacirama; u, Planotergum; v, Pseudomicippe; w, Schizophroida; x, Tumulosternum. Features labelled on a, p: aba, antennal basal article; ao, antorbital spine; i, intercalated spine; n, nephridiopore (antennal gland); po, preorbital spine; ps, postorbital spine; r, rostral spine.

– 23. – 24.

Carapace cardiac region not surrounded by deep trench. Indo-West Pacific��������������������������������������������������������������������� 23 Orbit open ventrally; antennal basal article widely separated from postorbital lobe (Fig. 14.87o)���������������������������������� 24 Orbit closed ventrally; antennal basal article in broad contact with postorbital lobe (Fig. 14.87i)���������������������������������� 36 Postorbital lobe a conical to acuminate spine, not cupped; cornea of retracted eye not protected on dorsal or ventral surfaces (Fig. 14.87s)���������������������������������������������������������������������������������������������������������������������������������������������� 25 – Postorbital lobe cupped anteriorly; cornea of retracted eye protected on dorsal and ventral surfaces or dorsal surface only (Fig. 14.87w)������������������������������������������������������������������������������������������������������������������������������������������� 29 25. Carapace gastric, cardiac and intestinal regions smooth. Antennal basal article apically notched, distolateral tooth short, rounded crenulate (Fig. 14.87s)��������������������������������������������������������������������������������� Paramithrax – Carapace gastric, cardiac and intestinal regions with tubercles and/or short spines. Antennal basal article not notched, distolateral angle spinelike������������������������������������������������������������������������������������������������������������������������������ 26 26. Preorbital angle acute. Male cheliped merus and carpus of male with upper crest of ~6 flat lobes/spines (Fig. 14.86c). Gonopod 1 aperture protected by a distally directed flap (Fig. 14.88g)�����������������������������������Ageitomaia

14 – Brachyura – crabs

– 27. – 28. – 29. – 30. – 31. – 32. – 33. – 34. –

627

Preorbital angle with or without blunt spine. Male cheliped merus with upper ridge entire or with 2–5 spines, outer lower margin entire or with tubercles. Gonopod 1 aperture margins lobe-like������������������������������������� 27 Antennal basal article broad, with 2 distal spines; articles 4, 5 stout, article 4 about as wide as long����������� Notomithrax Antennal basal article narrow, with distolateral spine only; articles 4, 5 slender, article 4 longer than wide (Fig. 14.87o)������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 28 Pereopods 2–5 propodi flexor margins distally expanded (Figs 14.85h, 14.88f). Carapace with at least 1 mesogastric tubercle, surface bare or with few short setae between groups of strong curled setae. Intercalated spine present (sometimes almost fused to eave) (Fig. 14.87o)���������������������������������������������������������������Naxia Pereopods 2–5 propodi flexor margins cylindrical. Carapace without mesogastric tubercle, surface tomentose and with groups of strong curled setae. Intercalated spine absent (Fig. 14.86j)���������������������� Microhalimus Preorbital angle smooth, rounded���������������������������������������������������������������������������������������������������������������������������������������������� 30 Preorbital angle produced as spine or tubercle��������������������������������������������������������������������������������������������������������������������������33 Rostral spines with 1 or 2 lateral accessory spines (Fig. 14.86q)����������������������������������������������������������������������������Schizophrys Rostral spines without lateral accessory spine���������������������������������������������������������������������������������������������������������������������������31 Carapace densely spinose or tuberculate (Fig. 14.85e). Maximum cl. 165 mm������������������������������������������������Leptomithrax Carapace with few blunt tubercles. Maximum cl. 37 mm�������������������������������������������������������������������������������������������������������32 Antennal basal article narrow, with 2 slender distal spines (Fig. 14.87w). Carapace intercalated spine larger than antorbital spine, not excluded from margin of orbit; branchial width ~1.5 times hepatic width (Fig. 14.86p)�����������������������������������������������������������������������������������������������������������������������������������������������Schizophroida Antennal basal article broad, with 2 short, blunt, distal spines (Fig. 14.87x). Carapace intercalated spine much smaller than antorbital spine or obsolete, excluded from margin of orbit; branchial width about twice hepatic width (Fig. 14.86s)�������������������������������������������������������������������������������������������������������������������� Tumulosternum Carapace suborbicular; preorbital angle with blunt tubercle; ostorbital lobe with accessory dorsal basal tooth (Fig. 14.85b). Rostrum short, 0.05–0.1 postrostral carapace length���������������������������������������������������������������Cyclax Carapace pyriform or suboval; preorbital angle with spine; postorbital lobe without spine on anterior edge. Rostrum 0.2 or more postrostral carapace length�������������������������������������������������������������������������������������������������������������� 34 Carapace suboval, dorsally with tubercles and few small spines (Fig. 14.86i); hepatic and branchial margins with tubercles��������������������������������������������������������������������������������������������������������������������������������������������������������������������Kimbla Carapace pyriform, with spines at least in midline; hepatic and branchial margins with spines or lobes����������������������35

Fig. 14.88.  Majidae. Right orbit, antennal basal article, eyestalk, dorsal view: a, Rathbunaja; b, Sakaija. c, male thoracic sternum, pleon, Planaja. Pereopod 2 merus: d, Maja; e, Paramaya. f, pereopod 2, Naxia. Gonopod 1 apex: g, Ageitomaia; h, Alcomaja; i, Choniognathus; j, Holthuija; k, Maja. Gonopod 1: l, Ovimaja; m, Paramaja; n, Planaja.

628

Marine Decapod Crustacea

35.

Rostral spines ~0.2 postrostral carapace length. Carapace branchial margin with 4–6 smooth sharp spines (last subdorsal) (Fig. 14.86r). Cheliped merus and carpus with spinous tubercles; male chela long, slender (palm ~4 times as long as high)��������������������������������������������������������������������������������������������������������������� Teratomaia – Rostral spines longer than 0.3 postrostral carapace length. Carapace branchial margin with 1–3 sharp or knobbed spines (last subdorsal), sometimes with anterior lamellate lobe (Fig. 14.86n). Cheliped merus and carpus usually with longitudinal ridges; male chela short, stout (palm ~1.5 times as long as high)�������������������������������������������������������������������������������������������������������������������������������������������������������������Prismatopus 36. Carapace branchial margin with tubercles; supraorbital eave with rounded antorbital angle, intercalated spine, postorbital spine almost fused. Rostral lobes 0.1 postrostral carapace length, broad, fused for most of length (Fig. 14.87g, l)����������������������������������������������������������������������������������������������������������������������������������Jacquinotia – Carapace branchial margin with at least 2 long spines; supraorbital eave strongly expanded to form tubular orbit. Rostral spines narrow, separate, 0.2–0.25 postrostral carapace length�����������������������������������������������������������������37 37. Carapace branchial margin with 2 spines; posterior margin with 1 spine (Fig. 14.86e)�������������������������������������Entomonyx – Carapace branchial margin with 4 spines; posterior margin with 2 spines (Fig. 14.86l)����������������������������Paraentomonyx Ageitomaia Griffin & Tranter, 1986 Diagnosis. Carapace tuberculate, with 2 mesogastric spines, pair of mesocardiac spines, 1 intestinal spine; Hepatic margin with 2 spines; branchial margin with 3 spines. Rostral spines divergent, at right angles. Preorbital angle acute; postorbital lobe spine-like, not cupped; cornea of retracted eye not protected on dorsal or ventral surfaces. Antennal flagellum excluded from orbit by antennal basal article process. Cheliped merus and carpus of male with upper crest of ~6 flat lobes/spines. Gonopod 1 aperture protected by a distally directed flap. Maximum cl. 21 mm. Shelf (59–119 m). Temperate South America (Juan Fernandez Islands, Chile). 1 species (Balss 1922; Garth 1958: as Paramithrax baeckstromi (Balss, 1923)).

Alcomaja Ng & Richer de Forges, 2015 Diagnosis. Carapace suboval, covered with numerous heterogeneous granules, branchial regions moderately swollen; cardiac region gently convex; regions clearly demarcated; branchial margins spinous. Rostral spines long, slender. Suborbital margin confluent with margin of postorbital spine, not demarcated by fissure or cleft; preorbital spine absent. Eyestalk long, more than twothirds exposed in dorsal view. Antennal basal article longer than wide; flagellum arising within orbit. Epistome as long as wide. Cheliped carpus, propodus smooth. Pereopods 2–5 meri upper margins unarmed. Gonopod 1 short, curved, with ventral projection, with scattered setae on distal surface. Maximum cl. 52 mm (Pl. 53a). Intertidal–slope (0–500 m). Western and Central Indo-Pacific. 7 species (Ng and Richer de Forges 2015: diagnosis, key to species).

Anacinetops Miers, 1879 Diagnosis. Carapace smooth; branchial margins unarmed. Rostral spines partly fused, short, triangular. Intercalated spine similar to postorbital spine. Eyestalk long, more than two-thirds exposed in dorsal view. Antennal basal article with anterolateral tooth directed laterally. Maximum cl. 26 mm.

Subtidal–shelf (3–36 m). Central Indo-Pacific, Temperate S Australia. 1 species (Baker 1905: description figures as Paramicippa hispida Baker, 1905; Griffin and Tranter 1974: comment).

Choniognathus Rathbun, 1932 Diagnosis. Carapace pyriform or suboval, with blunt or pointed tubercles; branchial margins with blunt or pointed tubercles; hepatic margin with broad blunt tuberculate lobe or 1 or 2 sharp processes. Rostral spines separated by U-shaped hiatus. Intercalated spine short (nearly obsolete) or close to eave and postorbital lobe. Antennal flagellum arising within orbit. Cheliped of male carpus, propodus spinulose. Maxilliped 3 ischium and merus fused. Gonopod 1 1 without recurved lobes or long setae, aperture simple and subterminal. Maximum cl. 25 mm. Shelf, slope (50–540 m). Western and Central Indo-Pacific, Temperate S Australia. 45 species (Griffin and Tranter 1986a: rediagnosis, key to 4 species; Richer de Forges et al. 2021: diagnosis, discussion and figures of species).

Cyclax Dana, 1851 Diagnosis. Carapace suborbicular, with few blunt tubercles; branchial margins spinose. Rostral spines short, 0.05–0.1 postrostral carapace length. Preorbital angle with tubercle; postorbital lobe with accessory dorsal basal tooth. Antennal flagellum excluded from orbit by antennal basal article process. Maximum cl. 46 mm (Pl. 53b). Intertidal, subtidal (0–6 m). Indo-West Pacific. 2 species (Forest and Guinot 1961: species differentiation, figures; Griffin and Tranter 1986a: key to species).

Entomonyx Miers, 1884 Diagnosis. Carapace with few spines in midline; branchial margin with 2 spines; posterior margin with 1 spine. Rostral spines separate from base, divergent. Orbit closed ventrally; antennal basal article in broad contact with postorbital lobe; supraorbital eave

14 – Brachyura – crabs

strongly expanded to form tubular orbit. Antennal flagellum excluded from orbit by antennal basal article process. Maximum cl. 22 mm. Shelf (60–150 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 1 species (Griffin and Tranter 1986a: key to 2 species, now Entomonyx and Paraentomonyx).

Eurynolambrus H. Milne Edwards & Lucas, 1841 Diagnosis. Carapace broadly subtrianglar, eroded, without spines,1.6 times as wide as long, depressed, vaulted, concealing pereopods; branchial margin lamellate, expanded, scalloped. Rostral spines partly fused, short, triangular. Antennal basal article broad proximally, projecting as lobe visible in dorsal view. Pereopods 2–5 flat, carinate. Maximum cl. 25 mm (Pl. 53c). Intertidal–shelf (0–92 m). Temperate Australasia (New Z ­ ealand endemic). 1 species (Griffin 1966).

Eurynome Leach, 1814 Diagnosis. Carapace with spines and large mushroom-shaped tubercles with crenulate edges, sometimes fused into plates; branchial margins with blunt or mushroom-shaped tubercles; hepatic margin with flat triangular plate or with large scutellate or mushroom-shaped tubercle. Rostral spines separated by U-shaped hiatus. Preorbital angle smooth, rounded; intercalated spine short (nearly obsolete). Maxilliped 3 ischium and merus free. Antennal basal article broad; flagellum arising within orbit. Cheliped of male carpus, propodus spinulose. Gonopod 1 with 1 or 2 recurved subdistal lobes and long setae. Maximum cl. 14 mm. Subtidal–slope (10–1216 m). Temperate Northern E Atlantic, including Mediterranean, Central Indo-Pacific, Temperate ­Australasia, Temperate Southern Africa. 6 species (Griffin and Tranter 1986a: features listed, key to species; Richer de Forges et al. 2021: diagnosis, discussion of species).

Holthuija Ng & Richer de Forges, 2015 Diagnosis. Carapace suboval, tuberculate, gastric and branchial regions delimited by grooves; branchial margins with 3 spines. Rostral spines long, slender. Supraorbital eave prominent, anterior part rounded; suborbital margin separated from margin of postorbital spine by deep fissure; preorbital spine absent. Eyestalk long, more than two-thirds exposed in dorsal view. Antennal basal article longer than wide; flagellum arising within orbit. Epistome wider than long. Male thoracic sternites 1–4 elongate, lateral junction between sternites 2 and 3 shallow; male sternoabdominal cavity margin with rim on thoracic somite 4. Cheliped carpus, propodus smooth. Pereopods 2–5 meri upper margins unarmed. Gonopod 1 stout, straight or slightly bent, distal part simple, with spinules and short setae. Maximum cl. 50 mm (Pl. 53d). Subtidal–slope (11–300 m). Temperate Northern W Pacific, Central Indo-Pacific. 6 species (Ng and Richer de Forges 2015: diagnosis, key to species).

629

Jacquinotia Rathbun, 1915 Diagnosis. Carapace pyriform, about as wide as long, granular; branchial margins with tubercles. Rostral spines 0.25–0.4 postrostral carapace length. Orbit closed ventrally; antennal basal article in broad contact with postorbital lobe; supraorbital eave with rounded antorbital angle, intercalated spine, postorbital spine almost fused. Antennal flagellum excluded from orbit by antennal basal article process. Maximum cl. 195 mm. Intertidal–slope (0–550 m). Temperate Australasia (S New ­Zealand only). 1 species (Griffin 1966).

Kasagia Richer de Forges & Ng, 2007 Diagnosis. Carapace suboval, with small rounded or flattened granules; branchial margins with simple or flat mushroom-shaped tubercles; hepatic margin with prominent triangular tooth. Rostral spines long, slender. Supraorbital eave overlapping postorbital spine; intercalated spine absent. Antennal flagellum arising within orbit. Maxilliped 3 ischium and merus free. Cheliped of male ~3 times carapace length. Maximum cl. 13.4 mm. Shelf, slope (148–300 m). Western and Central Indo-Pacific. 3 species (Richer de Forges and Ng 2007; Richer de Forges et  al. 2021: diagnosis, figures and descriptions of species).

Kimbla Griffin & Tranter, 1986 Diagnosis. Carapace suboval, with tubercles and few small spines; branchial margins with tubercles. Rostral spines 0.25–0.4 postrostral carapace length. Supraorbital eave with prominent antorbital spine; preorbital angle with sharp spine; postorbital lobe with tooth on posterior margin. Antennal flagellum excluded from orbit by antennal basal article process. Maximum cl. 19 mm. Shelf, slope (85–244 m). Central Indo-Pacific. 2 species (Griffin and Tranter 1986a: diagnosis; Richer de Forges 1993: second species).

Leptomithrax Miers, 1876 Diagnosis. Carapace densely spinose or tuberculate; branchial margins with spines. Rostral spines separate from base, divergent. Supraorbital eave with prominent antorbital spine; eave, intercalated spine, postorbital spine separated by narrow slits; preorbital angle with unarmed, smooth, rounded. Antennal flagellum excluded from orbit by antennal basal article process. Maximum cl. 165 mm (Pls 1f, 53e, 54a). Intertidal–slope (0–600 m). Temperate Northern Pacific, Central Indo-Pacific, Temperate Australasia. 15 species (Griffin 1963a: rediagnosis; Griffin 1963b: redescription; Griffin and Tranter 1986a: key to 13 species; Richer de Forges and Ng 2014). The genus has a disjunct distribution, inhabiting temperate and cold waters in north-western Asia and Australasia. Leptomithrax gaimardii (H. Milne Edwards, 1934) is one of few majids known to swarm in vast numbers in SE Australia (Pl. 1f; Poore 2004)

Maiopsis Faxon, 1893 Diagnosis. Carapace suborbicular, tuberculate; hepatic margin with 3 spines; branchial margin with 6 spines. Rostral spines with

630

Marine Decapod Crustacea

lateral spine. Supraorbital eave with spine behind preorbital spine. Eyestalk long, more than two-thirds exposed in dorsal view. Maximum cl. 112 mm. Slope (333 m). Tropical Eastern Pacific. 1 species (Rathbun 1925).

Maja Lamarck, 1801 Diagnosis. Carapace pyriform (rounder in adults), covered with numerous heterogeneous granules; branchial margins spinose. Rostral spines short, stout. Suborbital margin separated from margin of postorbital spine by deep fissure; preorbital spine absent. Eyestalk long, more than two-thirds exposed in dorsal view. Antennal basal article as wide as long; flagellum arising within orbit. Epistome wider than long. Cheliped carpus, propodus smooth. Pereopods 2–5 meri upper margins unarmed. Gonopod 1 long, with distal pad-like folds. Maximum cl. 150 mm. Intertidal–shelf (0–184 m). Temperate Northern and Tropical E Atlantic, including Mediterranean, Temperate Southern Africa. 4 species (Ng and Richer de Forges 2015: diagnosis, key to species). Some species of Maja swarm in huge numbers (González-­ Gurriarán et al. 1998; Sampedro and González-Gurriarán 2004).

2–5 propodi flexor margins distally expanded, prehensile. Maximum cl. 70 mm (Pl. 53f). Intertidal–slope (0–503 m). Tropical Atlantic (Brazil), Temperate Australasia. 5 species (Griffin and Tranter 1974: key to species; Poore 2004: figures; Tavares and Poore 2014: Atlantic species).

Neomaja Ng & Richer de Forges, 2015 Diagnosis. Carapace tuberculate; lateral margins spinose. Rostral spines long, slender. Suborbital margin separated from antennal basal antennal and margin of postorbital tooth by 2 gaps; preorbital spine absent. Eyestalk long, more than two-thirds exposed in dorsal view. Antennal basal article as wide as long; flagellum arising within orbit. Epistome as long as wide. Cheliped carpus, propodus smooth. Pereopods 2–5 meri upper margins with subdistal spine. Gonopod 1 long, slender, gently curved, distal part subspatuliform, without subdistal process. Maximum cl. 90 mm. Subtidal–slope (11–264 m). Temperate Northern and Tropical E Atlantic. 1 species (Ng and Richer de Forges 2015).

Notomithrax Griffin, 1963

Diagnosis. Carapace tuberculate; hepatic margin with 2 or 3 spines; branchial margin with 5 simple spines. Rostral spines with 1–3 small accessory spines on lateral margin. Supraorbital eave spinous; intercalated spine well developed. Antennal flagellum arising within orbit. Cheliped of male carpus, propodus spinulose. Maximum cl. 10 mm. Shelf (50–219 m). Temperate Northern W Pacific, Western Indo-Pacific (Madagascar, Mozambique Channel). 3 species (Ng et al. 2021: rediagnosis, figures of 3 species).

Diagnosis. Carapace tuberculate; hepatic margin with 1–3 spines; branchial margin with 2–5 spines. Rostral spines separate from base, divergent. Supraorbital eave with prominent antorbital spine; eave, intercalated spine, postorbital spine widely separated; preorbital angle smooth, rounded; postorbital lobe spinelike, not cupped; cornea of retracted eye not protected on dorsal or ventral surfaces. Antennal basal article broad, with 2 distal spines; peduncular articles 4, 5 stout, article 4 about as wide as long; flagellum excluded from orbit by antennal basal article process. Maximum cl. 52 mm (Pl. 53g). Intertidal–shelf (0–92 m). Temperate Australasia. 4 species (Griffin 1963a: rediagnosis; Griffin and Tranter 1986a: key to species).

Microhalimus Haswell, 1880

Ovimaja Ng & Richer de Forges, 2015

Diagnosis. Carapace without mesogastric tubercle, surface tomentose and with groups of strong curled setae; hepatic margin with 2 broad teeth. Rostral spines divergent, at right angles. Supraorbital eave with prominent antorbital spine; preorbital angle smooth, rounded; intercalated spine absent; postorbital spine, not cupped; cornea of retracted eye not protected on dorsal or ventral surfaces. Antennal flagellum excluded from orbit by antennal basal article process. Maximum cl. 15 mm. Intertidal–shelf (0–71 m). Temperate Australia. 1 species (Griffin and Tranter 1986a: comment; Poore 2004: diagnosis, figure).

Diagnosis. Carapace ovoid, longer than wide, tuberculate, gastric and branchial regions delimited by grooves; lateral margins spinose. Rostral spines short, stout. Suborbital margin separated from margin of postorbital spine by deep fissure; preorbital spine absent. Eyestalk long, more than two-thirds exposed in dorsal view. Antennal basal article longer than wide, with several large granules and tubercles; flagellum arising within orbit. Epistome longer than wide. Male thoracic sternites 1–4 not elongated, lateral junction between sternites 2 and 3 deep, forming waist-like structure; male sterno-abdominal cavity margin almost flat, without rim. Cheliped carpus, propodus smooth. Pereopods 2–5 meri upper margins unarmed. Gonopod 1 strongly bent, distal part dilated with 2 folds. Maximum cl. 59 mm (Pl. 53h). Shelf (20–84 m). Central Indo-Pacific. 1 species (Ng and Richer de Forges 2015).

Majella Ortmann, 1893

Naxia Latreille, 1825 Diagnosis. Carapace with at least 1 mesogastric tubercle, surface bare or with few short setae between groups of strong curled setae; hepatic margin with 2 spines. Rostral spines separate from base, divergent. Intercalated spine present (sometimes almost fused to eave); postorbital spine not cupped; cornea of retracted eye not protected on dorsal or ventral surfaces. Antennal flagellum excluded from orbit by antennal basal article process. Pereopods

Paraentomonyx Sakai, 1983 Diagnosis. Carapace with few spines in midline; branchial margin with 4 spines; posterior margin with 2 spines. Rostral spines

14 – Brachyura – crabs

separate from base, divergent. Orbit closed ventrally; antennal basal article in broad contact with postorbital lobe; supraorbital eave strongly expanded to form tubular orbit. Antennal flagellum excluded from orbit by antennal basal article process. Maximum cl. 25 mm. Shelf (20–179 m). Temperate Northern W Pacific. 1 species (Sakai 1983: figure).

Paramaja Kubo, 1936 Diagnosis. Carapace pyriform (rounder in adults), covered with numerous heterogeneous granules, branchial regions of adults swollen laterally and dorsally; cardiac region swollen; regions poorly demarcated; branchial margins tuberculate but without spines in adults. Rostral spines short, stout. Suborbital margin confluent with margin of postorbital tooth, not demarcated by fissure or cleft; preorbital spine absent. Eyestalk long, more than two-thirds exposed in dorsal view. Antennal basal article longer than wide; flagellum arising within orbit. Epistome wider than long. Cheliped carpus, propodus smooth. Pereopods 2–5 meri upper margins unarmed. Gonopod 1 straight, distal part conical, with scattered setae on distal surface. Maximum cl. 88 mm (Pl. 53i, j). Shelf, slope (100–580 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 3 species (Ng and Richer de Forges 2015: rediagnosis, key to species).

Paramaya De Haan, 1837 Diagnosis. Carapace suboval, tuberculate; lateral margins spinose. Rostral spines long, slender. Suborbital margin separated from margin of postorbital tooth by deep fissure; preorbital spine absent. Eyestalk long, more than two-thirds exposed in dorsal view. Antennal basal article longer than wide; flagellum arising within orbit. Epistome longer than wide. Cheliped carpus, propodus smooth. Pereopods 2–5 meri upper margins with subdistal spine. Gonopod 1 long, slender, gently curved, distal part with rounded tip, with dorsal projection. Maximum cl. 72 mm. Subtidal–slope (15–300 m). Temperate Northern W Pacific, Central Indo-Pacific. 4 species (Ng et al. 2018; Ng and Richer de Forges 2015: rediagnosis, key to 3 species).

631

Pippacirama Griffin & Tranter, 1986 Diagnosis. Carapace with tubercles and few small spines; branchial margin with 4 spines. Rostral spines, deflexed, short, truncate. Orbit open ventrally; antennal basal article widely separated from postorbital lobe; supraorbital eave weakly expanded, barely covering base of eyestalk; intercalated spine absent; postorbital lobe lamellate, with spine on anterior margin. Eyestalk long, with lobe on anterior margin. Antennal peduncular article 4 flat, broad; article 5 laterally directed. Maximum cl. 21 mm. Intertidal, subtidal (0–7 m). Temperate Australia. 1 species (Griffin and Tranter 1986a: diagnosis; Poore 2004: figure).

Planaja Ng & Richer de Forges, 2015 Diagnosis. Carapace suboval, tuberculate, gastric and branchial regions delimited by grooves; branchial margins with 6 spines. Rostral spines long, slender. Suborbital margin separated from margin of postorbital tooth by deep fissure; preorbital spine absent. Eyestalk long, more than two-thirds exposed in dorsal view. Antennal basal article longer than wide; flagellum arising within orbit. Epistome wider than long. Male thoracic sternites 1–4 elongate, lateral junction between sternites 2 and 3 shallow; male sterno-abdominal cavity margin with rim on thoracic somite 4. Cheliped carpus, propodus smooth. Pereopods 2–5 meri upper margins unarmed. Gonopod 1 slender, curved, distal part sharply tapering, margins chitinised, with numerous short spinules. Maximum cl. 47 mm. Shelf, slope (100–349 m). Central Indo-Pacific. 1 species (Ng and Richer de Forges 2015).

Planotergum Balss, 1935 Diagnosis. Carapace suboval, widest at midpoint, flat, smooth; lateral margins lamellate. Rostral spines projecting beyond orbits, lamellar, wide. Orbit not protecting short eyestalk. Antennal basal article broad; peduncular articles 4, 5 lamellar, both visible in dorsal view. Cheliped minute, not visible in dorsal view. Pereopods 2–5 short, flat, carinate, distal articles in same plane as carapace. Maximum cl. 13.4 mm (Pl. 53l). Subtidal, shelf (3–25 m). Western and Central Indo-Pacific. 2 species (Marin et al. 2019; Serène 1965).

Prismatopus Ward, 1933 Paramithrax H. Milne Edwards, 1834 sea toad Diagnosis. Carapace tuberculate, gastric, cardiac and intestinal regions smooth; branchial margin with 2 spines. Rostral spines partly fused, short, triangular. Preorbital angle smooth, rounded; postorbital spine not cupped; cornea of retracted eye not protected on dorsal or ventral surfaces. Antennal basal article apically notched, distolateral tooth short, rounded crenulate; flagellum excluded from orbit by antennal basal article process. Maximum cl. 30 mm. Subtidal, shelf (1–30 m). Temperate Australia. 1 species (Griffin 1963a: rediagnosis; Griffin 1963b: redescription of only species).

Diagnosis. Carapace with spines; branchial margin with 1–3 sharp or knobbed spines (last subdorsal), sometimes with anterior lamellate lobe. Rostral longer than 0.3 postrostral carapace length. Supraorbital eave with prominent antorbital spine; preorbital angle with sharp spine. Antennal flagellum excluded from orbit by antennal basal article process. Cheliped merus and carpus usually with longitudinal ridge; male chela short, stout (palm ~1.5 times as long as high). Maximum cl. 54 mm (Pl. 53k). Subtidal–slope (10–460 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia. 13 species (Griffin and Tranter 1986a: rediagnosis, key to species as junior synonym, Thacanophrys Griffin & Tranter, 1986; Takeda et al. 2022: species comparisons).

632

Marine Decapod Crustacea

Pseudomicippe Heller, 1861 Diagnosis. Carapace suboval, tuberculate; branchial margins with tubercles or short spines. Rostral spine up to third postrostral carapace length, partly fused, sometimes deflexed. Intercalated spine absent; postorbital lobe spine separated by a narrow hiatus. Eyestalk long, more than two-thirds exposed in dorsal view. Maximum cl. 29 mm. Intertidal–shelf (0–72 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australia. 14 species (Griffin and Tranter 1986a: rediagnosis, key to 11 species, figures of 9 species; Kazmi and Tirmizi 1999: species description).

Rathbunaja Ng & Richer de Forges, 2015 Diagnosis. Carapace densely spinose or tuberculate, gastric and branchial regions delimited by grooves; lateral margins spinose. Rostral spines short, stout, or spines long, slender. Suborbital margin confluent with margin of postorbital tooth, not demarcated by fissure or cleft; preorbital spine absent. Eyestalk long, more than two-thirds exposed in dorsal view. Antennal basal article longer than wide; flagellum just outside orbit, near rim. Epistome wider than long. Cheliped carpus, propodus smooth. Pereopods 2–5 meri upper margins with subdistal spine. Gonopod 1 long, slender, gently curved, distal part with rounded tip, with dorsal projection. Maximum cl. 54 mm. Shelf, slope (169–410 m). Temperate Northern W Pacific, Central Indo-Pacific. 4 species (Ng and Richer de Forges 2015: diagnosis, key to species).

Sakaija Ng & Richer de Forges, 2015 Diagnosis. Carapace pyriform (rounder in adults), covered with small uniform granules, gastric and branchial regions delimited by grooves; lateral margins spinous. Rostral spines short, stout. Suborbital margin confluent with margin of postorbital tooth, not demarcated by fissure or cleft; preorbital spine absent. Eyestalk long, more than two-thirds exposed in dorsal view. Antennal basal article longer than wide; flagellum arising within orbit. Epistome wider than long. Cheliped carpus, propodus smooth. Pereopods 2–5 meri upper margins unarmed. Gonopod 1 sinuous, with numerous long setae on distal surface. Maximum cl. 29 mm (Pl. 54b). Shelf, slope (30–567 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 7 species (Ng and Richer de Forges 2015: diagnosis, key to 6 species).

Schizophroida Sakai, 1933 Diagnosis. Carapace pyriform, branchial width ~1.5 times hepatic width, with few blunt tubercles. Rostral spines separate from base, divergent or subparallel; branchial margins with spines. Supraorbital eave with prominent antorbital spine; preorbital angle smooth, rounded; intercalated spine larger than antorbital spine, not excluded from margin of orbit. Antennal basal article narrow, with 2 slender distal spines; flagellum excluded from orbit by antennal basal article process. Maximum cl. 37 mm.

Intertidal–shelf (0–119 m). Temperate Northern W Pacific, Central and Eastern Indo-Pacific, Temperate Australasia. 5 species (Griffin and Tranter 1986a: comment; Ng and Ahyong 2018: key to species; Ng and Boyko 2017).

Schizophrys White, 1848 Diagnosis. Carapace tuberculate. Rostral spines with 1 or 2 lateral accessory spines. Supraorbital eave with prominent antorbital spine; preorbital angle smooth, rounded; postorbital lobe with basal accessory tooth. Lateral margins with spines. Antennal flagellum excluded from orbit by antennal basal article process. Maximum cl. 59 mm (Pl. 54c). Intertidal–shelf (0–126 m). Temperate Northern W Pacific, Indo-West Pacific, Temperate Australia. 6 species (Griffin and Tranter 1986a: comment, key and descriptions of 4 species; Tirmizi and Kazmi 1995).

Seiitaoides Griffin & Tranter, 1986 Diagnosis. Carapace with tubercles fused into 10–12 large tuberculate or spinulate plates. Rostral spines separated by U-shaped hiatus; branchial margins with blunt or mushroom-shaped tubercles; hepatic margin with bifid spine. Supraorbital eave denticulate; intercalated spine short, usually close to eave and postorbital lobe. Antennal flagellum arising within orbit. Maxilliped 3 ischium and merus free. Cheliped of male spinous, longer than ambulatory legs; chela palm spinous, fingers short, bent. Maximum cl. 12 mm. Shelf, slope (45–245 m). Temperate Northern W Pacific, Central Indo-Pacific. 2 species (Griffin and Tranter 1986a: diagnosis, key to species; Richer de Forges et al. 2021: diagnosis, descriptions and figures of species).

Temnonotus A. Milne-Edwards, 1875 Diagnosis. Carapace cardiac region surrounded, except anteriorly, by deep trench. Rostral spines long, slender. Antennal flagellum arising within orbit. Maximum cl. 23 mm. Shelf, slope (183–478). Tropical Atlantic (Caribbean). 1 species (Rathbun 1925).

Teratomaia Griffin & Tranter, 1986 Diagnosis. Carapace with dorsal spines; branchial margin with 4–6 smooth sharp spines (last subdorsal). Rostral spines ~0.2 postrostral carapace length. Supraorbital eave with prominent preorbital and antorbital spine; preorbital angle with sharp spine; postorbital lobe with tooth on posterior margin. Antennal flagellum excluded from orbit by antennal basal article process. Cheliped merus and carpus with spinous tubercles; male chela long, slender (palm ~4 times as long as high). Maximum cl. 102 mm. Slope (224–648, possibly 7160 m). Temperate Australasia (New Zealand, SE Australia). 1 species (Griffin 1966: species description; Griffin and Tranter 1986a: diagnosis). Teratomaia is usually recorded to slope depths of ~700 m, but the extraordinary record from the Kermadec Deep at 7160 m (Griffin and Tranter 1986b) requires corroboration.

14 – Brachyura – crabs

Tumulosternum McCulloch, 1913 Diagnosis. Carapace with few blunt tubercles; hepatic margin with strong spine or tubercle. Rostral spines short, stout. Preorbital angle smooth, rounded; intercalated spine much smaller than antorbital spine or obsolete, excluded from margin of orbit. Antennal basal article broad, with 2 short, blunt, distal spines; flagellum excluded from orbit by antennal basal article process. Maximum cl. 26 mm. Intertidal, subtidal (0–5 m). Temperate SE Australia. 3 species (Griffin and Tranter 1986a: rediagnosis, key to species). References Baker WH (1905) Notes on South Australian decapod Crustacea. Part II. Transactions of the Royal Society of South Australia 29, 116–130, pls 21–24. Balss H (1922) Decapoden von Juan Fernandez. In The Natural History of Juan Fernandez and Easter Island. (Ed. Skottsberg C) pp. 329–340. Almqvist & Wiksells, Uppsala. Forest J, Guinot D (1961) Crustacés décapodes Brachyoures de Tahiti et des Tuamotu. In Expédition française sur les récifs coralliens de la Nouvelle-Calédonie. Volume préliminaire. pp. 1–195, pls 1–18, Paris. Garth JS (1958) Brachyura of the Pacific coast of America. Oxyrhyncha. Allan Hancock Pacific Expeditions 21, 1–854. González-Gurriarán E, Fernández L, Freire J, Muiño R (1998) Mating and role of seminal receptacles in the reproductive biology of the spider crab Maja squinado (Decapoda, Majidae). Journal of Experimental Marine Biology and Ecology 220, 269–285. doi:10.1016/S0022-0981(97)00109-3 Griffin DJG (1963a) Notomithrax gen. nov. and the status of the genus Paramithrax H. Milne Edwards (Crustacea, Brachyura, Majidae). Transactions of the Royal Society of New Zealand 3, 229–237. Griffin DJG (1963b) Redescriptions of the Australian majid spider crabs Leptomithrax gaimardii (H. Milne Edwards) and Paramithrax barbicornis (Latreille). Records of the Australian Museum 26, 131–43, pls 6–7. Griffin DJG (1966) The marine fauna of New Zealand: spider crabs, family Majidae (Crustacea, Brachyura). New Zealand Oceanographic Institute Memoir 172, 1–111. Griffin DJG, Tranter HA (1974) Spider crabs of the family Majidae (Crustacea: Decapoda: Brachyura) from the Red Sea. Israel Journal of Zoology 23, 162–198. Griffin DJG, Tranter HA (1986a) The Decapoda Brachyura of the Siboga Expedition. Part VIII Majidae. Siboga-Expéditie 39 c4, 1–335. Griffin DJG, Tranter HA (1986b) Some majid spider crabs from the deep Indo-West Pacific. Records of the Australian Museum 38, 351–370. doi :10.3853/j.0067-1975.38.1986.186 Guinot D, Wicksten MK (2015) Chapter 71–11. Camouflage: carrying behaviour, decoration behaviour, and other modalities of concealment in Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 583–638. Brill, Leiden. Hultgren KM, Stachowicz JJ (2011) Camouflage in decorator crabs. In Animal camouflage. (Eds Stevens MA, Merilaita S) pp. 214–238. Cambridge University Press, Cambridge. Kazmi QB, Tirmizi NM (1999) Two new species and a new record of spider crabs with a note on Stilbognathus curvicornis (Herbst, 1803) from Pakistan waters of the Northern Arabian Sea. Crustaceana 72, 369–382. doi:10.1163/156854099503438 Marin IN, Spiridonov VA, Ng PKL (2019) A new species of the aberrant majid crab genus Planotergum Balss, 1935 (Crustacea: Brachyura,

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Majidae, Planoterginae) from the southern Red Sea. Zoological Studies 58(26), 1–11. Ng PKL, Ahyong S (2018) New species of reef spider crabs of the genus Schizophroida Sakai, 1933 (Crustacea: Brachyura: Majidae) from the Western Pacific. Records of the Australian Museum 70, 377–390. doi:10 .3853/j.2201-4349.70.2018.1712 Ng PKL, Boyko CB (2017) New species and records of crabs of the families Dromiidae, Dynomenidae, Homolidae, Aethridae, Parthenopidae, Majidae, and Epialtidae (Crustacea: Decapoda: Brachyura) from Easter Island, with a review of the majid genus Schizophroida Sakai, 1933. Pacific Science 71, 197–227. doi:10.2984/71.2.8 Ng PKL, Richer de Forges B (2015) Revision of the spider crab genus Maja Lamarck, 1801 (Crustacea: Brachyura: Majoidea: Majidae), with descriptions of seven new genera and 17 new species from the Atlantic and Indo-West Pacific. Raffles Bulletin of Zoology 63, 110–225. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Ng PKL, Prema M, Ravichandran S (2018) A new species of deep-water spider crab of the genus Paramaya De Haan, 1837 from the Bay of Bengal, India (Crustacea, Brachyura, Majidae). ZooKeys 769, 77–88. doi:10.3897/zookeys.769.26152 Ng PKL, Lee BY, Richer de Forges B (2021) Revision of Majella Ortmann, 1893 (Crustacea: Brachyura: Majidae), with description of two new species from the Indian Ocean. Zoological Studies 60, 15. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Rathbun MJ (1925) The spider crabs of America. Bulletin of the United States National Museum 129, 1–613. Richer de Forges B (1993) Deep sea crabs of the Tasman seamounts (Crustacea: Decapoda: Brachyura). Records of the Australian Museum 45, 11–24. doi:10.3853/j.0067-1975.45.1993.126 Richer de Forges B, Lee BY, Ng PKL (2021) The taxonomy of spider crabs of the genera Eurynome, Choniognathus, Seiitaiodes and Kasagia (Crustacea: Brachyura: Majidae) from southwest Indian Ocean. Zootaxa 5048, 301–333. Richer de Forges B, Ng PKL (2007) On a new genus and new species of deep-water spider crab from the Philippines (Crustacea: Decapoda: Brachyura: Majidae). Zootaxa 1644, 59–68. doi:10.11646/ zootaxa.1644.1.3 Richer de Forges B, Ng PKL (2014) The identity of Leptomithrax sinensis Rathbun, 1916, and the description of L. eldredgei, sp. nov. from Hong Kong (Crustacea: Decapoda: Brachyura: Majidae). Bishop Museum Bulletin in Zoology 9, 123–128. Sakai T (1983) Description of new genera and species of Japanese crabs together with systematically and biogeographically interesting species. (I). Researches on Crustacea 12, 1–44, pls 1–8. Sampedro M-P, González-Gurriarán E (2004) Aggregating behaviour of the spider crab Maja squinado in shallow waters. Journal of Crustacean Biology 24, 168–177. doi:10.1651/C-2404 Serène R (1965) Note sur un Oxyrhynche (Brachyura, Decapoda, Crustacea) aberrant du Sud-Est Asiatique. Les recherches de zoologie marine au Viet-Nam. Bulletin de la Societé Zoologique de France 90, 457–468, pls 1, 2. Takeda M, Ahyong ST, Ohtsuchi N, Komatsu H (2022) Crabs (Crustacea, Decapoda) from the Seas of East and Southeast Asia Collected by the RV Hakuhō Maru (KH–72–1 Cruise) 3. Sahul Shelf. Bulletin of the National Museum of Nature and Science. Series A, Zoology 48(2), (in press). Tavares M, Poore GCB (2014) A new species of Naxia Latreille, 1825 (Brachyura: Majidae) from deep water off Brazil. Zootaxa 3861, 86–90. doi:10.11646/zootaxa.3861.1.5

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Tirmizi NM, Kazmi QB (1995) A new species of Schizophrys from Pakistan with a note on Schizophrys aspera (Crustacea: Decapoda: Majidae). In The Arabian Sea: Living Marine Resources and the Environment. (Eds Thompson M-F, Tirmizi NM) pp. 136–141. Vanguard Books, Lahore.

Mithracidae MacLeay, 1838 Figures 14.89, 14.90, Plate 54d–k Mithracid spider crabs are confined to 60 species in 14 genera from the subtropical and tropical coasts of the Americas. The only exception is one species from the eastern Atlantic (Windsor and Felder 2014). Windsor and Felder (2014) excluded all previously included genera from the Indo-West Pacific (e.g. by Griffin and Tranter 1986), plus three from the Americas, on the basis of a thorough molecular and morphological analysis. They showed that the family is monophyletic, their work supplanting that of earlier phylogenetic studies (Baeza et  al. 2010; Hultgren and Stachowicz 2008). Windsor and Felder (2014) provided a key and diagnosed all genera. Their nomenclature has since been corrected (Windsor and Felder 2017) and another genus added (Klompmaker et al. 2015). Mithracids are found on intertidal and subtidal rocky and coral reefs, rarely to deep shelf depths. Some are known

to associate with sea anemones (Baeza et al. 2010). Many are very colourful (Windsor and Felder 2014). Diagnosis. Orbits formed dorsally by supraorbital eave, adjacent spines and postorbital spine or lobe, closed ventrally by expansion of broad basal antennal article forming floor of orbit. Thoracic pleurites 5–8 not visible dorsally, covered by posterolateral margin of carapace. Eyestalk hidden by supraorbital eave and spines. Male pleotelson formed by fused pleonite 6 and telson in males, fused pleonites 5–6 and telson in females. Pleonite 1 free from to carapace. Maxilliped 3 merus as wide as or wider than ischium. Gonopod 1 usually slender, straight or weakly curved, aperture terminal, apex simple or curved or expanded into 2 or 3 processes. Implicit genera attributes. Antennal article 4 subcylindrical or weakly depressed, if dorsally flattened longer than broad. Maxilliped 3 exopod mesial margin without flange overlapping merus. Cheliped of mature male less than twice as long as carapace. Cheliped propodus upper margin smooth; inner face smooth. Pereopods 2–4 meri without lamellate arcuate expansions of lower margins. Pereopod 2 longer than carapace length by less than dactylus plus half propodus.

Key to genera of Mithracidae 1. – 2. – 3. – 4. – 5. – 6. –

Antennal peduncular article 4 flat, wide, with lateral flange, sometimes broader than long (Figs 14.89i, 14.90g)�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Antennal peduncular article 4 subcylindrical or weakly depressed (Fig. 14.90f, h)��������������������������������������������������������������3 Antennal peduncular article 4 subquadrate, almost as wide as basal article (Fig. 14.90g). Carapace anterolateral margin with 0–3 blunt, tubercles. Pereopods 2–4 meri flat, with well defined upper and lower marginal carinae (Fig. 14.89b)����������������������������������������������������������������������������������������������������������������������������� Hemus Antennal peduncular article 4 rectangular, narrower than basal article. Carapace anterolateral margin with 4–6 acute or rounded hepatic and branchial teeth, continuous with posterolateral margin. Pereopods 2–4 meri cylindrical, without upper and lower marginal carinae (Fig. 14.89i)������������������������������������� Pitho Pereopods 2–4 meri flat, with lamellate expansion on lower margins (Fig. 14.90x)������������������������������������������������������Thoe Pereopods 2–4 meri cylindrical, sometimes spinose, usually on upper margins������������������������������������������������������������������4 Carapace subpyriform to subtriangular, strongly narrowing anteriorly to urn-shaped; anterolateral margin without or with few tubercles, posterolateral angle with 1 or 2 teeth (Fig. 14.89d, h). Rostrum comprising 2 elongate spines�����������������������������������������������������������������������������������������������������������������������������������������������������5 Carapace broadly subtriangular; anterolateral margin well ornamented, posterolateral tooth present or absent. Rostrum comprising 2 short teeth, lobes or spines��������������������������������������������������������������������������������������������������6 Maxilliped 3 merus distomesial margin truncate, palp set in triangular notch with mesiodistal tooth (Fig. 14.90u). Pereopods 2–4 carpi and propodi unarmed or with distal tooth on upper margin (Fig. 14.89h)����������������������������������������������������������������������������������������������������������������������������������������������������������� Omalacantha Maxilliped 3 merus convex, obliquely excavated for palp, with small mesiodistal tooth (Fig. 14.90r). Pereopods 2–4 carpi and propodi each bearing upper spine or angular tooth (Fig. 14.89d)������������������������Microphrys Carapace anterolateral margin with upper and lower parallel rows of small teeth. Rostrum of 2 broad lobes (Fig. 14.90c). Cheliped of mature male twice as long as carapace����������������������������������������������������������������� Petramithrax Carapace anterolateral margin single or with ill-defined submarginal row of angular lobes, tubercles or teeth. Rostrum of 2 teeth or spines. Cheliped of mature male usually less than twice as long as carapace�����������������7

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Fig. 14.89.  Mithracidae. a, Amphithrax pilosus (Rathbun, 1892); b, Hemus magalae Windsor & Felder, 2011; c, Maguimithrax spinosissimus (Lamarck, 1818); d, Microphrys branchialis Rathbun, 1898; e, Mithraculus coryphe (Herbst, 1801); f, Nemausa acuticornis (Stimpson, 1871); g, Nonala holderi (Stimpson, 1871); h, Omalacantha garthi (Lemos de Castro, 1953); i, Pitho aculeata (Gibbes, 1850).

7. – 8. –

Orbital margins without tubercles. Carapace anterolateral margin lobate or weakly dentate (Fig. 14.90d). Pereopods 2–4 meri upper margins with spines or lobes (Fig. 14.90w)������������������������������������������������������������ Teleophrys Orbital margins with teeth, spines or tubercles. Carapace anterolateral margin with 3 or more well defined lobiform or acute teeth. Pereopods 2–4 meri upper margins with tubercles or spines, not lobes���������������������������������8 Maxilliped 3 exopod mesial margin with rounded lamelliform flange overlapping merus (Fig. 14.90n). Carapace anterolateral margin with lamelliform lobes, posterolateral margin with broad lamellate expansion (Fig. 14.90a)���������������������������������������������������������������������������������������������������������������������������������������������������������� Ala Maxilliped 3 exopod mesial margin without flange overlapping merus (Fig. 14.90–o–u). Carapace anterolateral margin with teeth or lobes, posterolateral margin not expanded���������������������������������������������������������������9

636

Marine Decapod Crustacea

Fig. 14.90.  Mithracidae. Carapace, representative or only species: a, Ala; b, Mithrax; c, Petramithrax; d, Teleophrys; e, Thoe. f, left anterior with rostrum, epistome, antenna, eyestalk, subhepatic regions, Nemausa. Left antennal basal article, peduncular articles, 2, 3, flagellum, eyestalk, ventral view: g, Hemus; h, Thoe. Right antennal basal article: i, Amphithrax; j, Maguimithrax; k, Mithrax; l, Nemausa; m, Nonala. Maxilliped 3: n, Ala; o, Amphithrax; p, Hemus; q, Maguimithrax; r, Microphrys; s, Mithrax; t, Nonala; u, Omalacantha. v, male chela, Mithrax. w, pereopod 2, Teleophrys. x, pereopodal merus, Thoe. Features labelled on b, f: aba, antennal basal article; ao, antorbital spine; n, nephropore (antennal gland); po, preorbital spine; ps, postorbital spine; r, rostral spine.

9.

Carapace branchial regions with well defined, broad, uninterrupted, smooth, oblique sulci extending from anterolateral margins to weakly converge posteromesially, separated by raised, nearly continuous, ridges (Fig. 14.89e)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Mithraculus – Carapace branchial regions lacking posteromesially convergent oblique sulci or with ill-defined sulci roughened or interrupted by tubercles, obscured by partial anastomosing of separating ridges, or marked by broken ridges bearing incisions or depressions as deep and wide as sulci themselves������������������������������10 10. Maxilliped 3 merus mesial margin with tongue-like projection, palp set in deep square excavation, distolateral corner produced (Fig. 14.90q, t)�������������������������������������������������������������������������������������������������������������������������11 – Maxilliped 3 merus mesial margin truncate, palp set on oblique angle, distolateral corner rounded–square (Fig. 14.90o, s)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������13 11. Carapace with spine at widest part of carapace smaller than preceding spines (Fig. 14.89c). Antenna basal article with largest spine obliquely truncate (Fig. 14.90j)����������������������������������������������������������������������������� Maguimithrax – Carapace with spine at widest part of carapace larger than preceding spines (Fig. 14.89f). Antenna basal article with largest spine acute or rounded (Fig. 14.90l, m)����������������������������������������������������������������������������������������������� 12 12. Carapace anterolateral margin with 1 prominent hepatic hooked spine, 2–8 uneven tubercles, posterolateral tooth curved (Fig. 14.89g). Cheliped propodus smooth in adults���������������������������������������������������Nonala

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– 13. –

637

Carapace anterolateral margin with 3–5 acute spines, usually with accessory spinules, 1 tooth at posterolateral angle (Fig. 14.89f). Cheliped propodus upper margin with row of spines in adults���������������� Nemausa Carapace transversely ovoid, 1.2–1.5 times as wide as long; anterolateral margin with 4 lateral spines or teeth, sometimes with accessory spine; posterolateral spine minutely hooked (Fig. 14.90b)���������������������������� Mithrax Carapace subcircular-ovoid, as wide or little longer than wide; anterolateral margin with several, often compound, spines; posterolateral angle with single spine, not markedly larger than others (Fig. 14.89a)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Amphithrax Ala Lockington, 1877

Diagnosis. Carapace subtriangular, wider than long, branchial and gastric regions paved by low elevations densely covered by short setae; anterolateral margin with lamelliform lobes, posterolateral margin with broad lamellate expansion. Rostrum comprising 2 elongate spines. Orbital margin without teeth. Antennal basal article with 1 elongate spine at anterolateral corner, subequal to or exceeding rostrum length. Maxilliped 3 merus mesial margin broadly rounded, with shallow concavity for palp. Maxilliped 3 exopod mesial margin with rounded flange overlapping merus. Cheliped of mature male subequal to or exceeding carapace length. Cheliped propodus upper margin with row of tubercles or spines. Pereopods 2–4 meri margins with row of laminar spines. Maximum cl. 17 mm. Subtidal, shelf (11–60 m). Tropical Eastern Pacific. 1 species (Rathbun 1925: redescription as Anaptychus cornutus Stimpson, 1860; Windsor and Felder 2014: rediagnosis).

Amphithrax Windsor & Felder, 2017 Diagnosis. Carapace subcircular-ovoid, as wide as or little wider than long, dorsally paved with close-set granulated convex tubercles or flat granules, partially concealed by short stiff setae except in large adults; anterolateral margin with several, often compound, spines; posterolateral angle with single spine, not markedly larger than others. Rostrum comprising 2 spines, convergent in immature, short and blunt or truncate in large adults. Antennal basal article with 3 marginal spines or teeth, at least 2 visible in dorsal view: first at base of basal article; second forming strong preocular spine. Maxilliped 3 merus mesial margin straight, palp set in oblique excavation, distolateral angle rounded-square. Cheliped propodus upper margin with row of tubercles or spines, or smooth. Maximum cl. 116 mm. Intertidal–shelf (0–60 m). Tropical Atlantic. 10 species (Wagner 1990: description of some species as Mithrax; Windsor and Felder 2014: diagnosis as Mithrax; Windsor and Felder 2017: rediagnosis).

Hemus A. Milne-Edwards, 1875 Diagnosis. Carapace ovate, longer than wide, with granules and tubercles, elevated along part of midline, transversely elevated at cardiac region, with tufts of setae; anterolateral margin with 0–3 blunt tubercles. Rostrum broad, excavate. Orbital margin without teeth. Antennal basal article unarmed; antenna peduncular article 4 subquadrate, almost as wide as basal article. Maxilliped 3

merus mesial margin convex, not excavated for palp, distolateral margin lobe-like. Pereopods 2–4 meri flat, with well defined upper and lower marginal carinae. Maximum cl. 9 mm. Intertidal–shelf (0–60 m). Tropical W Atlantic, Tropical Eastern Pacific. 4 species (Windsor and Felder 2011: key to species; 2014: rediagnosis).

Maguimithrax Klompmaker, Portell, Klier, Prueter & Tucker, 2015 Diagnosis. Carapace subcircular, about as wide as long, covered with spines, especially medially; anterolateral margin with 6 spines, anteriormost ones with accessory spines at anterior bases, fifth and sixth spines weakest. Rostrum comprising 2 short teeth or lobes. Orbital margin with 4 or 5 spines behind preorbital tooth. Antennal basal article with 3 spines: first, near base of basal article reduced; second largest, truncate reaching as far forward as rostrum; third half as long as second. Maxilliped 3 merus mesial margin tongue-like, palp set in deep square excavation, distolateral corner produced. Cheliped propodus upper margin with row of tubercles or spines; inner face with 2–4 tubercles. Maximum cl. 170 mm. Subtidal, shelf (1–179 m). Temperate Northern W Atlantic, Tropical Atlantic. 1 species (Klompmaker et al. 2015).

Microphrys H. Milne Edwards, 1851 Diagnosis. Carapace pyriform, longer than wide, branchial region strongly inflated, uneven, with row of tubercles parallel to posterior margin; anterolateral margin with 4 or more tubercles or spines, 1 posterolateral spinulose spine. Rostrum comprising 2 elongate spines. Antennal basal article with 1 large anterior spine visible in dorsal view, often with smaller tooth near basal article, with or without anterolateral tooth. Maxilliped 3 merus mesial margin convex, obliquely excavated for palp, with small mesiodistal tooth. Cheliped of mature male less than twice as long as carapace. Pereopods 2–4 carpi and propodi each bearing distinct upper spine or angular tooth. Maximum cl. 34 mm. Intertidal–shelf (0–90 m). Tropical Eastern Pacific. 5 species (Garth 1958: key to species; Windsor and Felder 2014: rediagnosis).

Mithraculus White, 1847 Diagnosis. Carapace subhexagonal, usually wider than long, branchial regions with well defined, broad, uninterrupted, smooth, oblique sulci extending from anterolateral margins to weakly converge posteromesially, separated by raised, nearly

638

Marine Decapod Crustacea

continuous, ridges; anterolateral margin with 3 or 4 simple lobes or spines, sometimes setose. Rostrum comprising 2 blunt teeth. Antennal basal article anterolaterally produced as rounded or elongate tooth, not visible in dorsal view, usually with tubercle near base of basal article. Maxilliped 3 merus mesial margin straight, oblique at base of palp. Cheliped of mature male twice as long as carapace, or less than twice as long as carapace. Pereopods 2–4 carpi and propodi spinose. Maximum cl. 30 mm (Pl. 54d, e). Intertidal–shelf (0–90 m; associated with sponges, corals and sea anemones). Temperate W Northern Atlantic, Tropical W Atlantic, Tropical Eastern Pacific. 8 species (Garth 1958: key to 2 E Pacific species; Wagner 1990: key to 5 W Atlantic species; Windsor and Felder 2014: rediagnosis).

Mithrax Latreille, 1816 Diagnosis. Carapace transversely ovoid, 1.2–1.5 times as wide as long, smooth or tuberculate; anterolateral margin with 4 lateral spines or teeth, sometimes with accessory spine; posterolateral spine minutely hooked. Rostrum comprising 2 blunt teeth or lobes. Orbital margin with tubercles behind preorbital tooth. Antennal basal article with 2 lobes visible in dorsal view: first larger, reaching as far forward as rostrum, second third-half as long as first. Maxilliped 3 merus mesial margin straight, palp set in oblique excavation, distolateral angle rounded-square. Cheliped of mature male subequal to or exceeding carapace length. Maximum cl. 63 mm (Pl. 54f, g). Subtidal, shelf (1–65 m). Temperate Northern W Atlantic, Tropical Atlantic. 3 species (Windsor and Felder 2009: species differentiation; Windsor and Felder 2014: rediagnosis as Damithrax).

Nemausa A. Milne-Edwards, 1875 Diagnosis. Carapace pyriform, longer than wide, with numerous separate erect dorsal spiniform tubercles, slightly concealed by light setation lacking in large adults; anterolateral margin with 3–5 acute spines, usually with accessory spinules, 1 tooth at posterolateral angle. Rostrum comprising 2 spines, weakly convergent in immature, elongate in adults, about twice basal width. Orbital margin with 2 or 3 teeth behind preorbital tooth. Antennal basal article with 3 spines, longest conspicuous in dorsal view, similar but shorter than rostral horn. Maxilliped 3 merus mesial margin tongue-like, palp set in deep square excavation, distolateral corner produced. Cheliped propodus upper margin with row of tubercles or spines in adults. Pereopods 2–4 carpi and propodi spinose. Maximum cl. 93 mm (Pl. 54h). Subtidal–slope (1–1077 m). Tropical W Atlantic, Tropical Eastern Pacific. 4 species (Wagner 1990: description as Mithrax cornutus de Saussure, 1857; Windsor and Felder 2014: rediagnosis).

Nonala Windsor & Felder, 2014 Diagnosis. Carapace pyriform, longer than wide, tuberculate; anterolateral margin with 1 prominent hepatic hooked spine, 2–8 uneven tubercles, posterolateral tooth curved. Rostrum comprising 2 sharp short teeth. Orbital margin with 2 or 3 teeth behind

preorbital tooth. Antennal basal article with 1 elongate spine at anterolateral corner almost reaching end of rostrum, 2 smaller teeth. Maxilliped 3 merus mesial margin tongue-like, palp set in deep square excavation, distolateral corner produced. Cheliped of mature male subequal to or exceeding carapace length. Maximum cl. 35 mm. Intertidal–shelf (0–40 m). Tropical W Atlantic. 1 species (Rathbun 1925: description as M. holderi Stimpson, 1871; Windsor and Felder 2014: rediagnosis).

Omalacantha Streets, 1871 Diagnosis. Carapace pyriform, longer than wide, branchial regions weakly inflated, tuberculate; row of tubercles parallel to posterior margin; anterolateral margin with 1 well defined posterolateral tooth. Rostrum comprising 2 elongate spines. Antennal basal article with 1–3 spinulose spines, anterolateral spine visible in dorsal view. Maxilliped 3 merus mesial margin straight, palp set in triangular notch with mesiodistal tooth. Cheliped of mature male less than carapace length. Maximum cl. 14 mm. Subtidal, shelf (1–70 m). Temperate Northern and Tropical W Atlantic. 4 species (Lemos de Castro 1953: figure; Windsor and Felder 2014: rediagnosis).

Petramithrax Windsor & Felder, 2014 Diagnosis. Carapace ovoid, about as wide as long, smooth; anterolateral margin with upper and lower parallel rows of small teeth. Rostrum comprising 2 short broad lobes. Orbital margin with 2 small teeth behind small preorbital tooth. Antennal basal article with lateral tooth and distolateral tooth visible in dorsal view. Maxilliped 3 merus mesial margin straight, palp set in oblique excavation, distolateral angle rounded-square. Cheliped of mature male twice as long as carapace. Pereopods 2–4 meri with distal spine. Pereopod 2 longer than carapace length by about dactylus plus half propodus. Maximum cl. 13 mm. Subtidal, shelf (0–50 m). Tropical Eastern Pacific. 1 species (Garth 1958: description of as Mithrax pygmaeus Bell, 1836; Windsor and Felder 2014: rediagnosis).

Pitho Bell, 1836 Diagnosis. Carapace ovoid, as long or longer than wide, with truncate front, smooth or tuberculate; anterolateral margin with 4–6 acute or rounded hepatic and branchial teeth, continuous with posterolateral margin. Rostrum comprising 2 blunt teeth. Orbital margin without teeth. Antennal basal article with blunt teeth or tubercles on distal margin, with inner suborbital tooth; antenna peduncular article 4 rectangular, narrower than basal article. Maxilliped 3 merus mesial margin short, palp set on oblique margin. Maximum cl. 33 mm (Pl. 54i). Subtidal, shelf (1–82 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 10 species (Garth 1958: key to 3 E Pacific species; Rathbun 1925: key to 9 species; Windsor and Felder 2014: rediagnosis).

14 – Brachyura – crabs

Teleophrys Stimpson, 1860 Diagnosis. Carapace broadly pyriform, with wide front, tuberculate; anterolateral margin with 3 lobes or teeth, with tooth at or near posterolateral angle. Rostrum comprising 2 rounded teeth. Orbital margin without teeth. Antennal basal article narrower distally, with rounded distolateral lobe, with elongate tubercle or subacute tooth close to basal article. Maxilliped 3 merus mesial margin broadly rounded, without concavity for palp. Pereopods 2–4 meri upper margins with spines, cristae or lobes; carpi and propodi with prominent teeth or lobes. Maximum cl. 18 mm (Pl. 54j). Subtidal, shelf (1–46 m). Tropical W Atlantic, Tropical Eastern Pacific. 5 species (Garth 1958: key to 3 E Pacific species; Rathbun 1925: key to 4 species; Windsor and Felder 2014: rediagnosis).

Thoe Bell, 1835 Diagnosis. Carapace pyriform, longer than wide, smooth; anterolateral margin with low tubercles and granules, sometimes with 1 small tooth at posterolateral angle. Rostrum comprising 2 teeth, sometimes partially fused. Orbital margin not protecting eyestalk from above, without preorbital spine. Antennal basal article with 0–2 anterior teeth or lobes visible in dorsal view. Maxilliped 3 merus mesial margin short, palp set on oblique margin. Pereopods 2–4 meri flat, with lamellate expansion on lower margins. Maximum cl. 16 mm (Pl. 54k). Intertidal–shelf (0–27 m). Tropical W Atlantic, Tropical Eastern Pacific. 3 species (Garth 1958: key to species, all now T. erosa Bell, 1836; Rathbun 1925: key to 4 species; Windsor and Felder 2014: rediagnosis). References Baeza JA, Bolaños JA, Fuentes S, Hernandez JE, Lira C, et  al. (2010) Molecular phylogeny of enigmatic Caribbean spider crabs from the Mithrax–Mithraculus species complex (Brachyura: Majidae: Mithracinae): ecological diversity and a formal test of genera monophyly. Journal of the Marine Biological Association of the United Kingdom 90, 851–858. doi:10.1017/S0025315409991044 Garth JS (1958) Brachyura of the Pacific coast of America. Oxyrhyncha. Allan Hancock Pacific Expeditions 21, 1–854. Griffin DJG, Tranter HA (1986) The Decapoda Brachyura of the Siboga Expedition. Part VIII Majidae. Siboga-Expéditie 39 c4, 1–335. Hultgren KM, Stachowicz JJ (2008) Molecular phylogeny of the brachyuran crab superfamily Majoidea indicates close congruence with trees based on larval morphology. Molecular Phylogenetics and Evolution 48, 986–996. doi:10.1016/j.ympev.2008.05.004 Klompmaker AA, Portell RW, Klier AT, Prueter V, Tucker AL (2015) Spider crabs of the Western Atlantic with special reference to fossil and some modern Mithracidae. PeerJ 3, e1301. doi:10.7717/peerj.1301 Lemos de Castro A (1953) Fauna do Distrito Federal. IX. Descrição de uma nova espécie do gênero “Eucinetops” Stimpson (Decapoda, Majidae). Revista Brasileira de Biologia 13, 355–358. Rathbun MJ (1925) The spider crabs of America. Bulletin of the United States National Museum 129, 1–613. Wagner HP (1990) The genera Mithrax Latreille, 1818 and Mithraculus White, 1847 (Crustacea: Brachyura: Majidae) in the Western Atlantic Ocean. Zoölogische Verhandelingen 264, 1–65.

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Windsor A, Felder D (2009) Re-evaluation of species allied to Mithrax hispidus (Decapoda: Brachyura: Majoidea: Mithracidae) based on three mitochondrial genes. Zootaxa 2302, 61–68. doi:10.11646/zootaxa.​ 2302.1.5 Windsor AM, Felder DL (2011) A new species of Hemus (Majoidea: Majidae: Mithracinae) from the Pacific coast of Panamá, with a key to the genus. Zootaxa 2799, 63–68. doi:10.11646/zootaxa.2799.1.6 Windsor AM, Felder DL (2014) Molecular phylogenetics and taxonomic reanalysis of the family Mithracidae MacLeay (Decapoda : Brachyura : Majoidea). Invertebrate Systematics 28, 145–173. doi:10.1071/ IS13011 Windsor AM, Felder DL (2017) Corrigendum to: Molecular phylogenetics and taxonomic reanalysis of the family Mithracidae MacLeay (Decapoda: Brachyura: Majoidea). Invertebrate Systematics 31, 232. doi:10.1071/IS13011_CO

Oregoniidae Garth, 1958 Figures 14.91–14.93, Plate 55 Garth (1958) erected the subfamily Oregoniinae for three majoid genera from the Arctic, North Pacific and North Atlantic having the male pleon widened posteriorly, the telson recessed in the concave posterior margin of pleonite 6, and gonopod 1 having a longitudinal groove and prominent setae. A fourth genus was added to this group by Sakai (1978). Ng et al. (2008) raised the subfamily to familial status but retained the same generic composition. More recently, oregoniids have been found to share zoeal characters and have a close molecular relationship with Ergasticus and several other genera usually considered to be inachids (Marco-Herrero et al. 2013) prompting their inclusion as a subfamily Pleistacanthinae of Oregoniidae, alongside Oregoniinae. Reconsideration of these genera along with several ‘inachids’ revealed that they share with other oregoniids the convex posterior margin of male pleonite 6 into which the telson is recessed, although the degree of convexity ranges from marked to gentle – certainly not straight as in Inachidae sensu stricto. Therefore, we include these former inachid genera in Oregoniidae: Bothromaia, Cyrtomaia, Ergasticus, Grypacheus, Macrocheira, Parapleisticantha, Platymaia, Pleistacantha, Pleisticanthoides, Trichoplatus and Vitjazmaia. These former inachid genera would correspond to Pleistacanthinae and a currently unused name, Macrocheirinae. Notably, these genera contain most of the large ‘inachids’; most remaining inachids being small, 30 mm carapace length or less. Although existing taxonomic names are already available for groups within Oregoniidae, the systematics is complicated and requires further detailed study to enable meaningful subfamilial or separate familial delineation. Therefore, here, we do not use subfamilies. While all oregoniids have a gonopod 1 with a longitudinal groove, only those originally included have prominent subdistal setae. All those currently included have the telson

640

Marine Decapod Crustacea

more or less sunken into pleonite 6 but this feature is more pronounced in some genera than others. Species of Chionoecetes, Macroregonia, some species of Pleistacantha and Macrocheira are large crabs, the last exceptionally so, with leg spans of almost four metres. Diagnosis. Orbit absent, eyes unprotected though orbital margin usually with several small spines and postorbital spine; basal antennal article slender, usually more than twice

as long as wide, always fused distally. Thoracic pleurites 5–8 not visible dorsally, covered by posterolateral margin of carapace. Eyestalk visible. Male pleonite 6 distal margin gently to strongly concave, telson recessed into concavity (if pleonite 6 and telson fused, traces of concave posterior margin of pleonite 6 visible); pleonites 3–5 always free. Pleonite 1 free from carapace. Gonopod 1 with prominent longitudinal groove.

Key to genera of Oregoniidae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. – 9. – 10. – 11.

Pereopods 2–5 propodi distally expanded, subchelate (Fig. 14.92r). Supraorbital eave laterally convex (Fig. 14.92g, m)��������������������������������������������������������������������������������������������������������������������������������������������������������Trichoplatus Pereopods 2–5 propodi linear. Supraorbital eave straight or weakly curved (Fig. 14.92a–f, h)�������������������������������������������2 Supraorbital margin with spines between preorbital and postorbital spines. Rostrum usually with lateral spines (Fig. 14.92a–f)������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Supraorbital margin smooth. Rostrum without lateral spines (Figs 14.91c–h)����������������������������������������������������������������������8 Rostrum comprising pair of lobes with radiating cluster of spines. Carapace gastric and branchial regions smooth, with deep urogastric pit overhung by pair of cardiac spines and mesogastric spine (Fig. 14.92a, i)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������Bothromaia Rostrum comprising pair of slender spines, sometimes with accessory spines, antler-like. Carapace spinose and/or setose, without deep urogastric pit������������������������������������������������������������������������������������������������������������������������������4 Pereopod 5 shorter than others, prehensile (Fig. 14.92s)��������������������������������������������������������������������������������������Grypachaeus Pereopods 2–5 similar, not prehensile������������������������������������������������������������������������������������������������������������������������������������������5 Carapace with anterior margin of antennular sinus unarmed. N Atlantic������������������������������������������������������������Ergasticus Carapace with anterior margin of antennular sinus produced to form anterolaterally directed spine (Fig. 14.92 l). Indo-West Pacific������������������������������������������������������������������������������������������������������������������������������������������������6 Carapace dorsal surface granulate, with few spinules or setae (Fig. 14.92f). Pereopods 2–4 meri and propodi lined with long stiff setae. Gonopod 1 slender, with subapical flange, without subapical process (Fig. 14.93t)����������������������������������������������������������������������������������������������������������������������������������������� Pleisticanthoides Carapace dorsal surface spiny and granular (Fig. 14.92d, e). Pereopods 2–4 meri and propodi lined with spines and spinules. Gonopod 1 slender, with or without subapical process�����������������������������������������������������������7 Rostral spines various, usually longer than supraorbital margin (Fig. 14.92e). Gonopod 1 slender, with subapical process (Fig. 14.93s)��������������������������������������������������������������������������������������������������������������������Pleistacantha Rostral spines about as long as supraorbital margin (Fig. 14.92d). Gonopod 1 stout, without subapical process (Fig. 14.93q)���������������������������������������������������������������������������������������������������������������������Parapleisticantha Carapace circular to subcircular (Fig. 14.91c, g, h). Gonopod 1 without subdistal longitudinal row of stiff setae (Fig. 14.93m, r)�������������������������������������������������������������������������������������������������������������������������������������������������������9 Carapace pyriform (Fig. 14.91a, d, f). Rostral spines paired, long or short, not short and diverging. Gonopod 1 with subdistal longitudinal row of stiff setae (Fig. 14.93c, d)������������������������������������������������������������������������11 Interantennular spine deflexed in adults, usually shorter than lateral rostral spines; no spine much longer than width at base (Fig. 14.92o). Pereopods 2–5 propodi narrow, cylindrical�������������������������������������������������Cyrtomaia Interantennular spine horizontal, longer than lateral rostral spines (Fig. 14.92q). Pereopods 2–5 propodi widened, flat or oval in cross-section (Fig. 14.91h)��������������������������������������������������������������������������������������������������������������11 Cornea diameter greater than or equal to eyestalk length. Hepatic spine shorter than twice cornea diameter. Pereopods 3–5 smooth, flat, with setose margins (Fig. 14.91h)���������������������������������������������������������Platymaia Cornea diameter less than eyestalk length. Hepatic spine longer than 3 times cornea diameter (Fig. 14.92h). Pereopods 3–5 spinose, oval in cross-section, with long strong marginal setae���������������������������������������������Vitjazmaia Male telson less than twice as wide as long, triangular (Fig. 14.93e). Gonopod 1 without subdistal longitudinal row of stiff setae. Basal antennal article movable�����������������������������������������������������������������������Macrocheira

14 – Brachyura – crabs

641

Fig. 14.91.  Oregoniidae. Carapace, representative or only species: a, Bothromaia; b, Chionoecetes; c, Cyrtomaia; d, Hyas; e, Macroregonia; f, Oregonia. Habitus: g, Macrocheira kaempferi (Temminck, 1836); h, Platymaia aff. wyvillethomsoni Miers, 1885.



Male telson more than twice as wide as long, posterior margin ovate, broadly angular or emarginate (Fig. 14.93a, d, f, g). Gonopod 1 with subdistal longitudinal row of stiff setae (Fig. 14.93n, p). Basal antennal article fixed��������������������������������������������������������������������������������������������������������������������������������������������������������������� 12 12. Rostral spines spiniform, separate; orbit, postorbital spine absent (Fig. 14.91e). Male pleonite 6 with convex parallel sides (Fig. 14.93f)������������������������������������������������������������������������������������������������������������������������������������ Macroregonia – Rostral spines wide, triangular, lobate, separate or appressed; orbit, postorbital spine present (Fig. 14.91b, d, f). Male pleonite 6 with sides not parallel (Fig. 14.93a, d, g)����������������������������������������������������������������������������������������������13

642

Marine Decapod Crustacea

Fig. 14.92.  Oregoniidae. Rostrum, anterior carapace: a, Bothromaia; b, Ergasticus; c, Grypachaeus; d, Parapleisticantha; e, Pleistacantha; f, Pleisticanthoides; g, Trichoplatus; h, Vitjazmaia. Rostrum, right antennule, antenna, epistome: i, Bothromaia; j, Cyrtomaia; k, Grypachaeus; l, Parapleisticantha; m, Trichoplatus. Antennal basal article, first movable article: n, Hyas. Rostrum, anterior carapace, lateral view: o, Cyrtomaia; p, Ergasticus; q, Platymaia. Pereopod 2: r, Trichoplatus. Pereopod 5: s, Grypachaeus. Features labelled on b, m: aba, antennal basal article; ao, antorbital spine; i, intercalated spine; n, nephridiopore (antennal gland); po, preorbital spine; ps, postorbital spine; r, rostral spine.

13. – 14. –

Carapace wider than long, or very little longer than wide; rostrum about as long as wide at base (Fig. 14.91b). Cheliped of adult male much shorter than pereopods 2–5. Male pleonite 6 tapering distally (Fig. 14.93a)�������������������������������������������������������������������������������������������������������������������������������������������������������������Chionoecetes Carapace much longer than wide; rostrum longer than width at base (Fig. 14.91d, f). Cheliped of adult male longer or little shorter than pereopods 2–5. Male pleonite 6 dilating distally (Fig. 14.93d, g)�������������������������������������14 Carapace with constriction behind postorbital tooth; postorbital spine remote from eye, affording no concealment (Fig. 14.91f). Antenna, first movable article not broadened or flattened������������������������������������� Oregonia Carapace with hepatic region broad, merged with postorbital tooth; postorbital tooth close to eye, cupped, affording partial concealment (Fig. 14.91d). Antenna, first movable article broadened, flattened (Fig. 14.92n)���������������������������������������������������������������������������������������������������������������������������������������������������������������������������Hyas Bothromaia Williams & Moffitt, 1991

Diagnosis. Carapace pyriform, longer than wide; gastric and branchial regions smooth, with deep urogastric pit overhung by pair of cardiac spines and mesogastric spine. Rostrum comprising pair of short lobes with radiating cluster of spines. Interantennular spine curving anteriorly. Supraorbital eave with spines between preorbital and antorbital spines; supraorbital eave postorbital spine remote from eye, affording no concealment; supraorbital eave without intercalated spine. Pereopods 2–4 meri and propodi lined with spines and spinules. Gonopod 1 stout, with subapical

process. Male pleon almost parallel-sided over distal half at least, telson triangular. Maximum cl. 33 mm. Slope (366–379 m). Central Indo-Pacific (Mariana archipelago). 1 species (Williams and Moffitt 1991).

Chionoecetes Krøyer, 1838 snow crab, tanner crab Diagnosis. Carapace pyriform, wider than long or very little longer than wide. Rostrum comprising pair of short, wide, separate

14 – Brachyura – crabs

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Fig. 14.93.  Oregoniidae. Male pleon: a, Chionoecetes; b, Cyrtomaia; c, Ergasticus; d, Hyas; e, Macrocheira; f, Macroregonia; g, Oregonia; h, Parapleisticantha; i, Pleistacantha; j, Pleisticanthoides; k, Trichoplatus. Gonopod 1: l, Bothromaia; m, Cyrtomaia; n, Hyas (apex only); o, Grypachaeus; p, Macroregonia (with detail of apex); q, Parapleisticantha; r, Platymaia; s, Pleistacantha; t, Pleisticanthoides.

triangular or bluntly angular teeth. Supraorbital eave postorbital tooth close to eye, cupped, affording partial concealment. Cheliped of adult male much shorter than pereopods 2–5. Gonopod 1 almost straight, with subdistal longitudinal rows of stiff setae. Male pleon tapering, telson more than twice wider than long. Male pleonite 6 tapering distally. Maximum cl. 126 mm. Intertidal–bathyal (0–2222 m). Arctic, Temperate Northern Atlantic, Temperate Northern Pacific. 7 species (Garth 1958; Sakai 1978). Species of snow crabs are among the largest crabs known, even the largest invertebrates. All species of Chionoecetes are intensively studied as fishery subjects (e.g. Cote et  al. 2019; Murphy 2021). One species, C. opilio (Fabricius, 1788), has been introduced to the Barents Sea (Kuzmin et  al. 1998) probably by shipping and has invaded the Kara Sea (Zalota et al. 2018) as the water has warmed. The same species may be contracting its range elsewhere (Fedewa et al. 2020).

Cyrtomaia Miers, 1886 Diagnosis. Carapace subcircular; with dorsal spines. Rostrum comprising 2 spines or teeth. Interantennular spine deflexed, usually shorter than rostral spines. Supraorbital eave with short postorbital spine. Pereopods 3–5 propodi slender, cylindrical. Gonopod 1 slender, tapering, curved. Maximum cl. 80 mm (Pl. 55a, b).

Shelf, slope (120–1000 m). Indo-West Pacific, Temperate ­ ustralasia. 29 species (Griffin and Tranter 1986: key to 18 speA cies; Guinot and Richer de Forges 1982: rediagnosis; Richer de Forges et  al. 2009; Richer de Forges and Ng 2007: species list, discussion)

Ergasticus A. Milne-Edwards, 1882 Diagnosis. Carapace pyriform, longer than wide; surface granular, with pair of cardiac spines, pair of hepatic spines, 1 or 2 pairs of branchial spines, 1 mesogastric spine. Rostrum comprising pair of slender spines, with accessory spines. Interantennular spine curving anteriorly; anterior margin of antennular sinus unarmed. Supraorbital eave with spines between preorbital and antorbital spines; supraorbital eave postorbital spine remote from eye, affording no concealment; supraorbital eave without intercalated spine. Pereopods 2–4 meri and propodi lined with spines and spinules. Gonopod 1 slender, with recurved subapical process. Male pleon almost parallel-sided over distal half at least, telson ovate. Maximum cl. 25 mm. Shelf, slope (70–1000 m). Temperate Northern and Tropical Atlantic. 1 species (Manning and Holthuis 1981: rediagnosis; Marco-Herrero et al. 2013: affinities; Milne-Edwards and Bouvier 1900: figure).

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Marine Decapod Crustacea

Grypachaeus Alcock, 1895 Diagnosis. Carapace pyriform, longer than wide; with dorsal spines. Rostrum comprising 2 spines more or less basally fused. Interantennular spine present. Supraorbital eave with preorbital and antorbital spines; supraorbital eave postorbital spine remote from eye, affording no concealment; supraorbital eave with intercalated spine. Pereopods 2–4 meri and propodi lined with spines and spinules. Pereopod 5 shorter than others, prehensile. Gonopod 1 slender, with subapical flange. Male pleon tapering, telson and somite 6 fused. Maximum cl. 14 mm (Pl. 55c). Shelf, slope (50–600 m). Western Indo-Pacific, Central IndoPacific. 2 species (Griffin and Tranter 1986: key to 2 species; Takeda 1978: description of second species).

Hyas Leach, 1814 Diagnosis. Carapace pyriform, longer than wide; hepatic region broad, merged with postorbital lobe. Rostrum comprising pair of contiguous spines, one-quarter postorbital carapace length. Supraorbital eave postorbital spine remote from eye, affording no concealment. Cheliped of adult male little shorter than pereopods 2–5. Male pleon almost parallel-sided over distal half, telson wider than long. Gonopod 1 almost straight, with subdistal longitudinal rows of stiff setae. Male pleonite 6 dilating distally; telson more than twice wider than long. Maximum cl. 95 mm (Pl. 55f). Intertidal–slope (0–1650 m). Arctic, Temperate Northern Atlantic, Temperate Northern Pacific. 5 species (Garth 1958: key to 2 Pacific species; Rathbun 1925: key to 4 species). Hyas is one of few brachyuran species in the Arctic and has become more common during the last century (Berge et al. 2009).

Macrocheira De Haan, 1839 Japanese spider crab Diagnosis. Carapace pyriform; with dorsal tubercles. Rostrum comprising 2 short diverging spines. Interantennular spine present; anterior margin of antennular sinus unarmed. Supraorbital eave with preorbital and antorbital spines; supraorbital eave with short postorbital spine; supraorbital eave without intercalated spine. Basal antennal article movable. Male telson distinctly less than twice as wide as long, triangular. Gonopod 1 straight, tightly twisted, distally styliform, without subdistal longitudinal row of stiff setae. Maximum cl. 335 mm (Pl. 55d). Shelf, slope (50–300 m). Temperate Northern W Pacific. 1 species (figures: De Haan 1833–1850; Sakai 1976). Macrocheira kaempferi (Temminck, 1836) is the only species and is the largest crab known with a leg span of almost 4 m and a weight of 18 kg. The zoea was described by Clark and Webber (1991) leading them to propose family status for this species.

Macroregonia Sakai, 1978 Diagnosis. Carapace oval, longer than wide, hepatic region broad. Rostral spines short spiniform, widely separate. Supraorbital eave and postorbital spine absent. Cheliped of adult male longer than

pereopods 2–5. Male pleon almost parallel-sided over distal half, telson wider than long. Gonopod 1 thin, curved, with subdistal longitudinal rows of stiff setae. Male pleonite 6 parallel-sided; telson more than twice wider than long. Maximum cl. 122 mm. Slope (800–1100 m). Temperate Northern Pacific. 1 species (Sakai 1978).

Oregonia Dana, 1851 Diagnosis. Carapace pyriform, longer than wide; with constriction behind postorbital lobe. Rostrum comprising pair of largely contiguous spines, half postorbital carapace length or less. Supraorbital eave postorbital tooth close to eye, cupped, affording partial concealment. Cheliped of adult male longer than pereopods 2–5. Gonopod 1 almost straight, with subdistal longitudinal rows of stiff setae. Male pleon almost parallel-sided over distal half, telson more than twice wider than long. Male pleonite 6 dilating distally. Maximum cl. 60 mm. Intertidal–slope (0–436 m). Temperate Northern Pacific. 2 species (keys to species: Garth 1958; Sakai 1976).

Parapleisticantha Yokoya, 1933 Diagnosis. Carapace pyriform, longer than wide; dorsal surface covered with spiniform granules and spinules with several larger, scattered spines. Rostral spines divergent, widely separated along its entire length, with lateral spinules, about as long as supraorbital margin. Interantennular spine present; anterior margin of antennular sinus produced to form anterolaterally directed spine. Supraorbital eave with spines between preorbital and antorbital spines; supraorbital eave postorbital spine remote from eye, affording no concealment; supraorbital eave with intercalated spine. Pereopods 2–4 meri and propodi lined with spines and spinules. Gonopod 1 short, straight, without subdistal process, tip turning outwards at 90°. Maximum cl. 27 mm. Shelf, slope (152–540 m). Temperate Northern Pacific, Central Indo-Pacific. 2 species (Richer de Forges et al. 2013: rediagnosis, species descriptions)

Platymaia Miers, 1885 Diagnosis. Carapace subcircular; with dorsal spines, or without dorsal spines. Rostrum comprising 2 spines. Interantennular spine horizontal, longer than rostral spines. Supraorbital eave without intercalated spine. Postorbital spine less than twice eye diameter. Cornea diameter greater than or equal to eyestalk length. Pereopods 3–5 propodi smooth, flat, with setose margins. Gonopod 1 slender, tapering, curved. Maximum cl. 60 mm (Pl. 55e). Shelf, slope (184–880 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia. 9 species (Griffin and Tranter 1986: key to 6 species; Guinot and Richer de Forges 1986: rediagnosis, key to 9 species). Species of Platymaia use their long flat posterior pereopods with long marginal setae to swim off the bottom (Wicksten and Bingo 2021). Platymaia longimana Macpherson, 1984 from off south-west Africa is referable to Vitjazmaia.

14 – Brachyura – crabs

Pleistacantha Miers, 1879 Diagnosis. Carapace pyriform, longer than wide; dorsal surface densely covered by spines and granules. Rostral spines divergent or contiguous spines, with or without lateral and other secondary spines, usually longer than supraorbital margin. Interantennular present; anterior margin of antennular sinus produced to form anterolaterally directed spine. Supraorbital eave with spines between preorbital and antorbital spines; supraorbital eave postorbital spine remote from eye, affording no concealment; supraorbital eave with intercalated spine. Pereopods 2–4 meri and propodi lined with spines and spinules. Gonopod 1 slender, with subapical flange. Male pleon tapering, telson and pleonite 6 fused or free. Maximum cl. 146 mm (Pl. 55h). Shelf, slope (50–723 m). Temperate Northern Pacific, Western and Central Indo-Pacific. 13 species (Ahyong et al. 2019; Ahyong et al. 2005: key to 9 species; Ahyong and Lee 2006; Ahyong and Ng 2007; Takeda and Miyake 1969).

Pleisticanthoides Yokoya, 1933 Diagnosis. Carapace pyriform, longer than wide; dorsal surface with spinules or granules. Rostral spines short, divergent without lateral spinules, about as long as supraorbital margin. Interantennular divided, short; anterior margin of antennular sinus produced to form anterolaterally directed spine. Supraorbital eave with spines between preorbital and antorbital spines (usually); supraorbital eave postorbital spine remote from eye, affording no concealment; supraorbital eave without intercalated spine. Pereopods 2–4 meri and propodi lined with stiff setae. Gonopod 1 slender, without recurved subapical process. Male pleon almost parallel-sided over distal half at least, telson triangular. Maximum cl. 24 mm. Shelf, slope (123–780 m). Temperate Northern Pacific, Central Indo-Pacific. 3 species (Ng and Richer de Forges 2012: rediagnosis).

Trichoplatus A. Milne-Edwards, 1876 Diagnosis. Carapace pyriform, longer than wide. Rostrum comprising 2 spines. Interantennular spine present. Supraorbital eave laterally convex; supraorbital eave postorbital spine remote from eave; supraorbital eave intercalated spine with anterior basal spinule. Pereopods 2–4 propodi distally expanded, subchelate. Gonopod 1 slender, bulbous basally, curved distally, weakly expanded, truncate. Male pleon almost parallel-sided over distal half, telson wider than long. Male pleonite 6 longer than wide, fused with telson. Maximum cl. 82 mm. Intertidal–shelf (0–55 m). Temperate Australasia (New Zealand endemic). 1 species (Griffin 1966: redescription as Naxia huttoni A. Milne-Edwards, 1876).

Vitjazmaia Zarenkov, 1994 Diagnosis. Carapace subcircular; with dorsal spines. Rostral spines spiniform, suprarallel. Interantennular spine horizontal, longer than rostral spines. Supraorbital eave without intercalated spine. Hepatic spine longer than 3 times eye diameter. Cornea diameter less than eyestalk length. Pereopods 3–5 propodi spinose, oval in

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cross-section, with long strong marginal setae. Gonopod 1 slender, tapering, curved. Maximum cl. 80 mm (Pl. 55g). Shelf–bathyal (154–2058 m). Western Indo-Pacific, Temperate Australasia, South-east Atlantic. 3 species (Webber and Richer de Forges 1995). Platymaia longimana Macpherson, 1984 is a species of Vitjazmaia. References Ahyong ST, Lee T (2006) Two new species of Pleistacantha Miers, 1879 from the Indo-West Pacific region (Crustacea, Decapoda, Majidae). Zootaxa 1378, 1–17. doi:10.11646/zootaxa.1378.1.1 Ahyong ST, Ng PKL (2007) Pleistacantha ori, a new species of deep-water spider crab (Crustacea: Decapoda: Brachyura: Majidae) from the Western Indian Ocean. Raffles Bulletin of Zoology Supplement 16, 67–74. Ahyong ST, Chen H-L, Ng PKL (2005) Pleistacantha stilipes, a new species of spider crab from the South China Sea (Decapoda: Brachyura: Majidae). Zootaxa 822, 1–10. doi:10.11646/zootaxa.822.1.1 Ahyong S, Mitra S, Ng PKL (2019) Redescription of Echinoplax rubida Alcock, 1895, a valid species of Pleistacantha from the Andaman Sea (Crustacea: Brachyura: Oregoniidae). Raffles Bulletin of Zoology 67, 391–395. Berge J, Renaud PE, Eiane K, Gulliksen B, Cottier FR, et al. (2009) Changes in the decapod fauna of an Arctic fjord during the last 100 years (1908–2007). Polar Biology 32, 953–961. doi:10.1007/s00300-009-0594-5 Clark PF, Webber WR (1991) A redescription of Macrocheira kaempferi (Temminck, 1836) zoeas with a discussion of the classification of the Majoidea Samouelle, 1819 (Crustacea: Brachyura). Journal of Natural History 25, 1259–1279. doi:10.1080/00222939100770781 Cote D, Nicolas J-M, Whoriskey F, Cook A, Broome J, et al. (2019) Characterizing snow crab (Chionoecetes opilio) movements in the Sydney Bight (Nova Scotia, Canada): a collaborative approach using multiscale acoustic telemetry. Canadian Journal of Fisheries and Aquatic Sciences 76, 334–346. doi:10.1139/cjfas-2017-0472 De Haan W (1833–1850) Crustacea. In Fauna Japonica sive descriptio animalium, quae in itinere per Japoniam, jussu et auspiciis superiorum, qui summum in India Batava Imperium tenet, suscepto, annis 1823–1830 collegit, notis, observionibus et asumnrationibus illustravit. (Ed. von Siebold PF) pp. 243, pls 1–50, Leiden. Fedewa EJ, Jackson TM, Richar JI, Gardner JL, Litzow MA (2020) Recent shifts in northern Bering Sea snow crab (Chionoecetes opilio) size structure and the potential role of climate-mediated range contraction. Deep-sea Research. Part II, Topical Studies in Oceanography 181–182, 104878. doi:10.1016/j.dsr2.2020.104878 Garth JS (1958) Brachyura of the Pacific coast of America. Oxyrhyncha. Allan Hancock Pacific Expeditions 21, 1–854. Griffin DJG (1966) The marine fauna of New Zealand: spider crabs, family Majidae (Crustacea, Brachyura). New Zealand Oceanographic Institute Memoir 172, 1–111. Griffin DJG, Tranter HA (1986) The Decapoda Brachyura of the Siboga Expedition. Part VIII Majidae. Siboga-Expéditie 39 c4, 1–335. Guinot D, Richer de Forges B (1982) Révision du genre Indo-Pacifique Cyrtomaia Miers, 1886: Campagnes océanographiques du Challenger, de l’Albatross, du Siboga et du Vauban (Crustacea Decapoda Brachyura). Annales de l’Institut Océanographique, Monaco et Paris 58, 5–88. Guinot D, Richer de Forges B (1986) Crustacés Décapodes: Majidae (genres Platymaia, Cyrtomaia, Pleistacantha, Sphenocarcinus et Naxioides). In: Résultats des Campagnes MUSORSTOM 1 & 2. Mémoires du Muséum National d’Histoire Naturelle, Paris, ser. Zoologie 133, 83–178. Kuzmin SA, Akhtarin SM, Menis DT (1998) The first finding of snow crab Chionoecetes opilio (Decapoda, Majidae) in the Barents Sea. Zoologicheskii Zhurnal 77, 489–491.

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Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Marco-Herrero E, Torres AP, Cuesta JA, Guerao G, Palero F, et al. (2013) The systematic position of Ergasticus (Decapoda, Brachyura) and allied genera, a molecular and morphological approach. Zoologica Scripta 42, 427–439. doi:10.1111/zsc.12012 Milne-Edwards A, Bouvier E-L (1900) Crustacés décapodes. Première partie. Brachyures et Anomoures. In Expéditions scientifiques du Travailleur et du Talisman pendant les années 1880, 1881, 1882, 1883. (Ed. Milne-Edwards A) pp. 1–396, 32 pls. Masson, Paris. Murphy JT (2021) Temporal and spatial variability in size-at-maturity for the eastern Bering Sea snow and Tanner crab (Chionoecetes opilio and C. bairdi). Fisheries Research 234, 105761. doi:10.1016/j.fishres.2020.105761 Ng PKL, Richer de Forges B (2012) Pleisticanthoides Yokoya, 1933, a valid genus of deep-sea inachid spider crabs (Crustacea: Decapoda: Brachyura: Majoidea), with descriptions of two new species from the Philippines, Papua New Guinea and Vanuatu. Zootaxa 3551, 65–81. doi:10.11646/zootaxa.3551.1.5 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Rathbun MJ (1925) The spider crabs of America. Bulletin of the United States National Museum 129, 1–613. Richer de Forges B, Ng PKL (2007) Notes on deep-sea spider crabs of the genus Cyrtomaia Miers, 1886, from the Philippines (Crustacea: Decapoda: Brachyura: Majidae), with description of a new species. Raffles Bulletin of Zoology Supplement 16, 55–65. Richer de Forges B, Li K, Ng PKL (2009) Cyrtomaia largoi Richer de Forges & Ng, 2007 (Decapoda, Inachidae): a new record for Taiwan. Crustaceana 82, 1205–1208. doi:10.1163/156854009X454649 Richer de Forges B, Ng PKL, Ahyong ST (2013) Parapleisticantha Yokoya, 1933, a valid genus of deep-sea inachid spider crab from Japan and the Philippines (Crustacea: Decapoda: Brachyura: Majoidea), with the description of a new species. Zootaxa 3635, 15–26. doi:10.11646/ zootaxa.3635.1.2 Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo. Sakai T (1978) Decapod Crustacea from the Emperor Seamount Chain. Researches on Crustacea Supplement 8, 1–39, pls 1–4. Takeda M (1978) A new spider crab from off Iriomote Island, the Ryukyu Islands. Bulletin of the National Science Museum, Ser. Zoology 2, 117–121. Takeda M, Miyake S (1969) Crabs from the East China Sea. III Brachygnatha Oxyrhyncha. Journal of the Faculty of Agriculture, Kyushu University 15, 469–521. doi:10.5109/22797 Webber WR, Richer de Forges B (1995) Deep sea Majidae (Decapoda: Brachyura) new to New Zealand with a description of Oxypleurodon wanganella sp. nov. Journal of the Royal Society of New Zealand 25, 501–516. doi:10.1080/03014223.1995.9517500 Wicksten MK, Bingo S (2021) Platymaia Miers, 1885 (Decapoda: Brachyura: Inachidae) can swim. Journal of Crustacean Biology 41, ruaa090. Williams AB, Moffitt RB (1991) Crabs from the Mariana Archipelago: Bothromaia griffini new genus and species (Brachyura; Majidae), and remarks on Poupinia hirsuta Guinot (Homoloidea, Poupiniidae). Proceedings of the Biological Society of Washington 104, 569–582. Zalota AK, Spiridonov VA, Vedenin AA (2018) Development of snow crab Chionoecetes opilio (Crustacea: Decapoda: Oregonidae) invasion in the Kara Sea. Polar Biology 41, 1983–1994. doi:10.1007/s00300-018-2337-y

Calappoidea but is probably not closely related (Bellwood 1996; Guinot et al. 2013; Ng et al. 2008). The paddle-shaped last legs are used for burrowing in sand but not for swimming. Diagnosis. Carapace longitudinally ovate-subhexagonal, regions well defined, granular. Buccal cavity trapezoid-­ triangular, margins distinctly convergent anteriorly. Maxilliped 3 merus triangular, tapering distally; exopod without flagellum. Pereopods 2–4 dactyl slender, pointed; pereopod 5 dactylus as wide, flattened paddle. Male pleon short, filling only part of cavity. Female vulva exposed, not concealed by pleon.

Orithyiidae Dana, 1852 tiger crab Figure 14.94, Plate 56a Orithyia Fabricius, 1798 Maximum cl. 90 mm (Pl. 56a). Subtidal, shelf (10–30 m). Temperate North W Pacific. 1 species (Dai and Yang 1991: diagnosis; Shen 1932: description, figure). Orithyia sinica (Linnaeus, 1771) is confined to continental environments and does not occur on offshore islands (Ng et al. 2008). The species is fished commercially in China and South Korea. Consequently, it is much studied.

Orithyioidea Dana, 1852 The superfamily contains just one family and one species but this rank is justified on its peculiar features. It resembles

Fig. 14.94.  Orithyiidae, Orithyia sinica (Linnaeus, 1771). a, habitus; b, female thoracic sternum, pleon with vulvae (shaded).

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References Bellwood O (1996) A phylogenetic study of the Calappidae H. Milne Edwards 1837 (Crustacea: Brachyura) with a reappraisal of the status of the family. Zoological Journal of the Linnean Society 118, 165–193. doi:10.1111/j.1096-3642.1996.tb00225.x Dai A-Y, Yang S-L (1991) Crabs of the China Seas. English edn. China Ocean Press and Springer-Verlag, Beijing and Berlin. Guinot D, Tavares M, Castro P (2013) Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higherranked podotreme taxa. Zootaxa 3665, 1–414. doi:10.11646/zootaxa.​ 3665.1.1 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Shen CJ (1932) The brachyuran Crustacea of North China. Zoologia Sinica. Series A. Invertebrates of China 9, 1–320, pls 1–10.

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Palicoidea Bouvier, 1898 Palicoids are typically crabs with a flat carapace and strongly unequal walking legs. The first pair of walking legs are smaller than the next two pairs and the last pair are considerably smaller. The two families differ in the degree of reduction of the last pair and their position relative to more anterior legs. The systematics of what was then a single family with two subfamilies was treated in detail by Castro (2000) who erected new genera for some Indo-West Pacific species. Diagnosis. Carapace subquadrate to ovate; dorsal surface typically flat, granular, often with prominent regions, without transverse ridges. Antennule folding transversely. Buccal cavity approximately quadrate. Maxilliped 3 merus subquadrate. Pereopods 2–4 as walking legs. Pereopod 5 strongly reduced or rudimentary.

Key to families of Palicoidea 1. –

Pereopod 5 shorter but similar to pereopods 2–4, articulating with carapace at same level (Fig. 14.95c). Pleonites 1, 2 of both sexes about as long as pleonite 3 (Fig. 14.95b)������������������������������������������������������� Crossotonotidae Pereopod 5 much shorter, more slender than pereopods 2–4, articulating with carapace more dorsally (Fig. 14.96c, f, h). Pleonites 1, 2 of both sexes much shorter than other pleonites, about half or less as long as pleonite 3 (Fig. 14.96j, k)���������������������������������������������������������������������������������������������������������������������������������Palicidae

Reference Castro P (2000) Crustacea Decapoda: a revision of the Indo-West Pacific species of palicid crabs (Brachyura Palicidae). In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 21. Mémoires du Muséum National d’Histoire Naturelle, Paris 184, 437–610.

Crossotonotidae Moosa & Serène, 1981 Figure 14.95, Plate 56b The last legs of crossonototids are not as modified in the same way as palicids, instead being similar to the two pairs of larger walking legs. Crossonototids inhabit rocky and coral reefs of the Indo-West Pacific, from intertidal to upper shelf depths.

Castro’s (2000) revision included two genera as members of the palicid subfamily Crossotonotinae; few species have been added since. Differences between the genera were discussed by Takeda and Tachikawa (2015). Diagnosis. Pereopod 5 shorter but similar to pereopods 2–4, and articulating with carapace at same level. Pleonites 1, 2 of both sexes about as long as pleonite 3, not dorsoventrally compressed, not much narrower than pleonites 3–6. Crossotonotus A. Milne-Edwards, 1873 Diagnosis. Carapace subquadrate, with 10–20 small, rounded or pointed tubercles along posterior border (Fig. 14.95a). Maximum cl. 31 mm (Pl. 56b).

Fig. 14.95.  Crossotonotidae. Crossotonotus: a, carapace; b, pleon. c, carapace, left pereopod 5, right pereopod 4, Pleurophricus.

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Intertidal–shelf (0–146 m). Indo-West Pacific. 4 species (Castro 2000: rediagnosis, key to species)

Pleurophricus A. Milne-Edwards, 1873 Diagnosis. Carapace subcircular, with 5 or 6 tubercles along posterior border (Fig. 14.95c). Maximum cl. 9 mm. Subtidal (3–18 m). Central Indo-Pacific. 3 species (Castro 2000: rediagnosis, key to 2 species; Takeda and Tachikawa 2015: third species, modified diagnosis). References Castro P (2000) Crustacea Decapoda: a revision of the Indo-West Pacific species of palicid crabs (Brachyura Palicidae). In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 21. Mémoires du Muséum National d’Histoire Naturelle, Paris 184, 437–610. Takeda M, Tachikawa H (2015) Crabs of the families Palicidae and Crossotonotidae (Crustacea, Decapoda, Brachyura, Palicoidea) from the Ogasawara Islands, Japan, with the description of a new species. Zootaxa 3918, 141–150. doi:10.11646/zootaxa.3918.1.7

Palicidae Bouvier, 1898 Figure 14.96, Plate 56c–f Palicidae is the larger of the two families of Palicoidea. They differ from crossotonotids in having the last pair of legs much more slender than the others. Palicids range from subtidal to mid-slope depths where they inhabit mostly

sandy and muddy benthic environments. The long thin legs enable them to slide across the mud. Species of Exopalicus tend to live on harder coral substrates (Castro 2000). Castro (2000) thoroughly reviewed the systematics of Indo-West Pacific species, now numbering 38. He rediagnosed genera already proposed by Moosa and Serène (1981) and added others. Castro (2000) did not address the systematics of Palicus, the only genus found in the Eastern Pacific and Atlantic. The 25 species of Palicus are also variable as documented by Rathbun (1918) who used the earlier preoccupied genus name Cymopolia Roux, 1828. She proposed six species groups for the American species, without considering the type species C. caronii Roux, 1828, from the northeastern Atlantic. The phylogeny of North American species of Palicus based on mitochondrial genes and gonopods was addressed by Pecnik (2016). Consistent differences between Palicus and other genera are hard to find in Castro’s (2000) table of characters. Diagnosis. Pereopod 5 much shorter, more slender than pereopods 2–4, and articulating with carapace more dorsally. Pleonites 1, 2 of both sexes much shorter than other pleonites, half or less as long as pleonite 3, dorsoventrally compressed, much narrower than pleonites 3–6. Implicit generic attributes. Carapace dorsally slightly convex. Eyestalks flattened, cornea wider than eyestalk. Pereopods 3, 4 propodi extensor margins not convexly expanded. Male pleon with subparallel sides. Female pleonites free.

Key to genera of Palicidae 1. – 2. – 3. – 4. – 5. – 6.

Pereopods 2–4 with flattened carpi and propodi; meri with unequal tubercles or teeth along margins; dactyli sometimes armed with teeth or spiniform setae along complete length of flexor margins (Fig. 14.96h)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Pereopods 2–4 slender with elongate meri, filiform propodi and dactyli; meri with equal or subequal, small or flat tubercles along margins; dactyli without teeth or spines along complete length of flexor margins (Fig. 14.96f)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������7 Cornea spherical, rarely flattened. East Pacific, Atlantic����������������������������������������������������������������������������������������������� Palicus Cornea flattened. Indo-West Pacific���������������������������������������������������������������������������������������������������������������������������������������������3 Carapace transversely hexagonal, noticeably wider than long, hexagonal; anterolateral margin strongly curved, with 3 or 4 teeth (Fig. 14.96a, g, h)�����������������������������������������������������������������������������������������������������������������������������4 Carapace quadrate-trapezoid, little wider than long; anterolateral margin straight or slightly curved, with truncate teeth (Fig. 14.96e, d, i)�������������������������������������������������������������������������������������������������������������������������������������������������5 Carapace anterolateral margin with 3 or 4 acute teeth; supraorbital margin with 2 V-shaped incisions (Fig. 14.96g, h)�������������������������������������������������������������������������������������������������������������������������������������������������������Pseudopalicus Carapace anterolateral margin with 3 teeth; supraorbital margin with 1 narrow incision (Fig. 14.96a)����������Exopalicus Supraorbital margin with 2 incisions (Fig. 14.96i). Pereopods 2–4 propodi, dactyli flexor margins with short spines (Fig. 14.96l)�����������������������������������������������������������������������������������������������������������������������������������������Rectopalicus Supraorbital margin with triangular lobes. Pereopods 2–4 propodi, dactyli flexor margins entire (Fig. 14.96c, f)������������������������������������������������������������������������������������������������������������������������������������������������������������������������������6 Eyestalks with 1 granular, median tubercle and 2 smaller, granular tubercles, cornea spherical, almost as wide as eyestalk. Pereopods 3, 4 propodi expanded (Fig. 14.96c)��������������������������������������������������������������������� Neopalicus

14 – Brachyura – crabs

– 7. – 8. –

649

Eyestalks with thin, crescent-shaped process on anterior margin, cornea spherical, almost as wide as eyestalk (Fig. 14.96d). Pereopods 3, 4 propodi not expanded������������������������������������������������������������������������������ Palicoides Carapace anterolateral margin with 1 triangular or rounded tooth (Fig. 14.96f). Male pleon triangular (Fig. 14.96k)��������������������������������������������������������������������������������������������������������������������������������������������������������������� Parapalicus Carapace anterolateral margin with 3 rounded teeth (Fig. 14.96). Male pleon with subparallel sides (Fig. 14.96j)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������8 Supraorbital margin with 1 shallow broad concavity. Carapace anterolateral margin with 3 rounded teeth (Fig. 14.96b). Female pleonites free������������������������������������������������������������������������������������������������������������������������Miropalicus Supraorbital margin with 1 or 2 rounded or pointed lobes. Carapace anterolateral margin with 2–4 acute teeth (Fig. 14.96e). Female pleonites 3–6 or 4–6 fused��������������������������������������������������������������������������������������������Paliculus Exopalicus Castro, 2000

Diagnosis. Carapace wider than long, hexagonal; anterolateral margin strongly curved, with 3 teeth; supraorbital margin with 1 incision. Pereopods 2–4 with flattened carpi and propodi; meri with unequal tubercles or teeth along margins; propodi, dactyli flexor margins with short spines. Maximum cl. 8.4 mm. Intertidal, subtidal (0–18 m). Indo-West Pacific. 1 species (Castro 2000).

Miropalicus Castro, 2000 Diagnosis. Carapace subovate; anterolateral margin with 3 rounded teeth; supraorbital margin with 1 shallow broad concavity. Pereopods 2–4 slender with elongate meri, filiform propodi and dactyli; meri with equal or subequal, small or flat tubercles

along margins; propodi, dactyli flexor margins with short spines. Maximum cl. 10.5 mm. Shelf, slope (245–710 m). Central Indo-Pacific. 1 species (Castro 2000).

Neopalicus Moosa & Serène, 1981 Diagnosis. Carapace subquadrate; anterolateral margin almost straight, with 2 large truncate teeth, sometimes with similar smaller more posterior teeth; supraorbital margin with triangular lobe defined by 2 incisions. Eyestalks with 1 granular, median tubercle and 2 smaller, granular tubercles, cornea spherical, almost as wide as eyestalk. Pereopods 2–4 with flattened carpi and propodi; meri with unequal tubercles or teeth along margins; propodi, dactyli flexor margins entire. Pereopods 3, 4 propodi extensor margins convexly expanded. Maximum cl. 17.3 mm (Pl. 56c).

Fig. 14.96.  Palicidae. Carapace, pereopods. a, Exopalicus; b, Miropalicus; c, Neopalicus; d, Palicoides whitei (Miers, 1879) e, Paliculus; f, Parapalicus; g, Pseudopalicus; h, Pseudopalicus macromeles Castro, 2000; i, Rectopalicus. Male pleon: j, Miropalicus; k, Parapalicus. l, pereropod 4 dactylus, Rectopalicus.

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Marine Decapod Crustacea

Slope (360–840 m). Indo-West Pacific. 4 species (Castro 2000: rediagnosis, key to 2 species; Castro and Naruse 2014: fourth species).

Palicoides Moosa & Serène, 1981 Diagnosis. Carapace subquadrate; anterolateral margin almost straight, with 2 large truncate teeth; supraorbital margin with triangular lobe defined by 2 incisions. Eyestalks with thin, calcified, crescent-shaped process on anterior margin, cornea spherical, almost as wide as eyestalk. Pereopods 2–4 with flattened carpi and propodi; meri with unequal tubercles or teeth along margins; propodi, dactyli flexor margins entire. Female pleonites 3–6 fused. Maximum cl. 20.5 mm. Subtidal, shelf (2–70 m). Western and Central Indo-Pacific. 2 species (Castro 2000: rediagnosis, key to species).

margin with 2 V-shaped incisions. Pereopods 2–4 with flattened carpi and propodi; meri with unequal tubercles or teeth along margins; propodi, dactyli flexor margins with tubercles or teeth. Maximum cl. 22.3 mm (Pl. 56e, f). Shelf, slope (30–660 m). Temperate Northern W Pacific, IndoWest Pacific, Temperate Australasia. 12 species (Castro 2000: rediagnosis, key to 11 species).

Rectopalicus Castro, 2000 Diagnosis. Carapace rounded-subquadrate; anterolateral margin almost straight, with short truncate teeth; supraorbital margin with 2 incisions. Pereopods 2–4 with flattened carpi and propodi; meri with unequal tubercles or teeth along margins; propodi, dactyli flexor margins with short spines. Maximum cl. 9.7 mm. Shelf, slope (85–430 m). Central Indo-Pacific. 3 species (Castro 2000: key to species).

Paliculus Castro, 2000 Diagnosis. Carapace subquadrate; anterolateral margin with 2–4 acute teeth; supraorbital margin with 1 or 2 rounded or pointed lobes. Pereopods 2–4 slender with elongate meri, filiform propodi and dactyli; meri with equal or subequal, small or flat tubercles along margins; propodi, dactyli flexor margins with tubercles or teeth. Female pleonites 3–6 fused, or 4–6 fused. Maximum cl. 5 mm (Pl. 56d). Subtidal–slope (3–710 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 2 species (Castro 2000: diagnosis, key to species).

Palicus Philippi, 1838 Diagnosis. Carapace subquadrate; anterolateral margin with 2 or 3 large flat teeth; supraorbital margin with 2 V-shaped incisions. Eyestalks tuberculate, cornea spherical, wider than eyestalk. Pereopods 2–4 with flattened carpi and propodi; meri with unequal tubercles or teeth along margins; propodi, dactyli flexor margins entire. Male pleon triangular, or with subparallel sides. Maximum cl. 13 mm. Subtidal–slope (7–686 m). Temperate Northern and Tropical Atlantic (including Mediterranean), Tropical Eastern Pacific. 25 species (Rathbun 1918: key to 20 American species as Cymopolia).

Parapalicus Moosa & Serène, 1981 Diagnosis. Carapace subhexagonal; anterolateral margin with 1 triangular or rounded tooth; supraorbital margin without or with short incisions. Pereopods 2–4 slender with elongate meri, filiform propodi and dactyli; meri with equal or subequal, small or flat tubercles along margins; propodi, dactyli flexor margins entire. Male pleon triangular. Female pleonites 3–5 fused. Maximum cl. 10.5 mm. Shelf, slope (49–950 m). Western and Central Indo-Pacific. 13 species (Castro 2000: rediagnosis, key to species).

Pseudopalicus Moosa & Serène, 1981 Diagnosis. Carapace wider than long, hexagonal; anterolateral margin strongly curved, with 3 or 4 acute teeth; supraorbital

References Castro P (2000) Crustacea Decapoda: a revision of the Indo-West Pacific species of palicid crabs (Brachyura Palicidae). In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 21. Mémoires du Muséum National d’Histoire Naturelle, Paris 184, 437–610. Castro P, Naruse T (2014) New species of Latreillopsis Henderson, 1888 (Brachyura: Homolidae) and Neopalicus Moosa & Serène, 1981 (Brachyura: Palicidae) from the Hawaiian Islands. Zootaxa 3764, 169–180. doi:10.11646/zootaxa.3764.2.4 Moosa K, Serène R (1981) Observations on the Indo-West-Pacific Palicidae (Crustacea: Decapoda) with descriptions of two new subfamilies, four new general and six new species. Marine Research in Indonesia 22, 21–66. doi:10.14203/mri.v22i0.432 Pecnik SJ (2016). Phylogenetic analysis of North American representatives of the brachyuran genus Palicus, with focus on gonopod morphology and mitochondrial gene sequences. MSc thesis. University of Louisiana at Lafayette, Lafayette. Rathbun MJ (1918) The grapsoid crabs of America. Bulletin of the United States National Museum 97, 1–461, pls 1–161.

Parthenopoidea MacLeay, 1838 The superfamily contains two families, many of whose genera are highly ornamented. Until recently, close relationships were presumed between parthenopid, aethrids and several leucosiids owing to similar habitus and seemingly intergrading mouthpart structures, but more detailed anatomical comparisons and phylogenetic studies, rooted in initial observations by Guinot (1966, 1967a, 1967b), justified clearer separation of the groups (Ng et  al. 2008). Guinot (1978) noted similarities between Dairoides and Daira, but also remarked on resemblance to parthenopoids, especially the wide thoracic sternum with incomplete sutures, a functional press-button attachment between the male pleon and sternum, a stout gonopod 1 and gonopod 2 with a long flagellum. Ng et al. (2008) regarded these similarities as convergences treating dairoidids as eriphioids, but Lai et  al.

14 – Brachyura – crabs

(2014) proposed that they may have a phylogenetic basis allying them to parthenopoids. Diagnosis. Carapace triangular, subpentagonal or ovatesubpentagonal, sometimes almost semicircular or semielliptical, anteriorly angular, regions usually demarcated with prominent grooves, usually uneven, tuberculate, granulate

651

or eroded. Antennule folding transversely or obliquely. Buccal cavity approximately quadrate. Maxilliped 3 merus irregularly subquadrate or rectangular. Cheliped palm triangular to ovate in cross-section. Male pleonites 3–4 or 3–5 fused, immovable, even if some or all sutures visible; pleonite 3 without transverse ridge. Gonopod 1 stout.

Key to families of Parthenopoidea 1. –

Carapace surface densely covered in interlocking, perforate tubercles forming pavement-like surface. Chelipeds short; minor chela with slender fingers (Fig. 14.97a). Male pleonites 3–4 fused, immovable, sutures visible. Gonopod 2 longer than gonopod 1 (Fig. 14.97c)����������������������������������������������������������������������Dairoididae Carapace surface smooth, or variously ornamented, not with interlocking, perforate tubercles. Chelipeds short or long; minor chela with short, stout fingers (Figs 14.98–14.103). Male pleonites 3–5 fused, immovable, even if sutures visible. Gonopod 2 usually shorter than (Fig. 14.103j, k), occasionally longer than, gonopod 1 (Fig. 14.103i, l)������������������������������������������������������������������������������������������������������������������������ Parthenopidae

References Guinot D (1966) Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures. I. Les affinités des genres Aethra, Osachila, Hepatus, Hepatella et Actaeomorpha. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 38, 744–762. Guinot D (1967a) Recherches préliminaires sur les groupements naturels chez les Crustacés décapodes brachyoures. III. A propos des affinités des genre Dairoides Stebbing et Daira De Haan. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 39, 540–563. Guinot D (1967b) Recherches préliminaires sur les groupements naturels chez les Crustacés. Décapodes Brachyoures I. Les affinités des genres Aethra, Osachila, Hepatus, Hepatella et Actaeomorpha (suite et fin). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 38, 828–845. Guinot D (1978) Principes d’une classification évolutive des Crustacés Décapodes Brachyoures. Bulletin Biologique de la France et de la Belgique 112, 211–292. Lai JCY, Thoma BP, Clark PF, Felder DL, Ng PKL (2014) Phylogeny of eriphioid crabs (Brachyura, Eriphioidea) inferred from molecular and

morphological studies. Zoologica Scripta 43, 52–64. doi:10.1111/ zsc.12030 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286.

Dairoididae Števčić, 2005 Figure 14.97, Plate 57a Dairoides is the only genus, recognisable by perforated, interlocking tubercles covering much of the carapace except for grooves separating some regions, and the prominently angular anterior part of the carapace. The species inhabit shelf and upper slope depths. Dairoidids have variously been regarded as xanthoids, eriphioids and parthenopoids, the last hypothesis first suggested by Guinot (1967) and more recently corroborated by molecular phylogenetics (Lai et al. 2014).

Fig. 14.97.  Dairoididae. a, Dairoides kusei (Sakai, 1938). Dairoides margaritatus Stebbing, 1920: b, carapace; c, gonopods 1, 2.

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Marine Decapod Crustacea

Diagnosis. Carapace approximately diamond-shaped to subtriangular, anteriorly strongly angular, densely covered in interlocking, perforate tubercles forming pavment-like surface. Cheliiped palm ovate in cross-section; minor chleliped with slender fingers. Minor chleliped with slender fingers. Male pleonites 3, 4 fused, immovable, sutures visible. Gonopod 2 longer than gonopod 1, distal portion strongly recurved, as long as proximal portion. Dairoides Stebbing, 1920 Maximum cl. 75 mm (Pl. 57a). Shelf, slope (20–633 m). Indo-West Pacific, Temperate Southern Africa. 3 species (Barnard 1950; Ng and Tan 1999: redescription; Sakai 1976; Takeda and Ananpongsuk 1991: species compared). References Barnard KH (1950) Descriptive catalogue of South African decapod Crustacea (crabs and shrimps). Annals of the South African Museum 38, 1–837. Guinot D (1967) Recherches préliminaires sur les groupements naturels chez les Crustacés décapodes brachyoures. III. A propos des affinités des genre Dairoides Stebbing et Daira De Haan. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 39, 540–563. Lai JCY, Thoma BP, Clark PF, Felder DL, Ng PKL (2014) Phylogeny of eriphioid crabs (Brachyura, Eriphioidea) inferred from molecular and morphological studies. Zoologica Scripta 43, 52–64. doi:10.1111/zsc.12030 Ng PKL, Tan SH (1999) The Hawaiian parthenopid crabs of the genera Garthambrus Ng, 1996, and Dairoides Stebbing, 1920 (Crustacea: Decapoda: Brachyura). Proceedings of the Biological Society of Washington 112, 120–132. Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo. Takeda M, Ananpongsuk S (1991) A new deep-sea crab from the Andaman Sea off Thailand. Bulletin of the National Science Museum, Ser. Zoology 17, 93–100.

Parthenopidae MacLeay, 1838 elbow crabs, buckler crabs Figures 14.98–14.103, Plates 57b–l, 58 Parthenopids are striking crabs displaying a variety of forms. They have in common long chelipeds that protrude laterally when folded, which gives them their common

name, elbow crabs. Many have an almost triangular carapace, some are highly sculptured while others are quite smooth. The sculptured species resemble pieces of coral rubble. The smooth species are commonly called buckler crabs. Most parthenopids live in broken coral or rock sediments, usually subtidally but have been recorded down to upper slope depths. Little is known of their ecology but some may be predators (Anker and Ivanov 2020; Mura and Orrù 1998; Zipser and Vermeij 1978). From above, some of the smooth, flattened forms, such as Celatopisa and Cryptopodia closely resemble leucosioids, such as Cryptocnemus and Nursia, so mouthparts should always be carefully checked. Papers dealing with the parthenopid fauna of the Americas (Garth 1958; Gore and Scotto 1979; Rathbun 1925) and of the Indo-West Pacific (Flipse 1930) are important sources for species identification but generic placements have changed significantly. The taxonomy of the family was synthesised in detail in a PhD thesis (Tan 2004) where new genera were suggested. Two major revisions resulted from Tan’s thesis. The first diagnosed the family and erected 12 new genera of one of the two subfamilies, Parthenopinae (Tan and Ng 2007a). The second dealt with Daldorfiinae, erecting two new genera (Tan and Ng 2007b). Two genera have been added since 2007 (McLay and Tan 2009). The two subfamilies are distinguished by the relative size of the basal articles of the antenna and antennule (Fig. 14.98c– e). Generic and specific differences between taxa in Parthenopidae depend on attention to the regions, margins and ridges of the carapace. Figure 14.98 explains the terminology used. Our key is based on those already published (Tan and Ng 2007a, 2007b), removing some ambiguity and adding more recently described genera. Genera are arranged alphabetically by subfamily. Keys to species of most genera can be found in Tan (2004). Diagnosis. Carapace approximately triangular to cordiform, variously tuberculate and eroded. Cheliped palm usually triangular in cross-section. Male pleonites 3–5 fused. Gonopod 2 usually shorter than, occasionally slightly longer than, gonopod 1, distal portion straight or gently curved, usually shorter than proximal portion.

Key to subfamilies and genera of Parthenopidae 1. – 2. – 3.

Antennal article 2 length as long or longer than antennular article 1 (Fig. 14.98c)�������������������������������� Daldorfiinae … 2 Antennal article 2 less than half as long as antennular article 1 (Fig. 14.98d, e)����������������������������������Parthenopinae … 5 Antennal article 2 proximal half enlarged, about twice width of the distal half (Fig. 14.99f). Gonopod 2 about half length of gonopod 1. Female telson longer than wide (Fig. 14.99j)���������������������������������������������������� Niobafia Antennal article 2 proximal half not enlarged (Fig. 14.99e, g, h). Gonopod 1 longer than half length of gonopod 2. Female telson wider than long (Fig. 14.99i, k, l)������������������������������������������������������������������������������������������������3 Cheliped manus slender, about half length of merus; upper meral spine reduced to granule(s), not overlapping epibranchial teeth (Fig. 14.99d)�������������������������������������������������������������������������������������������������� Thyrolambrus

14 – Brachyura – crabs



653

Cheliped manus about as long as merus; upper meral spine strong, overlapping epibranchial teeth (Fig. 14.99a, c)������������������������������������������������������������������������������������������������������������������������������������������������������������������������������4 4. Carapace transversely ovate (Fig. 14.99c). Antennal article 2 without anterolateral tooth (Fig. 14.99g). Female telson subtriangular, distolateral margins of tip constricted (Fig. 14.99k)�������������������������������������������Olenorfia – Carapace subtriangular to subpentagonal (Fig. 14.99a). Antennal article 2 with anterolateral tooth (less than half length of article 3; Fig. 14.99e). Female telson semicircular (Fig. 14.99i)������������������������������������������ Daldorfia 5. Rostrum longer than postorbital carapace length. Eyes not enclosed by shallow orbits (Fig. 14.101c)���������������������������������������������������������������������������������������������������������������������������������������������������������Lambrachaeus – Rostrum shorter than half carapace length. Eyes enclosed within deep orbits����������������������������������������������������������������������6 6. Carapace dorsal surface with 3 low, parallel ridges, one on each epibranchial region and one on cardiac region (Fig. 14.101e)������������������������������������������������������������������������������������������������������������������������������������������������Leiolambrus – Carapace dorsal surface without parallel ridges��������������������������������������������������������������������������������������������������������������������������7 7. Maxilliped 3 exopod completely hidden by maxilliped (Fig. 14.103g)������������������������������������������������������������� Mimilambrus – Maxilliped 3 exopod exposed or partially hidden by maxilliped���������������������������������������������������������������������������������������������8 8. Suborbital region with diagonal pterygostomial ridge angled 45° to sagittal plane, visible in dorsal view (Fig. 14.100a)������������������������������������������������������������������������������������������������������������������������������������������������������������ Agolambrus – Suborbital region without diagonal ridge�������������������������������������������������������������������������������������������������������������������������������������9 9. Epistome with projecting tubercle below antennular article 1 (Fig. 14.103a)�����������������������������������������������������������������������10 – Epistome without projecting tubercle below antennular article 1 (Fig. 14.103b)�����������������������������������������������������������������11 10. Carapace pterygostomial region excavated, forming distinct inhalent channel; pterygostomial ridge with dense, long setae covering inhalent channel; last epibranchial tooth much longer than more anterior teeth (Fig. 14.100b)���������������������������������������������������������������������������������������������������������������������������������������������Aulacolambrus – Carapace pterygostomial region not excavated, without distinct inhalent channel; pterygostomial ridge without dense setae; last epibranchial tooth slightly longer than more anterior teeth (Fig. 14.102b)������Piloslambrus 11. Carapace epibranchial marginal teeth subrectangular or weakly indicated, with narrow gaps between adjacent teeth (Fig. 14.102d)�����������������������������������������������������������������������������������������������������������������������������Pseudolambrus – Carapace epibranchial marginal teeth tubercular, triangular or spine-like, with or without gaps between adjacent teeth���������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 12 12. Carapace dorsal surface smooth, except for ridges on the gastric and branchial regions��������������������������������������������������13 – Carapace dorsal surface spinose, tuberculate or rugose����������������������������������������������������������������������������������������������������������21 13. Maxilliped 3 merus irregularly subtriangular (Fig. 14.103d)��������������������������������������������������������������������������������� Celatopesia – Maxilliped 3 merus rectangular (Fig. 14.103e)��������������������������������������������������������������������������������������������������������������������������14 14. Carapace lateral margin strongly expanded, posterolateral (meso-, metabranchial) margins convex, forming dome-shaped ventral depression, concealing pereopods 2–5 (Fig. 14.100f)����������������������������������Cryptopodia – Carapace lateral carapace not expanded, posterolateral margins concave, not forming dome-shaped ventral depression, pereopods 2–5 not or partially hidden�������������������������������������������������������������������������������������������������������������15 15. Carapace gastric region with distinct V-shaped ridge (Fig. 14.100h, 14.101d, f, 14.102f)����������������������������������������������������16 – Carapace gastric region without V-shaped ridge����������������������������������������������������������������������������������������������������������������������19 16. Carapace last epibranchial marginal tooth coincident with end of epibranchial ridge������������������������������������������������������17 – Carapace last epibranchial marginal tooth anterior and lateral to end of epibranchial ridge�������������������������������������������18 17. Epistome broad, lower margin straight. Carapace subbranchial region without large round tubercle between epibranchial margin and subbranchial ridge (Fig. 14.100h)�����������������������������������������������������������Distolambrus – Epistome narrow, lower margin V-shaped. Carapace subbranchial region with large round tubercle between epibranchial margin and subbranchial ridge (Fig. 14.101d)������������������������������������������������������������Latulambrus 18. Maxilliped 3 merus distalomesial corner margin entire, forming a blunt angle; palp hidden behind merus (Fig. 14.103f). Thoracic sternum smooth����������������������������������������������������������������������������������������������������������������Mesorhoea – Maxilliped 3 merus distalmesial margin with concave notch for palp; palp not hidden behind merus (Fig. 14.103h). Thoracic sternum tuberculate�������������������������������������������������������������������������������������������������Solenolambrus

654

19. – 20. – 21. – 22. – 23. – 24. –

Marine Decapod Crustacea

Carapace epibranchial ridge prominent, granular, ending posterior to last epibranchial tooth (Fig. 14.100l)������������������������������������������������������������������������������������������������������������������������������������������������������������Heterocrypta Carapace epibranchial ridge obsolete, ending in tooth at last epibranchial tooth�������������������������������������������������������������� 20 Carapace triangular, hepatic and epibranchial margins straight (Fig. 14.102j)����������������������������������������� Zarenkolambrus Carapace transversely ovate, hepatic and epibranchial margins convex (Fig. 14.100e)��������������������������������� Costalambrus Carapace ovate (Fig. 14.101a, k)��������������������������������������������������������������������������������������������������������������������������������������������������� 22 Carapace triangular to pyriform-subtriangular���������������������������������������������������������������������������������������������������������������������� 23 Carapace hepatic margin with last tooth almost touching first tooth of anterolateral margin, forming a circular posthepatic notch. Cheliped outer margin with closely spaced teeth (Fig. 14.101a). Female with large capitate tubercle on thoracic sternite 4������������������������������������������������������������������������������� Hypolambrus Carapace hepatic margin with last tooth not touching first tooth of anterolateral margin, not forming a circular posthepatic notch. Cheliped outer margin without closely spaced teeth (Fig. 14.101k). Female without large capitate tubercle on thoracic sternite 4������������������������������������������������������������������������������������������Parthenope Carapace posterior and metabranchial margins almost straight or evenly curved, smooth or with few tubercles at most (Fig. 14.100j, 14.102i)��������������������������������������������������������������������������������������������������������������������������������� 24 Carapace posterior and metabranchial margins produced posteriorly, forming strongly curved or sinuous margin, with or without teeth������������������������������������������������������������������������������������������������������������������������������������������������ 25 Cheliped merus outer margin proximal teeth fused, forming wing-like expansion (Fig. 14.100j). Mature female telson less than 1.5 times longer than pleonite 6 (Fig. 14.103o). Antennal article 4 about half length of article 3��������������������������������������������������������������������������������������������������������������������������������������������������������Furtipodia Cheliped merus outer margin proximal teeth not forming wing-like expansion (Fig. 14.102i). Mature female telson about twice as long as pleonite 6 (Fig. 14.103q). Antennal article 4 as long as article 3�������������������������������������������������������������������������������������������������������������������������������������������������������������������� Velolambrus

Fig. 14.98.  Parthenopidae. a, marginal features of typical daldorfiine; b, major regions of typical parthenopine. Left antennular article 1 (A), antenna (articles 1, 2 labelled a1, a2), ventral view: c, Daldorfinae; d, Parthenopinae, Lambrachaeus; e, Parthenopinae, Parthenope.

14 – Brachyura – crabs

655

Fig. 14.99.  Parthenopidae, Daldorfiinae. a, Daldorfia excavata (Baker, 1905); b, Niobafia erosa (Miers, 1879); c, Olenorfia cariei (Bouvier, 1914); d, Thyrolambrus verrucibrachium Zimmerman & Martin, 1999. Left antennular article 1 (A), antenna (articles 1, 2 labelled a1, a2), ventral view: e, Daldorfia; f, Niobafia; g, Olenorfia; h, Thyrolambrus. Female telson: i, Daldorfia; j, Niobafia; k, Olenorfia; l, Thyrolambrus.

25. Carapace posterior and metabranchial margins gently convex; hepatic, epibranchial and metabranchial margins with similar triangular teeth, none prolonged (Fig. 14.100d)������������������������������������������������������� Certolambrus – Carapace posterior and metabranchial margins produced; hepatic, epibranchial and metabranchial margins with or without teeth or spines������������������������������������������������������������������������������������������������������������������������������ 26 26. Carapace flat. Cheliped manus outer margin with large, flat, smooth, closely spaced teeth (Fig. 14.102c)Platylambrus – Carapace regions, especially epibranchial region inflated. Cheliped manus outer margin with well spaced, conical, often spinose or granulate teeth������������������������������������������������������������������������������������������������������������������������������ 27 27. Carapace lateral extremity rounded, without strong spine (Fig. 14.101i). Cheliped marginal teeth short, rounded, pearl-like (Fig. 14.101i). Male telson longer than wide. Gonopod 1 longer than gonopod 2, with thinner tip beyond lamelliform process (Fig. 14.103j)�������������������������������������������������������������������������������������Nodolambrus – Carapace lateral extremity with strong spine (Fig. 14.101b, j). Cheliped marginal teeth long, triangular. Male telson as wide or wider than long. Gonopod 1 shorter or longer than gonopod 2, evenly tapering, without lamelliform process (Fig. 14.103i)��������������������������������������������������������������������������������������������������������������������������� 28 28. Carapace gastric and branchial regions and margins with tall erect spines (Fig. 14.101b)������������������������ Hispidolambrus – Carapace gastric and branchial regions smooth or with low tubercles; margins with teeth or short spines����������������� 29 29. Carapace protogastric region with 2 median tubercles, both higher than mesogastric median tubercle (Fig. 14.101j)�������������������������������������������������������������������������������������������������������������������������������������������������������� Ochtholambrus – Carapace protogastric region with median tubercle(s), if any, lower than mesogastric median tubercle����������������������� 30 30. Carapace widest between last epibranchial teeth. Cheliped merus outer margin with proximal diagonal row of tubercles (Fig. 14.100k, 14.102h). Gonopod 1 shorter than gonopod 2 (Fig. 14.103l)�����������������������������������������31 – Carapace not widest between last epibranchial teeth. Cheliped merus outer margin without proximal diagonal row of tubercles. Gonopod 1 longer or about as long as gonopod 2�����������������������������������������������������������������32 31. Epistome with median transverse lamelliform protrusion. Suborbital region with diagonal lamelliform protrusion����������������������������������������������������������������������������������������������������������������������������������������Tutankhamen

656

Marine Decapod Crustacea

– 32. – 33. –

Epistome flat. Suborbital region without lamelliform protrusion����������������������������������������������������������������������Garthambrus Cheliped manus outer margin with 2 large teeth, with or without intermediate teeth (Fig. 14.101h, l)��������������������������33 Cheliped manus outer margin with more than 2 large teeth, with smaller intermediate teeth��������������������������������������� 34 Carapace hepatobranchial notch wide, V-shaped; epibranchial margin evenly rounded (Fig. 14.101h)������������ Neikolambrus Carapace hepatobranchial notch narrow; epibranchial margin asymmetrical, widest posteriorly (Fig. 14.101l)��������������������������������������������������������������������������������������������������������������������������������������������������������Parthenopoides 34. Carapace epibranchial margin slightly convex; lateral tooth and last epibranchial short, tips rounded, blunt. Pereopods 2–5 strongly compressed laterally (Fig. 14.100g)���������������������������������������������������������������� Derilambrus – Carapace epibranchial margin strongly convex; lateral tooth and last epibranchial long, tips blunt or sharp. Pereopods 2–5 subcylindrical to slightly compressed laterally�����������������������������������������������������������������������������������������35 35. Mature female telson lateral margins straight or concave (Fig. 14.103n)����������������������������������������������������������������������������� 36 – Mature female telson lateral margins convex (Fig. 14.103p)����������������������������������������������������������������������������������������������������37 36. Mature female telson lateral margins straight. Postorbital region with notch (Fig. 14.102e)����������������������� Rhinolambrus – Mature female telson lateral margins concave (Fig. 14.103n). Postorbital region without notch (Fig. 14.100i)��������������������������������������������������������������������������������������������������������������������������������������������������������Enoplolambrus 37. Carapace suborbital region concave; metagastric region not inflated (Fig. 14.102a)���������������������������������������Patulambrus – Carapace suborbital region not concave; metagastric region inflated (Fig. 14.102g)������������������������������������� Spinolambrus

Subfamily Daldorfiinae Ng & Rodríguez, 1986 Diagnosis. Antennal article 2 length as long or longer than antennular article 1 (Ng and Rodríiguez 1986; Tan and Ng 2007b). Daldorfia Rathbun, 1904 Diagnosis. Carapace subtriangular to subpentagonal. Antennal article 2 proximal half not enlarged, with anterolateral tooth (less than half length of article 3). Gonopod 2 longer than half length of gonopod 2. Female telson wider than long, semicircular. Cheliped manus about as long as merus; upper meral spine strong, overlapping epibranchial teeth. Maximum cl. 50 mm (Pl. 57d). Intertidal, subtidal (0–176 m; coral and rocky reef). Indo-West Pacific, Temperate Australia. 13 species (Ng and Boyko 2017; Ng et al. 2017; Tan and Ng 2007b).

Cheliped manus about as long as merus; upper meral spine strong, overlapping epibranchial teeth. Maximum cl. 22 mm (Pl. 57k). Subtidal (0–5 m; coral rubble). Western and Central IndoPacific. 1 species (Tan and Ng 2007b).

Thyrolambrus Rathbun, 1894 Diagnosis. Carapace subpentagonal, wider than long. Antennal article 2 proximal half not enlarged. Gonopod 2 longer than half length of gonopod 2. Female telson wider than long. Cheliped manus slender, about half length of merus; upper meral spine reduced to granule(s), not overlapping epibranchial teeth. Maximum cl. 21 mm (Pl. 58e). Subtidal–slope (1–500 m). Tropical W Atlantic, Western and Central Indo-Pacific, Tropical Eastern Pacific. 3 species (Alcock and Anderson 1895: figure; Tan and Ng 2007b: rediagnosis).

Niobafia Tan & Ng, 2007

Subfamily Parthenopinae MacLeay, 1838

Diagnosis. Carapace subtriangular, pitted. Antennal article 2 proximal half enlarged, about twice width of the distal half. Gonopod 2 about half length of gonopod 1. Female telson longer than wide. Cheliped manus lower margin dentate, teeth broad, low, irregular. Maximum cl. 19 mm. Intertidal, subtidal (0–5 m; coral reef). Indo-West Pacific. 1 species (Tan and Ng 2007b).

Diagnosis. Antennal article 2 less than half as long as antennular article 1 (Tan and Ng 2007a). Implicit generic attributes of Parthenopinae. Rostrum shorter than half carapace length. Gastric region without distinct V-shaped ridge. Suborbital region without diagonal ridge. Subbranchial region without large round tubercle between epibranchial margin and subbranchial ridge; pterygostomial region not excavated, without distinct afferent (inhalent) channel; pterygostomial ridge without dense setae. Epistome not protruding below antennular article 1. Eyes enclosed within deep orbits. Maxilliped 3 merus rectangular. Maxilliped 3 exopod exposed or partially hidden.

Olenorfia Tan & Ng, 2007 Diagnosis. Carapace transversely ovate. Antennal article 2 proximal half not enlarged, without anterolateral tooth. Gonopod 2 longer than half length of gonopod 2. Female telson wider than long, subtriangular, distolateral margins of tip constricted.

14 – Brachyura – crabs

657

Fig. 14.100.  Parthenopidae, Parthenopinae. a, Agolambrus agonus (Stimpson, 1871); b, Aulacolambrus diacanthus (De Haan, 1837); c, Celatopesia concava (Stimpson, 1871); d, Certolambrus pugilator (A. Milne-Edwards, 1873); e, Costalambrus tommasii (Rodrigues da Costa, 1959); f, Cryptopodia fornicata (Fabricius, 1787); g, Derilambrus angulifrons (Latreille, 1825); h, Distolambrus maltzami (Miers, 1881); i, Enoplolambrus carinatus (Herbst, 1796); j, Furtipodia gemma Tan & Ng, 2003; k, Garthambrus cidaris (Garth & Davie, 1995); l, Heterocrypta macrobrachia Stimpson, 1871.

Agolambrus Tan & Ng, 2007

Aulacolambrus Paulson, 1875

Diagnosis. Carapace angular-subcircular, slightly wider than long, regions inflated, sparsely tuberculate, pitted. Suborbital region with diagonal ridge. Epibranchial margin rounded. Cheliped margins dentate, 3 or 4 large teeth interspaced with smaller teeth, merus upper surface tuberculate. Maximum cl. 11 mm. Subtidal, shelf (7–201 m). Tropical W Atlantic. 1 species (Tan and Ng 2007a).

Diagnosis. Carapace rounded-triangular, slightly wider than long, regions inflated, tuberculate; pterygostomial region excavated, forming distinct afferent channel; pterygostomial ridge with dense, long setae covering afferent channel; epibranchial margin convex, last tooth much longer than more anterior teeth. Epistome protruding below antennular article 1. Cheliped outer margins with flat, long, blade-like teeth, sometimes alternating, merus upper margin tuberculate. Maximum cl. 25 mm (Pl. 57b).

658

Marine Decapod Crustacea

Fig. 14.101.  Parthenopidae, Parthenopinae. a, Hypolambrus hyponcus (Stimpson, 1871); b, Hispidolambrus mironovi (Zarenkov, 1990); c, Lambrachaeus ramifer Alcock, 1895 (with cheliped, pereopod 4, rostrum incomplete); d, Latulambrus occidentalis (Dana, 1854); e, Leiolambrus nitidus Rathbun, 1901; f, Mesorhoea bellii (A. Milne-Edwards, 1878); g, Mimilambrus wileyi Williams, 1979; h, Neikolambrus polemistes Tan & Ng, 2003; i, Nodolambrus nodosus (Jacquinot, 1853); j, Ochtholambrus triangularis (Stimpson, 1871); k, Parthenope longimanus (Linnaeus, 1758); l, Parthenopoides massena (Roux, 1830).

Intertidal–slope (0–290 m). Indo-West Pacific. 8 species (Tan 2004: key to species; Tan and Ng 2003: discussion of generic attributes).

Celatopesia Chiong & Ng, 1998 Diagnosis. Carapace pentagonal, wider than long, smooth except for ridges on gastric and branchial regions; epibranchial margin obtusely angled, slightly more produced than last epibranchial tooth, with fused rectangular teeth. Maxilliped 3 merus irregularly subtriangular. Cheliped margins with small, closely spaced

teeth, merus upper surface smooth. Mature female telson lateral margins concave. Maximum cl. 8 mm. Subtidal, shelf (7–73 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Chiong and Ng 1998: key to species).

Certolambrus Tan & Ng, 2003 Diagnosis. Carapace subtriangular, wider than long, regions inflated, tuberculate; epibranchial margin gently convex, angled at last tooth, with teeth triangular separated by triangular gaps; posterior and metabranchial margins produced posteriorly, gently

14 – Brachyura – crabs

659

Fig. 14.102.  Parthenopidae, Parthenopinae. a, Patulambrus nummiferus (Rathbun, 1906); b, Piloslambrus depressiusculus (Stimpson, 1871); c, Platylambrus serratus (H. Milne Edwards, 1834); d, Pseudolambrus calappoides (Adams & White, 1849); e, Rhinolambrus spinifer (Haswell, 1880); f, Solenolambrus portoricensis Rathbun, 1924; g, Spinolambrus meridionalis (Boschi, 1965); h, Tutankhamen cristatipes (A. MilneEdwards, 1880); i, Velolambrus tuberculatus (Flipse, 1930); j, Zarenkolambrus minutus McLay & Tan, 2009.

convex; hepatic, epibranchial and metabranchial margins with similar triangular teeth. Cheliped outer margins with triangular teeth, longer teeth interspaced with shorter teeth. Maximum cl. 7.4 mm. Intertidal–shelf (0–27 m). Central Indo-Pacific. 1 species (Tan and Ng 2003).

tooth, with rectangular teeth. Cheliped margins with regular, rectangular teeth not separated by gaps, merus upper surface smooth. Maximum cl. 10 mm. Subtidal (2–9 m). Tropical W Atlantic. 1 species (Bolaños et al. 2006: description of species, distribution; Tan and Ng 2007a: diagnosis, figure).

Costalambrus Tan & Ng, 2007

Cryptopodia H. Milne Edwards, 1834

Diagnosis. Carapace ovate, wider than long, smooth, except for mesogastric ridge and obsolete epibranchial ridge ending in tooth at last epibranchial tooth; epibranchial margin strongly convex, posteriorly obtusely angled, more produced than last epibranchial

buckler crabs Diagnosis. Carapace pentagonal to subtriangular, much wider than long, gastric, epibranchial regions inflated, smooth, tuberculate or

660

Marine Decapod Crustacea

Fig. 14.103.  Parthenopidae, Parthenopinae. Epistome, antennule (fossa shaded), antenna, suborbital region: a, Aulacolambrus; b, Certolambrus; c, Tutankhamen. Maxilliped 3: d, Celatopesia; e, Cryptopodia; f, Mesorhoea; g, Mimilambrus; h, Solenolambrus. Gonopods 1, 2: i, Hispidolambrus; j, Nodolambrus; k, Ochtholambrus; l, Tutankhamen. Male pleonite 6, telson: m, Ochtholambrus. Mature female pleonite 6, telson: n, Enoplolambrus; o, Furtipodia; p, Spinolambrus; q, Velolambrus.

pitted; lateral margin strongly expanded, posterolateral margins convex, forming dome-shaped ventral depression, concealing pereopods 2–5. Cheliped margins with fused rectangular teeth, merus upper surface smooth or tuberculate. Mature female telson lateral margins straight. Maximum cl. 41 mm (Pl. 57c). Subtidal–slope (7–500 m). Temperate Northern W Pacific, Indo-West Pacific, ?Temperate Australasia. 13 species (Chiong and Ng 1998: rediagnosis of genus and species, key to 12 species). The range of Cryptopodia angulata (H. Milne-Edwards & Lucas 1841) may be spreading in India due to translocation by shipping (Silambarasan et al. 2015; Sivasankar et al. 2019).

Derilambrus Tan & Ng, 2007 Diagnosis. Carapace subpyriform, slightly wider than long, inflated, tuberculate; hepatobranchial notch narrow; epibranchial margin asymmetrical, widest posteriorly; posterior and metabranchial margins produced posteriorly. Cheliped margins with short, triangular teeth, larger teeth interspaced with smaller teeth; merus upper surface tuberculate. Maximum cl. 28 mm. Subtidal, shelf (2–45 m). Temperate Northern Atlantic (including Mediterranean). 1 species (Tan and Ng 2007a; Zariquiey Alvarez 1968: description, figures of only species as Parthenope anguilifrons Latreille, 1825).

Distolambrus Tan & Ng, 2007 Diagnosis. Carapace subpentagonal, smooth, except for ridges on the gastric and branchial regions; gastric region with distinct

V-shaped ridge; last epibranchial tooth coincident with end of epibranchial ridge. Epistome broad, lower margin straight. Cheliped margins with short broadly circular teeth with denticulate edges, merus upper surface smooth. Maximum cl. 11 mm. Subtidal–slope (5–550 m). Temperate Northern E Atlantic (including Mediterranean). 1 species (Massi et al. 2010: distribution; Tan and Ng 2007a: diagnosis, figure).

Enoplolambrus A. Milne-Edwards, 1878 Diagnosis. Carapace subtriangular to subrhomboidal, wider than long, regions inflated, tuberculate; postorbital region without notch. Suborbital region concave. Epibranchial margin strongly convex. Lateral tooth and last epibranchial long; posterior and metabranchial margins produced posteriorly. Cheliped margins with long, triangular, alternating large and smaller teeth, merus upper surface tuberculate. Mature female telson lateral margins concave. Maximum cl. 61 mm (Pl. 57e). Intertidal–shelf (0–82 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Southern Africa. 7 species (Tan 2004: key to 6 species; Tan and Ng 2007a: rediagnosis of genus for Indo-West Pacific species previously assigned to Platylambrus).

Furtipodia Tan & Ng, 2003 Diagnosis. Carapace subtriangular, wider than long, gastric, hepatic and epibranchial regions strongly inflated, rugose; epibranchial margin anterior straight, posterior half strongly convex, with triangular spaced teeth; posterior and metabranchial

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margins produced posteriorly. Antennal article 4 about half length of article 3. Cheliped merus outer margin proximal teeth fused, forming wing-like expansion. Mature female telson slightly longer than pleonite 6. Maximum cl. 11 mm (Pl. 57f). Intertidal–shelf (0–62 m), shelf. Indo-West Pacific. 2 species (Tan and Ng 2003: diagnosis, 2 species described, figured).

Garthambrus Ng, 1996 Diagnosis. Carapace triangular, widest between last epibranchial teeth, regions inflated; epibranchial margin slightly convex, posterior seventh acutely angled, with narrow triangular teeth; posterior and metabranchial margins produced posteriorly. Cheliped margins with short, triangular teeth, merus upper surface smooth to tuberculate. Gonopod 1 shorter than gonopod 2. Maximum cl. 30 mm (Pl. 57g). Shelf, slope (50–765 m). Temperate Northern W Pacific, IndoWest Pacific. 11 species (McLay and Tan 2009: key to 11 species; Ng 1996; Ng and Tan 1999: key to 8 species).

Heterocrypta Stimpson, 1871 Diagnosis. Carapace subtriangular, smooth, except for granular gastric and epibranchial ridges; epibranchial margin with posterior quarter right-angled, more produced than last epibranchial tooth, with fused rectangular teeth. Cheliped margins with fused pentagonal teeth, merus upper surface smooth. Maximum cl. 18 mm (Pl. 57h). Intertidal–shelf (0–137 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 5 species (Chiong and Ng 1998: in key to similar genera; Tan 2004: key to 6 species).

Hispidolambrus McLay & Tan, 2009 Diagnosis. Carapace triangular, gastric and branchial regions and margins with tall erect spines, with acute hepatic tooth on shoulder; anterior epibranchial margin divided into 2 sections, anterior section distinctly arcuate, the posterior section straight, posterior margin with 5 or 6 spines; posterior and metabranchial margins produced posteriorly. Cheliped outer margins spinose. Male telson wider than long. Gonopod 1 shorter than gonopod 2, evenly tapering. Maximum cl. 32 mm. Shelf, slope (162–800 m). Eastern Indo-Pacific (Nazca and Sala y Gómez submarine ridges). 1 species (McLay and Tan 2009).

Hypolambrus Tan & Ng, 2007 Diagnosis. Carapace ovate; hepatic region with last tooth almost touching first tooth of anterolateral margin, forming a circular posthepatic notch; epibranchial margin convex, posterior one third region obtusely angled, more produced than last epibranchial tooth, with subtriangular to circular teeth; posterior and metabranchial margins almost straight or evenly curved, smooth or with few tubercles at most. Cheliped margins dentate, with short, triangular, alternating large and small teeth, merus upper surface tuberculate. Maximum cl. 29 mm. Subtidal–shelf (9–41 m). Tropical Eastern Pacific. 1 species (Tan and Ng 2007a: diagnosis, figure).

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Lambrachaeus Alcock, 1895 Diagnosis. Rostrum about half carapace length. Carapace pyriform, much longer than wide, gastrohepatic notch deep, wide, forming distinct neck; pterygostomial region excavated, forming distinct afferent channel; pterygostomial ridge with dense, long setae covering afferent channel. Eyes not enclosed by shallow orbits. Maximum cl. 13 mm (Pl. 57i). Subtidal, shelf (9–92 m). Temperate Northern W Pacific, IndoWest Pacific, Temperate Southern Africa. 1 species. The only species was mistaken for an inachid spider crab because of its exceptionally long ‘neck’ (Garth 1958: description, figures as Parthenope hyponcus Stimpson, 1871; Ng and McLay 2003: figures, rediagnosis).

Latulambrus Tan & Ng, 2007 Diagnosis. Carapace widely triangular-pentagonal, smooth, except for ridges on the gastric and branchial regions; gastric region with distinct V-shaped ridge; subbranchial region with large round tubercle between epibranchial margin and subbranchial ridge; last epibranchial tooth coincident with end of epibranchial ridge. Epistome narrow, lower margin V-shaped. Cheliped margins with short, triangular teeth, merus upper surface smooth. Maximum cl. 21 mm. Intertidal–shelf (0–174 m). Tropical Eastern Pacific. 1 species (Garth 1958; Tan and Ng 2007a: diagnosis).

Leiolambrus A. Milne-Edwards, 1878 Diagnosis. Carapace rhomboidal, with 3 low, parallel ridges, one on each epibranchial region and one on cardiac region; epibranchial margin slightly convex anteriorly, multidentate, strong spine at midpoint, posterior half concave, with tooth slightly above margin. Maximum cl. 12 mm. Subtidal, shelf (7–185 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Gore and Scotto 1979: 1 species; Tan 2004: key to species).

Mesorhoea Stimpson, 1871 Diagnosis. Carapace pentagonal, smooth except for mesogastric and epibranchial ridges; gastric region with distinct V-shaped ridge; epibranchial margin convex, last tooth anterior and lateral to end of epibranchial ridge. Thoracic sternum smooth. Maxilliped 3 merus distomesial margin entire, forming blunt angle; palp hidden behind merus. Cheliped margins with short, pentagonal contiguous teeth, merus upper surface smooth. Maximum cl. 15 mm. Subtidal, shelf (2–100 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Garth 1958; Gore and Scotto 1979; Tan 2004).

Mimilambrus Williams, 1979 Diagnosis. Carapace subcircular, regions inflated, tuberculate; pterygostomial region excavated, forming distinct afferent channel;

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pterygostomial ridge with dense, long setae covering afferent channel; epibranchial margin strongly convex, posterior third obtusely angled, with triangular separated teeth. Maxilliped 3 exopod completely hidden by maxilliped. Maximum cl. 26 mm. Subtidal, shelf (3–31 m). Tropical W Atlantic. 1 species (Ng and Rodríiguez 1986; Williams 1979: figures).

Neikolambrus Tan & Ng, 2003 Diagnosis. Carapace triangular-pyriform, as wide as long, regions inflated, tuberculate; hepatobranchial notch wide, V-shaped; epibranchial margin evenly rounded; posterior and metabranchial margins produced posteriorly. Cheliped manus outer margin with 2 large teeth, with or without intermediate teeth. Maximum cl. 10 mm (Pl. 57j). Shelf (27–50 m). Central Indo-Pacific. 1 species (Tan and Ng 2003).

Nodolambrus Tan & Ng, 2007 Diagnosis. Carapace subpyriform, slightly wider than long, regions inflated, tuberculate; epibranchial margin strongly convex, irregularly nodular; posterior and metabranchial margins produced posteriorly. Cheliped marginal teeth short, rounded, pearl-like. Male telson longer than wide. Gonopod 1 longer than gonopod 2, with thinner tip beyond lamelliform process. Maximum cl. 28 mm. Subtidal, shelf (9–22 m). Central Indo-Pacific (N Australia). 1 species (Campbell and Stephenson 1970: figure; Tan and Ng 2007a: diagnosis).

Ochtholambrus Tan & Ng, 2007 Diagnosis. Carapace triangular, protogastric region with 2 median tubercles, both higher than mesogastric median tubercle; epibranchial margin convex, angled at last epibranchial tooth, with triangular, closely spaced teeth; posterior and metabranchial margins produced posteriorly. Cheliped manus outer margin with 2 or 3 triangular, widely spaced teeth, merus upper margin with oblique row of tubercles or cristae. Maximum cl. 32 mm. Subtidal, shelf (4–128 m). Tropical W Atlantic, Tropical Eastern Pacific. 3 species (Garth 1958; Milne-Edwards 1868: description, figure of Atlantic species; Tan and Ng 2007a: diagnosis).

Parthenope Weber, 1795 Diagnosis. Carapace ovate, tuberculate; hepatic region with last tooth not touching first tooth of anterolateral margin, not forming a circular posthepatic notch; epibranchial margin rounded; posterior and metabranchial margins almost straight or evenly curved, smooth or with few tubercles at most. Cheliped margins with alternating large and smaller teeth; merus upper surface tuberculate. Maximum cl. 30 mm (Pl. 57l). Intertidal–slope (0–300 m). Western and Central Indo-Pacific. 3 species Parthenope was previously home to many species but restricted to just three by Tan (2004) who provided a key. Other than including Parthenope in their key to genera the genus was

not rediagnosed by Tan and Ng (2007a) when they moved many species to other genera.

Parthenopoides Miers, 1879 Diagnosis. Carapace subtriangular, wider than long, regions inflated, tuberculate; hepatobranchial notch narrow; epibranchial margin asymmetrical, widest posteriorly; posterior and metabranchial margins produced posteriorly. Cheliped manus outer margin with 2 large teeth, with or without intermediate teeth. Maximum cl. 16 mm. Subtidal–slope (3–445 m). Temperate Northern E Atlantic (including Mediterranean). 1 species (Tan 2004: rediagnosis; Zariquiey Alvarez 1968: description of only species as Parthenope massena (Roux, 1830)).

Patulambrus Tan & Ng, 2007 Diagnosis. Carapace subtriangular, wider than long, regions inflated, smooth to tuberculate. Suborbital region concave. Epibranchial margin strongly convex. Lateral tooth and last epibranchial long; posterior and metabranchial margins produced posteriorly. Cheliped merus outer margin dentate, 3 or 4 teeth larger than interspersed teeth; merus upper surface smooth, finely pitted. Mature female telson lateral margins convex. Maximum cl. 31 mm. Shelf, slope (50–500 m). Temperate Northern W Pacific, IndoWest Pacific. 2 species (Tan and Ng 2007a).

Piloslambrus Tan & Ng, 2007 Diagnosis. Carapace subellipitical to triangular, wider than long, regions inflated, tuberculate; last epibranchial tooth slightly longer than more anterior teeth. Epistome with protruding tubercle below antennular article 1. Cheliped manus outer margin with flat teeth, longer teeth interspaced with smaller, teeth, merus upper margin tuberculate. Maximum cl. 21 mm. Subtidal, shelf (3–55 m). Tropical W Atlantic, Tropical Eastern Pacific. 3 species (Tan and Ng 2007a).

Platylambrus Stimpson, 1871 Diagnosis. Carapace subovate, wider than long, flat, tuberculate; epibranchial margin convex, posterior third acutely angled, more produced than last epibranchial tooth, with triangular teeth; posterior and metabranchial margins produced posteriorly. Cheliped manus outer margin with large, flat, smooth, closely spaced teeth. Maximum cl. 23 mm (Pl. 58a). Subtidal–slope (2–824 m). Tropical W Atlantic. 2 species (Gore 1977: 2 species of Parthenope (Platylambrus); Tan 2004).

Pseudolambrus Paulson, 1875 Diagnosis. Carapace semi-elliptical, pentagonal or triangular, usually wider than long, gastric and cardiac regions inflated, tuberculate; epibranchial teeth subrectangular, long, close or with

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narrow gaps between adjacent teeth. Cheliped manus outer margin indistinct, sometimes with 2 or 3 teeth. Maximum cl. 30 mm. Intertidal–slope (0–500 m). Temperate Northern W Pacific, Indo-West Pacific, Temperate Australasia. 19 species (Takeda and Komatsu 2015: new species; Tan 2004: rediagnosis, four species groups, keys to species; Tan 2008). Parthenolambrus A. MilneEdwards, 1878 is treated as a junior synonym.

Rhinolambrus A. Milne-Edwards, 1878 Diagnosis. Carapace pyriform, regions inflated, tuberculate; postorbital region with notch; hepatobranchial notch wide, V-shaped; epibranchial margin strongly convex, dentate. Lateral tooth and last epibranchial long; posterior and metabranchial margins produced posteriorly. Cheliped manus outer margin variably dentate, with smaller intermediate teeth. Mature female telson lateral margins straight. Maximum cl. 56 mm (Pl. 58b). Intertidal–slope (0–500 m). Temperate Northern W Pacific, Indo-West Pacific, Temperate Australasia. 13 species (Tan 2004: key to species; Tan et al. 1999: figures of 2 species).

Solenolambrus Stimpson, 1871 Diagnosis. Carapace pentagonal, wider than long, smooth, except for mesogastric ridge and prominent epibranchial ridge ending posterior to last epibranchial tooth; gastric region with distinct V-shaped ridge; last epibranchial tooth anterior and lateral to end of epibranchial ridge. Maxilliped 3 merus distalmesial margin with concave notch for palp; palp not hidden behind merus. Cheliped margins with short triangular spaced teeth, merus upper surface smooth or tuberculate. Maximum cl. 11.5 mm (Pl. 58c). Subtidal–slope (5–618 m). Tropical Atlantic, Tropical Eastern Pacific. 6 species (Gore and Scotto 1979: 3 species described; Tan 2004: key to 6 species).

Spinolambrus Tan & Ng, 2007 Diagnosis. Carapace triangular, metagastric region inflated. Suborbital region convex. Epibranchial margin strongly convex. Lateral tooth and last epibranchial long; posterior and metabranchial margins produced posteriorly. Cheliped manus outer margins spinose, with long alternating teeth, merus upper surface spinose to tuberculate. Mature female telson lateral margins convex. Maximum cl. 40 mm (Pl. 58d). Subtidal–slope (7–750 m). Temperate Northern and Tropical Atlantic (including Mediterranean), Tropical Eastern Pacific. 8 species (Tan 2004: key to species; Tan and Ng 2007a: diagnosis).

Tutankhamen Rathbun, 1925 Diagnosis. Carapace triangular, widest between last epibranchial teeth, regions inflated, smooth. Suborbital region with diagonal lamelliform process. Epibranchial margin slightly convex, posterior seventh acutely angled, with narrow triangular teeth; posterior and metabranchial margins produced posteriorly. Epistome with median transverse lamelliform protrusion. Cheliped margins dentate to cristate, merus with broadly triangular teeth over

663

proximal half only, merus upper surface smooth. Gonopod 1 shorter than gonopod 2. Maximum cl. 14 mm. Slope (227–366 m). Tropical Atlantic (Caribbean). 1 species (McLay and Tan 2009: rediagnosis, figures).

Velolambrus Tan & Ng, 2007 Diagnosis. Carapace triangular, wider than long, regions slightly inflated-flat, smooth to slightly tuberculate; epibranchial margin slightly convex, with triangular teeth separated by triangular gaps; posterior and metabranchial margins produced posteriorly. Antennal article 4 about as long as article 3. Cheliped manus outer margin with 2 or 3 lobate or triangular teeth; merus upper surface without row of tubercles. Mature female telson about twice as long as pleonite 6. Maximum cl. 7 mm. Subtidal, shelf (13–125 m). Temperate Northern Atlantic and Tropical (including Mediterranean), Central Indo-Pacific. 2 species (Flipse 1930; figures: Miers 1879; Tan 2004; Tan and Ng 2007a: diagnosis).

Zarenkolambrus McLay & Tan, 2009 Diagnosis. Carapace triangular, slightly wider than long, smooth, except for mesogastric ridge and obsolete epibranchial ridge ending in tooth at last epibranchial tooth. Maximum cl. 5 mm. Slope (400 m). Eastern Indo-Pacific. 2 species (McLay and Tan 2009: diagnosis, key to species). References Alcock A, Anderson ARS (1895) Illustrations of the Zoology of the Royal Indian Marine Surveying Steamer Investigator, under the Command of Commander A. Carpenter, R.N., D.S.O., of the Late Commander R.F. Hoskyn, R.N., and of Commander C.F. Oldham. Crustacea, Part III. Pls IX–XV. Trustees of the Indian Museum, Calcutta. Anker A, Ivanov Y (2020) First record of the predation upon sea slugs (Cephalaspidea and Nudibranchia) by the peculiar elbow crab Lambrachaeus ramifer Alcock, 1895 (Decapoda: Parthenopidae). Marine Biodiversity 50, doi:10.1007/s12526-020-01047-x. Bolaños J, Lira C, Hernández G, López R (2006) El cangrejo Heterocrypta tommasii (Decapoda: Parthenopidae): adición a la carcinofauna venezolana. Revista de Biología Tropical 54, 131–134. Campbell BM, Stephenson W (1970) The sublittoral Brachyura (Crustacea: Decapoda) of Moreton Bay. Memoirs of the Queensland Museum 15, 235–301, pl. 22. Chiong WL, Ng PKL (1998) A revision of the buckler crabs of the genus Cryptopodia H. Milne Edwards, 1834 (Crustacea: Decapoda: Brachyura: Parthenopidae). Raffles Bulletin of Zoology 46, 157–216. Flipse HJ (1930) Decapoda F. (Decapoda Brachyura continued) Oxyrhyncha: Parthenopidae. Siboga-Expéditie 39c, 1–96. Garth JS (1958) Brachyura of the Pacific coast of America. Oxyrhyncha. Allan Hancock Pacific Expeditions 21, 1–854. Gore RH (1977) Studies on decapod Crustacea from the Indian River region of Florida. VI. The identity of Parthenope (Platylambrus) serrata (H. Milne Edwards, 1834) and Parthenope (Platylambrus) granulata (Kingsley, 1879). Proceedings of the Biological Society of Washington 90, 505–531. Gore RH, Scotto LE (1979) Crabs of the family Parthenopidae (Crustacea Brachyura: Oxyrhyncha) with notes on specimens from the Indian River region of Florida. Memoirs of the Hourglass Cruises 3, 1–98.

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Massi D, Micalizzi R, Giusto GB, Pipitone C (2010) First record of Heterocrypta maltzami Miers, 1881 (Decapoda, Brachyura, Parthenopidae) in the Strait of Sicily. Crustaceana 83, 1141–1145. doi:10.1163/001121610X520966 McLay C, Tan SH (2009) Revision of the crab genus Garthambrus Ng, 1996, with the description of two new genera and discussion of the status of Tutankhamen Rathbun, 1925 (Crustacea: Brachyura: Parthenopidae). Zootaxa 2122, 1–50. doi:10.11646/zootaxa.2122.1.1 Miers EJ (1879) Descriptions of new or little-known species of maioid Crustacea (Oxyrhyncha) in the collection of the British Museum. Annals and Magazine of Natural History (ser. 5) 4, 1–28, pls 4, 5. Milne-Edwards A (1868) Observations sur la faune carcinologiques des îles du Cap-Vert. Nouvelles Archives du Muséum d’Histoire Naturelle, Paris 4, 49–68. doi:10.5962/bhl.part.15909 Mura M, Orrù F (1998) Feeding habits of Parthenope macrochelos (Brachyura, Parthenopidae) in the central-western Mediterranean Sea. Biologia Marina Mediterranea 5, 570–573. Ng PKL (1996) Garthambrus, a new genus of deep water parthenopid crabs (Crustacea: Decapoda: Brachyura) from the Indo-Pacific, with description of a new species from the Seychelles. Zoölogische Mededeelingen 70, 155–168. Ng PKL, Boyko CB (2017) New species and records of crabs of the families Dromiidae, Dynomenidae, Homolidae, Aethridae, Parthenopidae, Majidae, and Epialtidae (Crustacea: Decapoda: Brachyura) from Easter Island, with a review of the majid genus Schizophroida Sakai, 1933. Pacific Science 71, 197–227. doi:10.2984/71.2.8 Ng PKL, McLay CL (2003) On the systematic position of Lambrachaeus Alcock, 1895 (Brachyura, Parthenopidae). Crustaceana 76, 897–915. doi:10.1163/156854003771997819 Ng PKL, Rodríiguez G (1986) New records of Mimilambrus wileyi Williams, 1979 (Crustacea: Decapoda: Brachyura), with notes on the systematics of the Mimilambridae Williams, 1979 and Parthenopoidea MacLeay, 1838 sensu Guinot, 1978. Proceedings of the Biological Society of Washington 99, 88–99. Ng PKL, Tan SH (1999) The Hawaiian parthenopid crabs of the genera Garthambrus Ng, 1996, and Dairoides Stebbing, 1920 (Crustacea: Decapoda: Brachyura). Proceedings of the Biological Society of Washington 112, 120–132. Ng PKL, Prema M, Tan SH, Ravichandran S (2017) The taxonomy of two poorly known species of elbow crabs, Daldorfia spinosissima (A. MilneEdwards, 1862) and D. triangularis Sakai, 1974 (Brachyura, Parthenopidae). Crustaceana 90, 1779–1791. doi:10.1163/15685403-00003721 Rathbun MJ (1925) The spider crabs of America. Bulletin of the United States National Museum 129, 1–613. Silambarasan K, Sundaramanickam A, Sujatha K, Senthilkumaar P (2015) First record of buckler crab Cryptopodia angulata (Decapoda: Brachyura: Parthenopidae) from Chennai coast (Bay of Bengal), India. Journal of Asia-Pacific Biodiversity 8, 102–104. doi:10.1016/j.japb.2015.02.002 Sivasankar R, Ezhilarasan P, Kanuri VV, Kumaraswami M, Rao VR, et  al. (2019) On the occurrence of buckler crab Cryptopodia angulata in the coastal waters of India. Indian Journal of Geo-Marine Sciences 48, 464–467. Takeda M, Komatsu H (2015) A new species of the genus Pseudolambrus (Crustacea, Decapoda, Brachyura, Parthenopidae) from the Ogasawara Islands, Japan. Bulletin of the National Museum of Nature and Science. Series A, Zoology 41, 15–20.

Tan SH (2004) A systematic revision of the Parthenopidae (Crustacea: Decapoda: Brachyura). PhD thesis. National University of Singapore, Singapore. Tan SH (2008) A new species of Pseudolambrus (Crustacea: Brachyura: Parthenopidae: Parthenopinae) from Panglao, Bohol, the Philippines. Zootaxa 1731, 63–68. doi:10.11646/zootaxa.1731.1.6 Tan SH, Ng PKL (2003) The Parthenopinae of Guam (Crustacea: Decapoda: Brachyura: Parthenopidae). Micronesica 35–36, 385–416. Tan SH, Ng PKL (2007a) Descriptions of new genera from the subfamily Parthenopinae (Crustacea: Decapoda: Brachyura: Parthenopidae). Raffles Bulletin of Zoology 16, 95–119. Tan SH, Ng PKL (2007b) Review of the subfamily Daldorfiinae Ng & Rodriguez, 1986 (Crustacea: Decapoda: Brachyura: Parthenopidae). Raffles Bulletin of Zoology Supplement 16, 121–167. Tan SH, Huang J-F, Ng PKL (1999) Crabs of the Family Parthenopidae (Crustacea: Decapoda: Brachyura) from Taiwan. Zoological Studies 38, 196–206. Williams A (1979) A new crab family from the shallow waters of the West Indies (Crustacea: Decapoda: Bachyura). Proceedings of the Biological Society of Washington 92, 399–413. Zariquiey Alvarez R (1968) Crustáceos decápodos ibéricos. Investigaciones Pesqueras 32, 1–510. Zipser E, Vermeij GJ (1978) Crushing behavior of tropical and temperate crabs. Journal of Experimental Marine Biology and Ecology 31, 155–172. doi:10.1016/0022-0981(78)90127-2

Pilumnoidea Samouelle, 1819 The pilumnoids currently comprise three families united by the usually sinuous gonopod 1 and very short, sigmoid gonopod 2 that is typically about one-quarter the length of gonopod 2 or less, and which has a slightly ‘cupped’ apex (Davie et al. 2015; Ng et al. 2008). Tanaochelids are easy to recognise by the elongated chelipeds with distally spoontipped fingers, but distinctions between Pilumnidae and Galenidae are largely arbitrary. No single feature or combination of features distinguishes pilumnids from galenids and the two groups are unlikely to be reciprocally monophyletic. Diagnosis. Carapace variable in shape, from broadly pentagonal, subpentagonal to hexagonal or subquadrilateral, usually convex, sometimes setose. Antennule folding transversely, or folding obliquely forwards. Buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular. Male pleonites 1–6 and telson free. Gonopod 1 slender, usually sinuous, distal part without large spines or complex folds, apex usually recurved. Gonopod 2 minute, less than one-fourth (rarely one-third) as long as gonopod 1, sigmoidal, comma-shaped, apex slightly cupped.

Key to families of Pilumnoidea 1. –

At least one cheliped long and slender, twice carapace length or more; fingers spoon-tipped (Fig. 14.114)�����������������������������������������������������������������������������������������������������������������������������������������������������������Tanaochelidae Chelipeds about as long as or little longer than carapace; finger tips pointed, not spoon-tipped���������������������������������������2

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2. –

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Carapace variously shaped, widest anteriorly or posteriorly; surface glabrous or setose, usually densely setose (Figs 14.105–14.113). Endostomial ridges usually present; if carapace glabrous or sparsely setose, endostomial ridges present�������������������������������������������������������������������������������������������������������������������������������������Pilumnidae Carapace transversely ovate to subhexagonal, always widest near midlength; surface glabrous or sparsely setose, never densely setose (Fig. 14.104). Endostomial ridges absent����������������������������������������������������������������Galenidae

References Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology - Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286.

Galenidae Alcock, 1898 Figure 14.104, Plate 59a, b The galenids are a group of ‘atypical’ pilumnoids that superficially resemble goneplacids or xanthids. Unlike the

‘hairy’ pilumnids, galenids are sparsely setose at most, but nevertheless have typically pilumnoid gonopods and larvae (where known) (Lee and Ko 2016; Mohan and Kannupandi 1986; Terada 1985). Galenidae, however, is probably an artificial group and further research will probably show some of the constituent genera to be more closely allied to various pilumnids than to each other. Apart from one species that is captured in sufficient numbers to be of some interest to fisheries, others are rarely collected. They are mostly from muddy subtidal sediments. Diagnosis. Carapace transversely ovate, widest near midlength, glabrous or sparely setose (never densely setose). Endostomial ridges absent. Chelipeds about as long as or little longer than carapace; finger tips pointed, not spoon-tipped.

Key to genera of Galenidae 1. – 2. – 3.

Carapace anterolateral margin with single laterally directed tooth, otherwise evenly curved or weakly scalloped (Fig. 14.104a). Cheliped palm upper margin with thin, blade-like crest (Fig. 14.104e)�����������Dentoxanthus Carapace anterolateral margin with 2 or more anterolateral teeth. Cheliped palm upper margin tuberculate or bluntly rounded, without thin, blade-like crest�����������������������������������������������������������������������������������������������������������������2 Carapace surface smooth, regions weakly indicated; with 2 or 3 small, widely-spaced anterolateral teeth. Cheliped palm upper surface smooth or with spaced, low tubercles (Fig. 14.104b, Pl. 59a)�������������������������������� Galene Carapace surface rugose, regions distinctly indicated; with 4 or more large, closely-spaced anterolateral teeth (Fig. 14.104c, d). Cheliped palm upper surface prominently nodular or tubercular (Fig. 14.104f, g)������������������3 Carapace anterolateral margin with 4 lobes or narrow triangular teeth, third and fourth largest, separated by deep angles (Fig. 14.104c). Cheliped palm upper margin with 4 or more large tubercles; outer surface covered with flat or rounded tubercles or smooth; dactylus upper proximal margin with tubercles, usually prominent (Fig. 14.104f, Pl. 59b)������������������������������������������������������������������������������������������������������������������Halimede

Fig. 14.104.  Galenidae. a, Dentoxanthus iranicus Stephensen, 1946. Carapace, pereopods: b, Galene; c, Halimede; d, Parapanope. Cheliped, outer face: e, Dentoxanthus; f, Halimede; g, Parapanope. Male pleon: h, Dentoxanthus.

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Marine Decapod Crustacea

Carapace anterolateral margin with 4 broad triangular teeth or lobes separated by shallow angles (Fig. 14.104d). Cheliped palm upper margin bluntly rounded or with spaced low tubercles; outer surface with 2 or 3 longitudinally granulated carinae or smooth; dactylus upper proximal margin smooth, without tubercles (Fig. 14.104g)�����������������������������������������������������������������������������������������������������������������������������Parapanope Dentoxanthus Stephensen, 1946

Diagnosis. Carapace regions well indicated; anterolateral margin with single laterally directed tooth, otherwise evenly curved or weakly sinuous. Cheliped palm upper margin with smooth, thin, blade-like crest; outer surface smooth. Maximum cl. 18 mm. Subtidal, shelf (10–25 m). Western Indo-Pacific. 1 species (Ng et al. 2015: redescription).

Galene De Haan, 1833 Diagnosis. Carapace smooth, regions weakly indicated; anterolateral margin with numerous small granules, 2 or 3 distinct conical teeth (2 teeth distinct). Cheliped palm upper margin bluntly rounded or with spaced low tubercles; outer surface tuberculate. Maximum cl. 60 mm (Pl. 59a). Subtidal, shelf (1–100 m). Western and Central Indo-Pacific. 1 species (Ng 1998: diagnosis, figure). The only species, of low commercial value, can be abundant and is caught as trawl by-catch.

Halimede De Haan, 1835 Diagnosis. Carapace rugose, regions distinctly indicated; anterolateral margin with 4 lobes or narrow triangular teeth, third and fourth largest, separated by deep angles. Cheliped palm upper margin with 4 or more large tubercles; outer surface covered with flat or rounded tubercles, or smooth; dactylus upper proximal margin with tubercles, usually prominent. Maximum cl. 33 mm (Pl. 59b). Intertidal–shelf (0–50 m). Temperate Northern W Pacific, Western and Indo-Pacific; introduced to E Mediterranean. 4 species (Galil 2000: rediagnosis, key to 3 species).

Parapanope De Man, 1895 Diagnosis. Carapace rugose, regions distinctly indicated; anterolateral margin with 4 broad triangular teeth or lobes separated by shallow angles. Cheliped palm upper margin bluntly rounded or with spaced low tubercles; outer surface with 2 or 3 longitudinally granulated carinae, or smooth; dactylus upper proximal margin smooth, without tubercles. Maximum cl. 15 mm. Intertidal–shelf (0–35 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 8 species (Ng and Guinot 2021: rediagnosis, key to species). References Galil BS (2000) Halimede tyche (Herbst, 1801) (Decapoda; Xanthoidea; Halimedinae) from the Mediterranean Coast of Israel, with a review of the genus. Journal of Crustacean Biology 20, 325–331. doi:10.1163/​ 1937240X-90000034 Lee SH, Ko HS (2016) The first zoeal stages of Parapanope euagora and Halimede fragifer (Decapoda: Pilumnoidea: Galenidae) hatched in the

laboratory. Animal Systematics, Evolution and Diversity 32, 133–140. doi:10.5635/ASED.2016.32.2.133 Mohan R, Kannupandi T (1986) Complete larval development of the xanthid crab, Galene bispinosa (Herbst), reared in the laboratory. In Biology of benthic organisms. Techniques and methods as applied to the Indian Ocean. (Eds Thompson MF, Sarojini R, Nagabhushanam R) pp. 193–202. Oxford & IBH Publishing Co., New Delhi. Ng PKL (1998) Crabs. In FAO Species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 2. Cephalopods, crustaceans, holothurians and sharks. (Eds Carpenter KE, Niem VH) pp. 1045–1155. Food and Agriculture Organization, Rome. Ng PKL, Guinot D (2021) Parapanope De Man, 1895 (Decapoda: Brachyura: Pilumnoidea: Galenidae): revisited and revised, with descriptions of two new species. Journal of Crustacean Biology 41, doi:10.1093/ jcbiol/ruab020. Ng PKL, Trivedi JN, Vachhrajani KD (2015) Dentoxanthus iranicus Stephensen, 1946 (Crustacea: Brachyura: Galenidae): a new record from Gujarat, India, with systematic notes. Marine Biodiversity Records 8, e132. doi:10.1017/S1755267215001098 Terada M (1985) Early zoeal development of Halimede fragifer De Haan (Xanthidae, Xanthinae). Proceedings of the Japanese Society of Systematic Zoology 31, 30–37.

Pilumnidae Samouelle, 1819 Figures 14.105–14.113, Plates 59i, j, 60 The pilumnids, commonly known as hairy crabs, are most common in shallow waters on all types of substrate. They are particularly common in rock and coral interstices, and amongst sponge, algal and bryozoan encrustations, but also occur in muddy and estuarine habitats, or as echinoderm symbionts. Unlike galenids, pilumnids are usually distinctly setose dorsally, and usually have endostomial ridges (always absent in galenids). Given the morphological and ecological diversity of Pilumnidae, their classification has been fluid. Davie (2002) and Ng et  al. (2008) reviewed the numerous subfamilial schemes but the most recent synopsis recognised five (Davie et  al. 2015): Calmaniinae, Eumedoninae, Pilumnidae, Rhizopinae and Xenophthalmodinae. Of these we recognise the first four subfamilies. The elongated gonopods 1 that extend anteriorly beyond the pleon, believed to be diagnostic of the monogeneric Xenophthalmodinae (Števčić 2005), is a feature of only some members of Xenophthalmodes. As such, we are unable to distinguish Rhizopinae from Xenophthalmodinae and consider the two to be synonymous. However, we recognise the validity of the monogeneric Hapalonotinae as separate from Eumedoninae given the highly divergent morphology of Hapalonotus (Chia and Ng 1999; Števčić 2005).

14 – Brachyura – crabs

Pilumninae and Rhizopinae are the two largest pilumnid subfamilies but their separation is currently difficult. Ng (1987) showed the rhizopines to belong to Pilumnidae rather than Goneplacidae, but since then, Rhizopinae has neither been adequately diagnosed nor shown to be a natural group (Ng et  al. 2008). Pilumnines generally have a transversely ovate carapace with prominent anterolateral teeth, and a proximally narrow pleon in which sternite 8 is usually fully concealed by pleonite 2; they are usually most common on rocky and coral reefs. Rhizopines generally have a more quadrate carapace with obsolescent anterolateral teeth and a proximally widened pleon, albeit with male pleonite 2 narrowed such that sternite 8 is visible on either side; they are more common in muddy and estuarine habitats. These distinctions, however, are not consistent and the current subfamilial placement of several genera is largely arbitrary. For instance, the pilumnine Vellumnus, with prominent anterolateral carapace teeth appears to be closely related to the rhizopine, Lophoplax (also with prominent anterolateral teeth); and the rhizopine, Itampolus, closely resembles the pilumnine, Glabropilumnus. Also, some genera have been variously assigned to both, such as Heteropilumnus, Peleianus and Viaderiana (see Ng 1987; Ng et  al. 2008; Serène and

667

Umali 1972). As such, pending phylogenetic evaluation, Pilumninae and Rhizopinae are currently best considered as heuristic groupings that probably reflect a degree of phylogenetic and ecological reality, but which require refinement to be taxonomically effective. To this effect, we employ Pilumninae and Rhizopinae substantially following the taxonomic composition of Ng et  al. (2008) but with several pragmatically made generic realignments given under the accounts of the respective subfamilies. Following Ng et al. (2008), Heteropilumnus is included in Pilumninae but owing to morphological heterogeneity, appears twice in the key below, once among the pilumnines and once among the rhizopines. Identification of pilumnids, especially rhizopines and pilumnines, often relies on carapace and pereopod setation patterns as well as details of surface ornamentation. As such, setae may need to be removed from parts of a specimen in order to observe diagnostic surface features. Diagnosis. Carapace transversely ovate; usually densely setose. Chelipeds about as long as or little longer than carapace; finger tips pointed, not spoon-tipped. Endostomial ridges usually present; if carapace glabrous or sparsely setose, endostomial ridges present. Male pleon triangular, pleonites 3–6 with margins tapering.

Key to subfamilies and genera of Pilumnidae 1. – 2. – 3. – 4. – 5. – 6. – 7. –

Antennules folding obliquely forwards (Fig. 14.106m, n)����������������������������������������������������������������������������������������������������������2 Antennules folding transversely (Fig. 14.107b, 14.109l–n)�������������������������������������������������������������������������������������������������������14 Carapace front at most weakly produced beyond orbits, with fringe of long setae (Fig. 14.105a). Cheliped fingers with proximal gape when closed, margins of gape with dense brush of bristly setae (Fig. 14.105b). Pereopods 2–5 without dactylus-propodal lock�����������������������������������������������������������������������Calmaniinae … Calmania Carapace front strongly produced beyond orbits, often as single rostrum or pair of ‘horns’, without fringe of setae. Cheliped fingers without gape, occlusal margins at most sparsely setose. Pereopods 2–5 with dactylus-propodal lock (Fig. 14.106r, u)��������������������������������������������������������������Eumedoninae … 3 Carapace rounded, ovate-pyriform (Fig. 14.106h)������������������������������������������������������������������������������������������������ Rhabdonotus Carapace angular, subpentagonal-subhexagonal������������������������������������������������������������������������������������������������������������������������4 Carapace surface with pair of long upright protogastric spines (Fig. 14.106j)���������������������������������������������������� Tiaramedon Carapace surface smooth or variously ornamented, without long paired protogastric spines��������������������������������������������5 Carapace anterolateral margin with several incisions; frontal lobes separated by median incision, flanked on each side by blunt or pointed supraorbital lobe of varying length (Fig. 14.106a, f, g, i)���������������������������������������������6 Carapace anterolateral margin entire; front produced as long median rostrum (with or without short median emargination) or divided for almost whole length forming pair of long subparallel spines or flattened lobes; supraorbital lobes absent��������������������������������������������������������������������������������������������������������������������������������9 Supraorbital lobes blunt, rounded, appressed to and not reaching anteriorly beyond frontal lobes (Fig. 14.106g, i)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������7 Supraorbital lobes angular to pointed, well demarcated, reaching anteriorly to or beyond frontal lobes (Fig. 14.106a, f)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������8 Carapace widest at fourth anterolateral lobe (Fig. 14.106g). Cheliped carpus inner margin unarmed (Fig. 14.106o)������������������������������������������������������������������������������������������������������������������������������������������������������������ Permanotus Carapace widest at third anterolateral lobe (Fig. 14.106i). Cheliped carpus inner margin with blunt inner spine (Fig. 14.106p)������������������������������������������������������������������������������������������������������������������������������������������������������� Tauropus

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Marine Decapod Crustacea

Antennal article 2 2.0–2.7 times as long as wide (Fig. 14.106n)��������������������������������������������������������������������������������� Harrovia Antennal article 2 4.2–5.7 times as long as wide (Fig. 14.106m)���������������������������������������������������������������������� Ceratocarcinus Rostrum undivided or with shallow median emargination not exceeding one-third rostrum length (Fig. 14.106c, d, e, l)�������������������������������������������������������������������������������������������������������������������������������������������������������������������10 – Rostrum deeply and widely incised for almost whole length, forming pair of long subparallel spines or flattened lobes (Fig. 14.106b, k)�����������������������������������������������������������������������������������������������������������������������������������������������13 10. Pereopods 2–5 meri upper margin rounded, blunt, without longitudinal crest (Fig. 14.106r, t)���������������������������������������11 – Pereopods 2–5 meri upper margin with thin longitudinal crest (Fig. 14.106s, w)�������������������������������������������������������������� 12 11. Carapace lateral margin with prominent laterally directed tooth; rostrum usually short, with narrow to V-shaped median notch (Fig. 14.106e)����������������������������������������������������������������������������������������������������������������Gonatonotus – Carapace lateral margin with rounded or angular point; rostrum undivided, without median notch (Fig. 14.106c)�������������������������������������������������������������������������������������������������������������������������������������������������������������� Echinoecus 12. Pereopods 2–5 meri dorsal crest without distal spine (Fig. 14.106s)���������������������������������������������������������������������Eumedonus – Pereopod2 2–5 meri dorsal crest with distal spine (Fig. 14.106w)��������������������������������������������������������������������������Zebridonus 13. Rostral lobes flattened, apex pointed or rounded (Fig. 14.106k). Pereopods 2–5 with upper margins of merus, carpus and propodus cristate, often forming high lobes; propodus lower distal margin with triangular crest, occluding with dactylus forming prehensile structure (Fig. 14.106v)���������������������������������������Zebrida – Rostral lobes slender (Fig. 14.106b). Pereopods 2–5 linear (Fig. 14.106q)�������������������������������������������������������������Cebudonus 14. Carapace anterolateral teeth usually distinct, angular or sharp (Figs 14.108, 14.109a–k), or absent (Fig. 14.107a). Thoracic sternite 8 usually concealed by pleonite 2 in male; if carapace anterolateral teeth absent or poorly developed, thoracic sternite 8 not visible on each side of pleonite 2 in male�������������������������������������15 – Carapace anterolateral teeth absent or at most weakly indicated, low, obtusely convex. Thoracic sternite 8 always visible on each side of pleonite 2 in male����������������������������������������������������Rhizopinae … 53 15. Carapace subcircular-narrowly ovate, frontal and anterolateral margins smooth, unbroken, without teeth, granules or notches (Fig. 14.107a)������������������������������������������������������������ Hapalonotinae … Hapalonotus – Carapace variously shaped, front and anterolateral margins variously ornamented; if front entire, anterolateral teeth or lobes always present������������������������������������������������������������������������������������������������Pilumninae … 16 16. Carapace markedly wider anteriorly than posteriorly, asymmetrically hexagonal, appearing approximately triangular; front appearing trilobed, with narrow median fissure (Fig. 14.108c)����������������������������������Garthopilumnus – Carapace transversely ovate, subhexagonal or subquadrate; front undivided, bilobed or quadrilobate�������������������������17 17. Suborbital margin prominently produced anteriorly for full width forming wide shelf visible in dorsal view (Fig. 14.108b, j)���������������������������������������������������������������������������������������������������������������������������������������������������������������������������18 – Suborbital margin not prominently produced anteriorly for full width, at most with inner margin produced forwards as rounded or angular lobe�������������������������������������������������������������������������������������������������������������������������������������19 18. Carapace subhexagonal (Fig. 14.108j). Cheliped covered with rounded granules. Pereopods 2–5 smooth, without tubercles (Fig. 14.111i)���������������������������������������������������������������������������������������������������������������������������Colerolumnus – Carapace transversely ovate. Cheliped covered with acute granules and large, rounded or elongated, flat-topped tubercles. Pereopods 2–5 carpi, propodi with irregular, elongated, flat-topped tubercles (Fig. 14.108b)������������������������������������������������������������������������������������������������������������������������������������������������������ Balssopilumnus 19. Carapace glabrous or at most with few fine scattered setae or short tomentum around anterior margins, never covering most or entire surface; upper surface smooth, without setose ridges or patches of granules������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 20 – Carapace variously but obviously setose; upper surface usually rugose, with ridges, patches of granules or other ornamentation���������������������������������������������������������������������������������������������������������������������������������������������������������������� 29 20. Carapace rounded-subquadrate (Fig. 14.109c, j). Pereopod 3 longer than twice carapace length������������������������������������21 – Carapace transversely ovate. Pereopod 3 shorter than twice carapace length�������������������������������������������������������������������� 22 21. Carapace front entire, without median or lateral notches; anterolateral margin with low, blunt, granular lobes (Fig. 14.109j). Gonopod 1 apex strongly recurved, hooked (Fig. 14.111x)����������������������������������������Xestopilumnus 8. – 9.

14 – Brachyura – crabs

– 22. – 23. – 24. – 25. – 26. – 27. – 28. – 29. – 30. – 31. – 32. – 33. – 34. – 35. – 36. – 37.

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Carapace front with median and lateral notches; anterolateral margin with 2 anteriorly curved spines (Fig. 14.109c). Gonopod 1 apex straight (Fig. 14.111u)�������������������������������������������������������������������������������������������Peleianus Carapace anterolateral margin smooth, without teeth or lobes (Fig. 14.108p). Pereopod 5 ischium, merus with spines along lower margin (Fig. 14.111l)��������������������������������������������������������������������������������������������������Gorgonariana Carapace anterolateral margin with teeth or lobes. Pereopod 5 ischium, merus unarmed along lower margin�������������� 23 Thoracic sternite 8 of male partially exposed on each side of pleonite 2 (Fig. 14.110g)�����������������������������������Eurycarcinus Thoracic sternite 8 of male completely concealed by pleonite 2 (Fig. 14.110h)�������������������������������������������������������������������� 24 Male telson not reaching midlength of sternite 4 (Fig. 14.110d)������������������������������������������������������������������������ Heteropanope Male telson extending to or beyond midlength of sternite 4�������������������������������������������������������������������������������������������������� 25 Eyestalk as long as cornea (Figs 14.108o, 14.109f)�������������������������������������������������������������������������������������������������������������������� 26 Eyestalk longer than cornea (e.g. Fig. 14.108t)�������������������������������������������������������������������������������������������������������������������������� 28 Suborbital margin straight, inner margin not produced forwards. Antennal flagellum half as long as orbit width (Fig. 14.109f)���������������������������������������������������������������������������������������������������������������������������������������������� Serenolumnus Suborbital margin with blunt inner lobe produced forwards, visible in dorsal view. Antennal flagellum at least as long as orbit width (Fig. 14.108o)����������������������������������������������������������������������������������������������������������������������������� 27 Gonopod 1 apex simple, undivided (Fig. 14.111s)��������������������������������������������������������������������������������������������Glabropilumnus Gonopod 1 apex bifurcate (Fig. 14.111t)������������������������������������������������������������������������������������������������������������������������ Itampolus Eyestalk cylindrical (Fig. 14.108t). Cheliped palm outer surface entirely covered with sharp granules (Fig. 14.111d)����������������������������������������������������������������������������������������������������������������������������������������������������������� Lentilumnus Eyestalk pyriform, with distinct constriction at base of cornea (Fig. 14.109k). Cheliped palm outer surface with sharp granules proximally, smooth distally (Fig. 14.111g)����������������������������������������������������������������������������Xlumnus Pereopods 2–5 meri, carpi upper margins with thin, upraised crest (Fig. 14.111m)����������������������������������� Lophopilumnus Pereopods 2–5 meri, carpi upper margins smooth, granular or spinous����������������������������������������������������������������������������� 30 Carapace entirely covered with fine, short, velvet-like tomentum�����������������������������������������������������������������������������������������31 Carapace with long setae or mixture or short and long setae�������������������������������������������������������������������������������������������������32 Carapace front bilobed, with shallow median notch; surface smooth or coarsely granulate (Fig. 14.108a)������Actumnus Carapace front entire, without median notch; surface smooth (Fig. 14.109b)������������������������������������������������� Neoactumnus Gonopod 1 straight, at least in distal half, not recurved apically (Fig. 14.111p, v)���������������������������������������������������������������33 Gonopod 1 sinuous, usually recurved apically (Fig. 14.111w)�������������������������������������������������������������������������������������������������35 Pereopods 2–5 slender; meri and carpi upper margins spinose; pereopod 3 longer than twice carapace length, merus more than 4 times as long as wide (Fig. 14.111k)�������������������������������������������������������������������������� Danielum Pereopods 2–5 stout, merus and carpus unarmed; pereopod 3 shorter than twice carapace length, merus less than 3 times as long as wide�������������������������������������������������������������������������������������������������������������������������������������������� 34 Male telson about twice as long as wide, with straight lateral margins (Fig. 14.110i). Gonopod 1 gently tapering, evenly slender (Fig. 14.111p)������������������������������������������������������������������������������������������������������������Bathypilumnus Male telson ~1.4 times as long as wide, with concave lateral margins (Fig. 14.110o). Gonopod 1 proximal half strongly inflated, swollen, distal half slender (Fig. 14.111v)��������������������������������������������������������������� Priapipilumnus Carapace setation dense, short, forming vermiform or labyrinth-like pattern partially corresponding to underlying curved ridges on carapace surface (Fig. 14.109h)�����������������������������������������������������������������������������Vellumnus Carapace setation uniform or coarsely irregular in no distinct pattern, surface granular, spinular or nodular��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 36 Carapace anterolateral teeth narrow, separated by short intervals, not immediately abutting each other, spiniform or blunt-subconical, occasionally appearing as low angular lobes�����������������������������������������������������������������37 Carapace anterolateral teeth broad, closely spaced, bases abutting, appearing as prominent triangular teeth or lobes (lobes sometimes indistinct)����������������������������������������������������������������������������������������������������������������������������������� 43 Carapace outline almond-shaped; setation short, even, not obscuring surface; regions not indicated (Fig. 14.108m)��������������������������������������������������������������������������������������������������������������������������������������������������������� Ericiolumnus

670

– 38. – 39. – 40. – 41. – 42. – 43. – 44. – 45. – 46. – 47. – 48. – 49. – 50. –

Marine Decapod Crustacea

Carapace outline transversely ovate to subquadrate; variously with simple, plumose or clubbed setae, long or short, sometimes substantially obscuring surface; regions weakly to well marked������������������������������������������������� 38 Carapace surface irregularly ornamented with granules and prominent glabrous nodules or ‘islands’ in addition to long setae and tomentose patches (Figs 14.108u, 14.112i)���������������������������������������������������������������� Lophoplax Carapace surface granulation, if present, evenly sized, without large glabrous nodules��������������������������������������������������� 39 Carapace outline transversely ovate to subhexagonal. Walking legs stout; pereopod 3 shorter than twice carapace length (Pl. 60e–k)�����������������������������������������������������������������������������������������������������������������������������������������Pilumnus Carapace outline rounded-subquadrate. Walking legs slender; pereopod 3 longer than twice carapace length (Figs 14.108d, i, 14.109g, i)������������������������������������������������������������������������������������������������������������������������������������������ 40 Carapace front without anteriorly directed fringe of long setae���������������������������������������������������������������������������������������������41 Carapace front with anteriorly directed fringe of long setae (Fig. 14.108d)������������������������������������������������������������������������� 42 Carapace anterolateral teeth small, sharp (Fig. 14.109g). Cheliped palm outer surface covered with long, boletate or thorn-like spines (Fig. 14.111f). Pereopods 2–5 carpi, propodi upper margins spinose (Fig. 14.111n). Male telson longer than wide (Fig. 14.110q)���������������������������������������������������������������������������������� Takedana Carapace anterolateral teeth submarginal, subcylindrical, blunt (Fig. 14.108q). Cheliped palm outer surface with widely-spaced rounded or acute tubercles (Fig. 14.111b). Pereopods 2–5 carpi, propodi upper margins unarmed. Male telson wider than long (Fig. 14.110l)����������������������������������������������������������� Lamarckopilumnus Carapace exorbital angle bluntly rounded (Fig. 14.108i). Pereopods 2–5 meri upper margin unarmed (Fig. 14.111h). Male pleonite 1 twice as wide as pleonite 2 (Fig. 14.110j)����������������������������������������������������� Bossacarcinus Carapace exorbital angle forming small tooth (Fig. 14.109i). Pereopods 2–5 meri upper margin usually spinose (Fig. 14.111o). Male pleonite 1 little wider than pleonite 2 (Fig. 14.110r)�������������������������������������������Viaderiana Cheliped palm outer surface of one or both chelipeds smooth (Fig. 14.111e) or microscopically granular������������������� 44 Cheliped palm outer surface both chelipeds tuberculate or spinose, setose������������������������������������������������������������������������ 47 Carapace with 3 anterolateral teeth (Fig. 14.109d). Male pleonite 3 ~3 times as wide as pleonite 4 (Fig. 14.110p)������������������������������������������������������������������������������������������������������������������������������������������������������������ Rhizopoides Carapace with 4 or 5 anterolateral teeth (e.g. Fig. 14.108g, s). Male pleonite 3 less than 1.5 times as wide as pleonite 4����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 45 Carapace front narrower than one-quarter carapace width (Fig. 14.108s). Male pleonites 1 and 2 of equal width (Fig. 14.110n)����������������������������������������������������������������������������������������������������������������������������������������������������Leelumnus Carapace front exceeding one-third carapace width. Male pleonite 1 wider than pleonite 2������������������������������������������� 46 Carapace regions indicated, gastro-cardiac grooves distinct (Fig. 14.108g). Male telson extending beyond midlength of sternite 4 (Fig. 14.110b)��������������������������������������������������������������������������������������������������Benthopanope Carapace regions weakly indicated, gastro-cardiac grooves absent or at most, faint (Fig. 14.108f). Male telson not reaching midlength of sternite 4 (Fig. 14.110f)���������������������������������������������������������������������Pilumnopeus Pereopod 5 ischium, merus with spines along lower margin (Fig. 14.111j)�������������������������������������������������������������������������� 48 Pereopod 5 ischium, merus unarmed along lower margin����������������������������������������������������������������������������������������������������� 49 Suborbital margin straight, without inner projection (Figs 14.108k, 14.109o). Thoracic sternites 1–3 indistinguishably fused in male (Fig. 14.110c)��������������������������������������������������������������������������������������������� Cryptopilumnus Suborbital margin with rounded inner projection. Thoracic sternites 1 and 2 fused, separated from sternite 3 by distinct groove in male (Fig. 14.110a)��������������������������������������������������������������������������������������������Aniptumnus Carapace regions protuberant, forming large, naked, granular nodules, especially on anterior half; with close pile of short setae between and around nodules (Fig. 14.108e)������������������������������������������������� Lobopilumnus Carapace regions indicated or not, variously spiny, granulate or smooth, but not nodulose-protuberant; variously with simple, plumose or clubbed, long or short setae��������������������������������������������������������������������������������������� 50 Male pleonite 1 distinctly wider than pleonite 2 (Fig. 14.110k); thoracic sternite 8 exposed on each side of pleonite 2������������������������������������������������������������������������������������������������������������������������������������������������ Heteropilumnus (part) Male pleonite 1 as wide as or little wider than pleonite 1 (Fig. 14.110m), thoracic sternite 8 not exposed on each side of pleonite 2���������������������������������������������������������������������������������������������������������������������������������������������������������������51

14 – Brachyura – crabs

51. – 52. – 53. – 54. – 55. – 56. – 57. – 58. – 59. – 60. – 61. – 62. – 63. – 64. – 65. – 66. – 67. – 68. – 69. –

671

Carapace widest across fourth (last) anterolateral tooth, fourth tooth equal to or larger than third tooth (Fig. 14.109e)����������������������������������������������������������������������������������������������������������������������������������Serenepilumnus Carapace widest across third anterolateral tooth, fourth tooth smaller than third (Fig. 14.108r)������������������������������������52 Carapace frontal lobes anterior margin broadly convex (Fig. 14.108r). Male telson not reaching midlength of sternite 4 (Fig. 14.110e)������������������������������������������������������������������������������������������������������������������������������������ Latopilumnus Carapace frontal lobes anterior margin straight (Fig. 14.109a). Male telson extending beyond midlength of sternite 4�������������������������������������������������������������������������������������������������������������������������������������������������������������� Nanopilumnus Carapace semicircular, lateral margins distinctly divergent posteriorly (Fig. 14.112e, o)������������������������������������������������� 54 Carapace ovate to subquadrate, lateral margins subparallel or convergent posteriorly������������������������������������������������������55 Antennal flagellum slender, sparsely setose. Maxilliped 3 merus rounded distolaterally (Fig. 14.113o)�������������������������������������������������������������������������������������������������������������������������������������������������� Xenophthalmodes Antennal flagellum densely plumose (Fig. 14.112e). Maxilliped 3 produced distolaterally (Fig. 14.113m)����������Mertonia Carapace margin fully circumscribed by continuous narrow ridge (Fig. 14.112f)������������������������������������������� Paranotonyx Carapace margin at most with partial ridge around parts of anterior and posteror margins������������������������������������������ 56 Male pleonite 3 not reaching pereopod 5 coxa (Fig. 14.113e, g, h, i)��������������������������������������������������������������������������������������57 Male pleonite 3 reaching pereopod 5 coxa (Fig. 14.113j)��������������������������������������������������������������������������������������������������������� 64 Male pleonite 1 not reaching pereopod 5 coxa (Fig. 14.113g, i)���������������������������������������������������������������������������������������������� 58 Male pleonite 1 broadened, reaching pereopod 5 coxa (Fig. 14.113e, j)���������������������������������������������������������������������������������62 Maxilliped 3 merus distolaterally produced (Fig. 14.113k)�������������������������������������������������������������������������������������� Ceratoplax Maxilliped 3 merus not distolaterally produced (Fig. 14.113l, n)�������������������������������������������������������������������������������������������59 Maxilliped 3 exopod reaching to distal end of merus; merus wider than ischium at midlength (Fig. 14.113l). Epistome lower medial margin cut into 2 narrow rectangular lobes (Fig. 14.113a)�������������������������������������������������������� 60 Maxilliped 3 exopod reaching to midlength of merus; merus narrower than ischium at midlength (Fig. 14.113n, o). Epistome lower medial margin not cut into rectangular lobes, at most with shallow median notch (Fig. 14.113b, c)�������������������������������������������������������������������������������������������������������������������������������������������������61 Fronto-orbital width half carapace width (Fig. 14.112k). Carapace surface finely setose���������������������������������������� Rhizopa Fronto-orbital width two-thirds carapace width (Fig. 14.112d). Carapace surface glabrous������������������������ Luteocarcinus Carapace anterolateral margins smooth, unarmed, not lined with granules (Fig. 14.112g)�������������������������� Paraselwynia Carapace anterolateral margins irregular, lobate, dentate or lined with granules (Fig. 14.112n)��������������� Typhlocarcinus Pereopods 2–5 coxae with denticulate plate partially overlapping ischium (Fig. 14.113q)��������������������������������Cryptolutea Pereopods 2–5 coxae without denticulate plate������������������������������������������������������������������������������������������������������������������������ 63 Eye usually fixed, cornea reduced, sometimes pigmented (Fig. 14.113c)����������������������������������������������������� Typhlocarcinops Eye mobile, cornea well developed, pigmented (Fig. 14.112h)�������������������������������������������������������������������������������� Pronotonyx Carapace angular, subquadrate to subhexagonal, anterolateral margin distinctly shorter than posterolateral margin; front without fringe of long setae�������������������������������������������������������������������������������������������������� 65 Carapace transversely ovate, anterolateral margin as long as or slightly shorter than posterolateral margin; front usually with fringe of long setae���������������������������������������������������������������������������������������������������������������������������������� 67 Carapace quadrate, posterolateral margins subparallel (Fig. 14.112l)��������������������������������������������������������������������������������� Ser Carapace subhexagonal, posterolateral margins strongly convergent (Fig. 14.112b)��������������������������������������������������������� 66 Carapace surface smooth, polished; anterior carapace and chelipeds densely setose�����������������������Pseudocryptocoeloma Carapace surface rugose, with 2 low transverse ridges; anteriorly carapace and chelipeds glabrous����������������Camptoplax Eyes below anterior carapace margin, not visible in dorsal view (even after removal of anterior setae) (Fig. 14.113d)����������������������������������������������������������������������������������������������������������������������������� Cryptocoeloma Eyes on anterior carapace margin, visible in dorsal view (after removal of anterior setae)���������������������������������������������� 68 Carapace surface irregular, variously granular, regions distinct (Fig. 14.108d)���������������������������Heteropilumnus (in part) Carapace surface smooth, regions indistinct or absent (Fig. 14.112a, j)������������������������������������������������������������������������������� 69 Cheliped palm outer surface densely coarsely granulate (Fig. 14.113p)����������������������������������������������������������Pseudolitochira Cheliped palm outer surface smooth or finely granulate, with or without setae (Fig. 14.112a)������������������� Zehntneriana

672

Marine Decapod Crustacea

Fig. 14.105.  Pilumnidae, Calmaniinae, Calmania. a, carapace; b, chela, outer face.

Subfamily Calmaniinae Števčić, 1991

Ng and Chia (1999) tentatively retained the sea cucumber endosymbiont, Hapalonotus, in Eumedoninae on the basis of its echinoderm association, but acknowledged that it otherwise had little in common with other eumedonines and probably belonged in a separate subfamily. Števčić (2005) proposed the subfamily Hapalonotinae for Hapalonotus, followed here. Diagnosis. Carapace subpentagonal-subhexagonal to ovate; front strongly produced beyond orbits, often as single rostrum or pair of ‘horns’, without fringe of setae. Antennules folding obliquely forwards. Cheliped fingers without gape, occlusal margins at most sparsely setose. Pereopods 2–5 with dactylus-propodal lock. Cebudonus Ng, 2014 Diagnosis. Carapace lateral margin with angular point; front produced as 2 slender rostral spines. Maximum cl. 13 mm. Shelf, slope (80–350 m; host unknown, probably sea urchins). Central Indo-Pacific (Philippines). 1 species (Ng 2014).

Calmaniines are minute crabs occurring mostly in shallow tropical reefs habitats, amongst rubble and grit. They are recognisable by the combination of obliquely folding antennules, a fringe of long setae on the carapace front and the setose gape between the cheliped fingers. Two genera, Ralumnia and Calmania, have been recognised but Takeda (1989) showed them to be synonymous. Diagnosis. Carapace quadrate to longitudinally ovate; front at most weakly produced beyond orbits, with fringe of long setae; surface with tufts of setae. Antennules folding obliquely forwards. Cheliped fingers with proximal gape when closed, margins of gape with dense brush of bristly setae. Pereopods 2–5 without dactylus-propodal lock.

Diagnosis. Carapace lateral margin with angular point; front with 2 lobes separated by median incision, flanked on each side by supraorbital lobe; supraorbital lobes angular to pointed, well demarcated, reaching anteriorly to or beyond frontal lobes; anterolateral margin incised. Antennal article 2 4.2–5.7 times as long as wide. Maximum cl. 13 mm. Intertidal–shelf (0–53 m; obligate symbiont of crinoids). Central Indo-Pacific, Temperate Australasia. 3 species (Chia and Ng 1998: key to 3 species).

Calmania Laurie, 1906

Echinoecus Rathbun, 1894

Intertidal–shelf (0–190 m). Maximum cl. 7 mm. Western and Central Indo-Pacific. 5 species (Serène and Umali 1972: key to species included in Ralumnia, junior synonym, and Calmania).

Diagnosis. Carapace lateral margin with angular point; front produced as triangular rostrum with rounded apex. Pereopods 2–5 meri upper margin rounded, blunt, without longitudinal crest. Maximum cl. 16 mm. Intertidal–shelf (0–110 m; obligate symbiont of sea urchins). Indo-West Pacific. 3 species (Chia et al. 1999: key to species).

Subfamily Eumedoninae Dana, 1852 Eumedonines are ectosymbionts of echinoderms, specifically sea urchins (echinoids) and feather stars (crinoids). Although common, eumedonines are seldom seen because of their small-size (~10 mm or less) and concealment amongst the spines or sea-urchins or arms of feather stars. All occur in the Indo-West Pacific, mainly on coral reefs, but extend into temperate Australia. Eumedonines were once thought to be allied to parthenopids, or to represent a separate xanthoid family allied to Pilumnidae (Števčić et al. 1988). The group is now considered a pilumnid subfamily on the basis of larval and adult morphology (Ng and Clark 2000).

Ceratocarcinus White, 1847

Eumedonus H. Milne Edwards, 1834 Diagnosis. Carapace lateral margin with angular point; front produced as rostrum with apical notch. Pereopods 2–5 meri upper margin with thin longitudinal crest. Maximum cl. 15 mm. Subtidal–slope (30–235 m; obligate symbiont of sea urchins). Western and Central Indo-Pacific, Temperate SE Australia. 5 species (Chia and Ng 2000: rediagnosis, key to species).

Gonatonotus White, 1847 Diagnosis. Carapace lateral margin with prominent laterally directed tooth; front produced as short rostrum, with narrow to

14 – Brachyura – crabs

673

Fig. 14.106.  Pilumnidae, Eumedoninae. a, Ceratocarcinus longimanus White, 1847. Carapace: b, Cebudonus; c, Echinoecus; d, Eumedonus; e, Gonatonotus; f, Harrovia; g, Permanotus; h, Rhabdonotus; i, Tauropus; j, Tiaramedon; k, Zebrida; l, Zebridonus. Antennules, antennae, eyestalks: m, Ceratocarcinus; n, Harrovia. Cheliped: o, Permanotus; p, Tauropus. Pereopod 5: q, Cebudonus; r, Echinoecus; s, Eumedonus; t, Gonatonotus; u, Rhabdonotus; v, Zebrida; w, Zebridonus.

V-shaped median notch. Pereopods 2–5 meri upper margin rounded, blunt, without longitudinal crest. Maximum cl. 16 mm. Intertidal–shelf (0–182 m; obligate symbiont of sea urchins). Temperate Northern Pacific, Western Indo-Pacific, Temperate SE Australia. 3 species (Chia and Ng 2000: rediagnosis, key to species).

Harrovia Adams & White, 1849 Diagnosis. Carapace lateral margin with angular point; front with 2 lobes separated by median incision, flanked on each side by supraorbital lobe; supraorbital lobes angular to pointed, well demarcated, reaching anteriorly to or beyond frontal lobes; anterolateral margin incised. Antennal article 2 2.0–2.7 times as long as wide. Maximum cl. 14 mm.

Subtidal, shelf (1–154 m; obligate symbiont of crinoids). Western and Central Indo-Pacific. 7 species (Chia and Ng 1998: rediagnosis, key to species).

Permanotus Chia & Ng, 1998 Diagnosis. Carapace widest at fourth anterolateral lobe; front with 2 lobes separated by median incision, flanked on each side by supraorbital lobe; supraorbital lobes blunt, rounded, appressed to and not reaching anteriorly beyond frontal lobes; anterolateral margin incised. Cheliped carpus inner margin unarmed. Maximum cl. 7 mm. Subtidal, shelf (1–37 m; obligate symbiont of crinoids). Western and Central Indo-Pacific. 1 species (Chia and Ng 1998).

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Marine Decapod Crustacea

Rhabdonotus A. Milne-Edwards, 1879 Diagnosis. Carapace rounded, ovate-pyriform. Maximum cl. 12 mm. Subtidal–slope (5–571 m: obligate symbiont of crinoids). Western and Central Indo-Pacific. 3 species (Chia and Ng 1995: key to species).

Tauropus Chia & Ng, 1998 Diagnosis. Carapace widest at third anterolateral lobe; front with 2 lobes separated by median incision, flanked on each side by supraorbital lobe; supraorbital lobes blunt, rounded, appressed to and not reaching anteriorly beyond frontal lobes; anterolateral margin incised. Cheliped carpus inner margin with blunt inner spine. Maximum cl. 11 mm. Subtidal, shelf (1–82 m: host unknown, probably crinoids). Central Indo-Pacific (South China Sea, Japan). 1 species (Chia and Ng 1998).

Tiaramedon Chia & Ng, 1998 Diagnosis. Carapace lateral margin with angular point; front with 2 lobes separated by median incision, flanked on each side by supraorbital lobe; supraorbital lobes prominent, about half carapace length; surface with pair of long upright protogastric spines; anterolateral margin entire. Cheliped carpus inner margin with sharp inner spine. Maximum cl. 10 mm. Intertidal–shelf (0–57 m; obligate symbiont of crinoids). Western and Central Indo-Pacific. 1 species (Chia and Ng 1998).

Zebrida White, 1847 Diagnosis. Carapace lateral margin with anteriorly directed lamelliform lobe; front produced as flat rostrum, deeply and widely incised for almost whole length, forming pair of long subparallel acute or rounded lobes. Maximum cl. 14 mm. Intertidal–shelf (0–40 m; obligate symbiont of sea urchins). Western and Central Indo-Pacific, Temperate SW Australia. 3 species (Ng and Chia 1999: rediagnosis, key to species).

is an obligate endosymbiont of sea cucumbers, and on this basis, was tentatively assigned to Eumedoninae, despite its incongruent morphology (Chia and Ng 1999; Ng et  al. 2008). Aside from its association with echinoderms, Hapalonotus has little in common with eumedonines, particularly differing in having transversely rather than obliquely folding antennules, a weakly rather than strongly produced front and in its endo- rather than ecto-symbiotic ecology; as such, we follow Števčić (2005) in recognising a separate subfamily, Hapalonotinae, for Hapalonotus. Diagnosis. Carapace subcircular, ~1.2 times as wide as long; front smooth, without median notch; smooth, covered with short, fine translucent tomentum; anterolateral margin smooth, unbroken, without teeth, granules or notches. Thoracic sternite 8 of male completely covered by anterior pleonites, not visible on each side of pleonite 2. Antennules folding transversely. Cheliped fingers without gape, occlusal margins at most sparsely setose. Pereopods 2–5 with weakly developed dactylus-propodal lock. Hapalonotus Rathbun, 1897 sea-cucumber crab Maximum cl. 23 mm. Subtidal (2–11 m). Inside sea cucumbers (Holothuria). Central Indo-Pacific (Indonesia, Papua New Guinea). 1 species (Chia and Ng 1999: description, figures; Vandenspiegel et al. 1992: general biology).

Subfamily Pilumninae Samouelle, 1819 Pilumnines are the classic ‘hairy’ crabs, frequently having dense, wiry setae obscuring the carapace surface and margins. The carapace is generally transversely ovate with prominent anterolateral teeth, and the pleon proximally narrow with sternite 8 usually fully concealed by pleonite 2. They are common on rocky and coral reefs, especially in

Zebridonus Chia, Ng & Castro, 1995 Diagnosis. Carapace lateral margin with angular point; front produced as rostrum with apical notch. Pereopods 2–5 meri upper margin with thin longitudinal crest ending in distal spine. Maximum cl. 7 mm. Subtidal (45 m: host unknown, probably sea urchins). Central Indo-Pacific (Australia). 1 species (Chia et al. 1995).

Subfamily Hapalonotinae Števčić, 2005 The peculiar and poorly known genus Hapalonotus has been variously considered to be a pinnotherid (De Man 1879; Schmitt et  al. 1973; Tesch 1918) and xanthid (Balss 1933), but Vandenspiegel et  al. (1992) showed it to be a pilumnid based on the distinctive gonopods. Hapalonotus

Fig. 14.107.  Pilumnidae, Hapalonotinae, Hapalonotus. a, habitus; b, antennules, antennae, eyestalks.

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Fig. 14.108.  Pilumnidae, Pilumninae. a, Actumnus setifer (De Haan, 1835); b, Balssopilumnus bolitifer (Monod, 1956); c, Garthopilumnus palmeri (Garth, 1986); d, Heteropilumnus fimbriatus (H. Milne Edwards, 1834); e, Lobopilumnus agassizii (Stimpson, 1871) (Source: Fish. Bull. 65: 1–298); f, Pilumnopeus serratifrons (Kinahan, 1856). Carapace: g, Benthopanope; h, Bathypilumnus; i, Bossacarcinus; j, Colerolumnus; k, Cryptopilumnus; l, Danielum; m, Ericiolumnus; n, Eurycarcinus; o, Glabropilumnus; p, Gorgonariana; q, Lamarckopilumnus; r, Latopilumnus; s, Leelumnus; t, Lentilumnus; u, Lophoplax.

tight crevices and amongst sponge, algal and bryozoan encrustations. These setal coverings of many species are often loaded with sediment, enhancing camouflage but impeding identification. The taxonomic limits of Pilumninae and Rhizopinae with respect to each other remain to be phylogenetically

assessed but here, largely follow Ng et al. (2008) with some pragmatic modifications. Pilumninae as recognised here emphasises the presence of prominent anterolateral teeth on the carapace, and as such, Lophopilumnus and Rhizopoides, with well developed anterolateral teeth, are included in Pilumninae instead of Rhizopinae, as are Itampolus and

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Marine Decapod Crustacea

Fig. 14.109.  Pilumnidae, Pilumninae. Carapace (setae not shown): a, Nanopilumnus; b, Neoactumnus; c, Peleianus; d, Rhizopoides; e, Serenepilumnus; f, Serenolumnus; g, Takedana; h, Vellumnus; i, Viaderiana; j, Xestopilumnus; k, Xlumnus. Front, antennules, antennae, orbits: l, Colerolumnus; m, Takedana; n, Vellumnus. Right suborbital ridge, antenna: o, Cryptopilumnus; p, Lentilumnus; q, Vellumnus.

Peleianus, which are morphologically similar to the pilumnine Glabropilumnus. Heteropilumnus is included in Pilumninae, but, being morphologically heterogeneous (Davie and Humpherys 1997), appears twice in the key below, once among the pilumnines and once among the rhizopines. Heteropanope tuberculidens Monod, 1956 and H. acanthocarpus Crosnier, 1967, although congeneric, do not properly belong in Heteropanope sensu stricto (Davie 1989a) and are herein transferred to Latopilumnus. The genus Parapleurophrycoides Nobili, 1906 (type species: P. roseus Nobili, 1906) from French Polynesia is based on a juvenile, possibly of a known genus, but is presently unidentifiable based the brief type description; it is excluded from the key below. The unusual monotypic genus, Garthopilumnus, also based on a juvenile (but well illustrated; Garth 1986: Fig. 4) (Pl. 14.108c) is tentatively regarded as a pilumnine; its gonopods are currently unknown. Lastly, we recommend caution when identifying members of the speciose genera, Actumnus, Heteropilumnus and Pilumnus, many of which are currently poorly known and require redescription – some may prove to belong to other genera. Diagnosis. Carapace front usually divided (when entire, anterolateral teeth or lobes always present); anterolateral margin usually with distinct angular or sharp teeth. Thoracic sternite 8 of male usually completely covered by anterior pleonites, rarely visible on each side of pleonite 2 (when anterolateral teeth weak or absent, male sternite 8 always concealed by pleonite 2). Antennules folding transversely.

Cheliped fingers without gape, occlusal margins at most sparsely setose. Pereopods 2–5 with or without dactyluspropodal lock. Implicit pilumnine generic attributes. Carapace transversely ovate; front without anteriorly directed fringe of long setae; setose; upper surface usually rugose, with ridges, patches of granules or other ornamentation; suborbital margin not prominently produced anteriorly for full width, at most with inner margin produced forwards as rounded or angular lobe. Eyestalk mobile, cornea well developed. Gonopod 1 sinuous, usually recurved apically. Actumnus Dana, 1851 Diagnosis. Front bilobed, with median notch; carapace entirely covered with fine, short, velvet-like tomentum; smooth or coarsely granulate; posterolateral surface often gently sunken, accommodating folded pereopod 5; anterolateral margin with obsolete or triangular lobes. Maximum cl. 31 mm. Intertidal–shelf (0–200 m). Indo-West Pacific, Temperate Southern Africa, Temperate Australasia. 29 species. Local keys to species only are available (Dai and Yang 1991; Naderloo 2017; Sakai 1976; Takeda and Komatsu 2017).

Aniptumnus Ng, 2002 Diagnosis. Carapace rounded-subquadrate; front exceeding onethird carapace width, with anteriorly directed fringe of long setae; suborbital margin with rounded inner projection; anterolateral

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Fig. 14.110.  Pilumnidae, Pilumninae. Thoracic sternum (sternites 1–4 or more, with or without male pleon or telson): a, Aniptumnus; b, Benthopanope; c, Cryptopilumnus; d, Heteropanope; e, Latopilumnus; f, Pilumnopeus. Male pleonites 1–3, thoracic sternite 8 (shaded), coxa 5: g, Eurycarcinus; h, Heteropanope. Male pleon: i, Bathypilumnus; j, Bossacarcinus; k, Heteropilumnus; l, Lamarckopilumnus; m, Latopilumnus; n, Leelumnus; o, Priapipilumnus; p, Rhizopoides; q, Takedana; r, Viaderiana.

Fig. 14.111.  Pilumnidae, Pilumninae. Chela, outer face: a, Bathypilumnus; b, Lamarckopilumnus; c, Leelumnus; d, Lentilumnus; e, Rhizopoides; f, Takedana; g, Xlumnus. Pereopod 5: h, Bossacarcinus; i, Colerolumnus; j, Cryptopilumnus; k, Danielum; l, Gorgonariana; m, Lophopilumnus; n, Takedana; o, Viaderiana. Gonopod 1: p, Bathypilumnus; q, Danielum; r, Garthopilumnus; s, Glabropilumnus; t, Itampolus; u, Peleianus; v, Priapipilumnus; w, Vellumnus; x, Xestopilumnus.

margin with 2 lobes and 2 teeth. Thoracic sternites 1 and 2 fused, separated from sternite 3 by distinct groove in male. Cheliped palm outer surface both chelipeds tuberculate or spinose, setose. Pereopod 3 longer than twice carapace length. Pereopod 5 ischium, merus with spines along lower margin. Male pleonite 1 wider than pleonite 2, thoracic sternite 8 not exposed on each side of pleonite 2, pleonite 3 less than 1.5 times as wide as pleonite 4. Maximum cl. 10 mm. Intertidal–shelf (0–33 m). Western and Central Indo-Pacific. 3 species (Ng 2002: diagnosis, figures; Ng and Clark 2008: third species).

Balssopilumnus Števčić, 2011 Diagnosis. Carapace transversely ovate; suborbital margin prominently produced anteriorly for full width forming wide shelf visible in dorsal view; anterolateral margin with 3 teeth, first 2 large, triangular, lobiform, third tooth small, directed laterally. Cheliped covered with acute granules and large, rounded or elongated, flattopped tubercles. Pereopods 2–5 carpi, propodi with irregular, elongated, flat-topped tubercles. Maximum cl. 6 mm.

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Marine Decapod Crustacea

Subtidal (1–12 m). Tropical E Atlantic. 1 species (Monod 1956: figures of type species as Pilumnus boletifer Monod, 1956).

Bathypilumnus Ng & Tan, 1984 Diagnosis. Carapace subcircular to ovate; anterolateral teeth sharp, well spaced. Male telson about twice as long as wide, with straight lateral margins. Gonopod 1 obtusely bent, straight in distal half, not apically hooked. Maximum cl. 40 mm. Subtidal, shelf (1–128 m). Central Indo-Pacific. 3 species (Ng and Tan 1984).

Benthopanope Davie, 1989 Diagnosis. Carapace rounded-subquadrate; front exceeding onethird carapace width, with anteriorly directed fringe of long setae; regions indicated, gastro-cardiac grooves distinct; anterolateral margin with 4 lobes or teeth. Cheliped palm outer surface of one or both chelipeds smooth or microscopically granular, glabrous. Pereopod 3 longer than twice carapace length. Male pleonite 1 wider than pleonite 2, thoracic sternite 8 not exposed on each side of pleonite 2, pleonite 3 less than 1.5 times as wide as pleonite 4. Male telson extending beyond midlength of sternite 4. Maximum cl. 14 mm. Intertidal, subtidal (0–8 m). Western and Central Indo-Pacific. 6 species (Davie 1989a).

Bossacarcinus Števčić, 2011 Diagnosis. Carapace rounded-subquadrate; front with anteriorly directed fringe of long setae; anterolateral margin with 3 acute triangular teeth; exorbital angle bluntly rounded. Pereopod 3 longer than twice carapace length. Male pleonite 1 twice as wide as pleonite 2. Maximum cl. 6 mm. Subtidal, shelf (35–72 m). Central Indo-Pacific. 1 species (figures of type species: Serène 1964; Tesch 1918).

Colerolumnus Ng, 2010 Diagnosis. Carapace hexagonal; suborbital margin prominently produced anteriorly for full width forming wide shelf visible in dorsal view; anterolateral margin with 3 teeth, first 2 large, triangular, lobiform, third tooth small, directed laterally. Cheliped covered with rounded granules. Pereopods 2–5 smooth, without tubercles. Maximum cl. 5 mm. Subtidal. Central Indo-Pacific (Papua New Guinea). 1 species (Ng 2010).

Cryptopilumnus Hsueh, Huang & Ng, 2009 Diagnosis. Carapace rounded-subquadrate; front exceeding onethird carapace width, with anteriorly directed fringe of long setae; suborbital margin straight, without inner projection; anterolateral margin with 2 small teeth. Thoracic sternites 1–3 indistinguishably fused in male. Cheliped palm outer surface both chelipeds tuberculate or spinose, setose. Pereopod 3 longer than twice

carapace length. Pereopod 5 ischium, merus with spines along lower margin. Male pleonite 1 wider than pleonite 2, thoracic sternite 8 not exposed on each side of pleonite 2, pleonite 3 less than 1.5 times as wide as pleonite 4. Maximum cl. 5 mm. Intertidal–shelf (0–27 m). Western and Central Indo-Pacific. 3 species (Hsueh et al. 2009).

Danielum Vázquez-Bader & Gracia, 1995 Diagnosis. Carapace subcircular, about as wide as long; anterolateral margin with 4 acute teeth. Pereopods 2–5 slender, merus more than 4 times as long as wide; meri upper margin and carpi upper margins spinose. Pereopod 3 longer than twice carapace length. Gonopod 1 broadly curved, straight in distal half, not apically hooked. Maximum cl. 20 mm. Shelf (65–182 m). Tropical Atlantic (Gulf of Mexico). 1 species (Vázquez-Bader and Gracia 1995).

Ericiolumnus Ng & Rahayu, 2020 Diagnosis. Carapace almond-shaped; setation short, even, not obscuring surface; regions not indicated; anterolateral margin with 4 subdivided teeth. Maximum cl. 13 mm. Subtidal (to 20 m; associated with sponges). Central IndoPacific (Indonesia, New Caledonia). 1 species (Ng and Rahayu 2020a).

Eurycarcinus A. Milne-Edwards, 1867 Diagnosis. Carapace glabrous or at most with few fine scattered setae; upper surface smooth, without ridges; anterolateral margin with usually 3 teeth or lobes. Thoracic sternite 8 of male partially exposed on each side of pleonite 2. Pereopod 3 shorter than twice carapace length. Pereopod 5 ischium, merus unarmed along lower margin. Male telson not reaching beyond midlength of sternite 4. Maximum cl. 27 mm. Intertidal–shelf (0–32 m). Western and Central Indo-Pacific; one species exotic in Mediterranean (Ozcan et al. 2010). 4 species (Ng et al. 2018: differentiation of genus and 3 species).

Garthopilumnus Števčić, 2011 Diagnosis. Carapace markedly wider anteriorly than posteriorly, asymmetrically hexagonal, appearing approximately triangular; front appearing trilobed but with narrow median fissure; anterolateral margin with 4 complex teeth. Cheliped spinose. Maximum cl. 7 mm. Subtidal, shelf (4–27 m). Tropical Eastern Pacific (Ecuador) (Garth 1986: description of type species as Pilumnus palmeri Garth, 1986).

Glabropilumnus Balss, 1932 Diagnosis. Carapace glabrous or at most with few fine scattered setae; upper surface smooth, without ridges; suborbital margin with blunt inner lobe produced forwards, visible in dorsal view; anterolateral margin with 3 teeth, 2 broad, last smallest. Thoracic

14 – Brachyura – crabs

sternite 8 of male completely concealed by pleonite 2. Eyestalk as long as cornea. Antennal flagellum at least as long as orbit width. Pereopod 3 shorter than twice carapace length. Pereopod 5 ischium, merus unarmed along lower margin. Gonopod 1 sinuous, apex simple, undivided. Male telson extending to or beyond midlength of sternite 4. Maximum cl. 14 mm (Pl. 60a). Intertidal, subtidal (0–6 m). Indo-West Pacific. 6 species (Galil and Takeda 1988: key to 5 species).

Gorgonariana Galil & Takeda, 1988 Diagnosis. Carapace glabrous or at most with few fine scattered setae; upper surface smooth, without ridges; anterolateral margin smooth, without teeth or lobes. Pereopod 3 shorter than twice carapace length. Pereopod 5 ischium, merus with spines along lower margin. Maximum cl. 14 mm. Shelf, slope (32–398 m; associated with alcyonacean corals, Solenocaulon). Western and Central Indo-Pacific. 1 species (Galil and Takeda 1988).

Heteropanope Stimpson, 1858 Diagnosis. Carapace glabrous or at most with few fine scattered setae; upper surface smooth, without ridges; anterolateral margin with usually 3 teeth or lobes. Thoracic sternite 8 of male completely concealed by pleonite 2. Pereopod 3 shorter than twice carapace length. Pereopod 5 ischium, merus unarmed along lower margin. Male telson not reaching midlength of sternite 4. Maximum cl. 15 mm. Estuarine, intertidal. Western and Central Indo-Pacific. 2 species (Davie 1989a: rediagnosis, 2 species). The West African Heteropanope tuberculidens Monod, 1956 and H. acanthocarpus Crosnier, 1967 are transferred to Latopilumnus.

Heteropilumnus De Man, 1895 Diagnosis. Carapace rounded-subquadrate; front exceeding onethird carapace width, with anteriorly directed fringe of long setae; surface irregular, variously granular and setose, regions distinct; anterolateral margin with 4 or 5 broad, closely spaced teeth, bases abutting, or indistinct. Cheliped palm outer surface both chelipeds tuberculate or spinose, setose. Pereopod 3 longer than twice carapace length. Pereopod 5 ischium, merus unarmed along lower margin. Male pleonite 1 distinctly wider than pleonite 2 so thoracic sternite 8 exposed on each side of pleonite 2, pleonite 3 less than 1.5 times as wide as pleonite 4. Maximum cl. 19 mm (Pl. 60b). Intertidal–shelf (0–82 m). Western and Central Indo-Pacific. 20 species. The genus has often been treated as a member of Rhizopinae. Its species have not been reviewed. For comments see Ng (1987) and figures see Sakai (1976) and Davie and Humpherys (1997).

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with blunt inner lobe produced forwards, visible in dorsal view; anterolateral margin with 3 teeth, 2 broad, last smallest. Thoracic sternite 8 of male completely concealed by pleonite 2. Eyestalk as long as cornea. Antennal flagellum at least as long as orbit width. Pereopod 3 shorter than twice carapace length. Pereopod 5 ischium, merus unarmed along lower margin. Male telson extending beyond midlength of sternite 4. Gonopod 1 sinuous, apex bifurcate. Maximum cl. 4 mm. Intertidal, subtidal (0–5 m). Western Indo-Pacific (Réunion, Mauritius). 1 species (Serène and Peyrot-Clausade 1977).

Lamarckopilumnus Števčić, 2011 Diagnosis. Carapace rounded-subquadrate; anterolateral margin with submarginal, subcylindrical, blunt teeth. Cheliped palm outer surface with widely-spaced rounded or acute tubercles. Pereopods 2–5 carpi, propodi upper margins unarmed. Pereopod 3 longer than twice carapace length. Male telson wider than long. Maximum cl. 4 mm. Shelf (122–190 m). Central Indo-Pacific (Ogasawara Is, Japan). 1 species (Takeda and Ng 1997: description of type species).

Latopilumnus Türkay & Schuhmacher, 1985 Diagnosis. Carapace rounded-subquadrate, widest across third anterolateral tooth, fourth tooth smaller than third; front exceeding one-third carapace width, anterior margin broadly convex, with anteriorly directed fringe of long setae; anterolateral margin with 3 teeth, first usually low and broader than others. Cheliped palm outer surface both chelipeds tuberculate or spinose, setose. Pereopod 3 longer than twice carapace length. Pereopod 5 ischium, merus unarmed along lower margin. Male pleonite 1 wider than pleonite 2, thoracic sternite 8 not exposed on each side of pleonite 2, pleonite 3 less than 1.5 times as wide as pleonite 4. Male telson not reaching midlength of sternite 4. Maximum cl. 6 mm. Intertidal–shelf (0–55 m). Tropical E Atlantic, Western and Central Indo-Pacific. 6 species. (Ng and Clark 2008: rediagnosis; Türkay and Schuhmacher 1985). Heteropanope tuberculidens Monod, 1956 and H. acanthocarpus Crosnier, 1967, both from West Africa, are included here.

Leelumnus Mendoza & Ng, 2011 Diagnosis. Carapace rounded-subquadrate; front narrower than one-quarter carapace width, with anteriorly directed fringe of long setae; anterolateral margin with 4 anterolateral teeth, last 2 narrow. Cheliped palm outer surface of major cheliped smooth, glabrous, of minor cheliped granular, setose’. Pereopod 3 longer than twice carapace length. Male pleonite 1 and 2 of equal width, pleonite 3 less than 1.5 times as wide as pleonite 4. Maximum cl. 12 mm. Subtidal (on floating semi-submerged structures). Central Indo-Pacific (Singapore). 1 species (Mendoza and Ng 2011).

Itampolus Serène & Peyrot-Clausade, 1977

Lentilumnus Galil & Takeda, 1988

Diagnosis. Carapace glabrous or at most with few fine scattered setae; upper surface smooth, without ridges; suborbital margin

Diagnosis. Carapace glabrous or at most with few fine scattered setae; upper surface smooth, without ridges; anterolateral margin

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Marine Decapod Crustacea

with 3 blunt or sharp teeth. Thoracic sternite 8 of male completely concealed by pleonite 2. Eyestalk cylindrical, longer than cornea. Cheliped palm outer surface entirely covered with sharp granules. Pereopod 3 shorter than twice carapace length. Pereopod 5 ischium, merus unarmed along lower margin. Male telson extending beyond midlength of sternite 4. Maximum cl. 6 mm. Subtidal, shelf (18–48 m). Central Indo-Pacific. 2 species (Galil and Takeda 1988).

Lobopilumnus A. Milne-Edwards, 1880 Diagnosis. Carapace rounded-subquadrate; front exceeding onethird carapace width, with anteriorly directed fringe of long setae; regions protuberant, forming large, naked, granular nodules, especially on anterior half; with close pile of short setae between and around nodules; anterolateral margin with 3 acute triangular teeth. Cheliped palm outer surface both chelipeds tuberculate or spinose, setose. Pereopod 3 longer than twice carapace length. Pereopod 5 ischium, merus unarmed along lower margin. Male pleonite 1 wider than pleonite 2, thoracic sternite 8 not exposed on each side of pleonite 2, pleonite 3 less than 1.5 times as wide as pleonite 4. Maximum cl. 23 mm. Intertidal–shelf (0–51 m). Tropical W Atlantic. 1 species (Rathbun 1930: rediagnosis, figures; Williams 1984: figure).

Lophopilumnus Miers, 1886 Diagnosis. Carapace anterolateral margin with 4 lobes or teeth. Pereopods 2–5 meri upper margin and carpi upper margins with thin, upraised crest. Maximum cl. 43 mm. Intertidal–shelf (0–46 m). Central Indo-Pacific. 3 species (Davie 1988: rediagnosis, 3 species).

Lophoplax Tesch, 1918 Diagnosis. Carapace irregularly ornamented with granules and prominent glabrous nodules or ‘islands’ in addition to long setae and tomentose patches; anterolateral margin with 3 prominent blunt or sharp teeth. Maximum cl. 9 mm. Intertidal–shelf (0–100 m). Indo-Pacific. 6 species (figures: Takeda 1977; Takeda and Kurata 1984; Deb 1989: figure, as Myopilumnus, junior synonym; Pseudocryptocoeloma symmetrinudus Edmondson, 1951, is tentatively placed here).

Nanopilumnus Takeda, 1974 Diagnosis. Carapace rounded-subquadrate, widest across third anterolateral tooth, fourth tooth smaller than third; front exceeding one-third carapace width, anterior margin straight, with anteriorly directed fringe of long setae; anterolateral margin with 3 stout, sometimes lobate teeth. Cheliped palm outer surface both chelipeds tuberculate or spinose, setose. Pereopod 3 longer than twice carapace length. Pereopod 5 ischium, merus unarmed along lower margin. Male pleonite 1 wider than pleonite 2, thoracic sternite 8 not exposed on each side of pleonite 2, pleonite 3 less than 1.5 times as wide as pleonite 4. Male telson extending beyond midlength of sternite 4. Maximum cl. 6 mm.

Subtidal, shelf (1–211 m). Western and Central Indo-Pacific. 5 species (Takeda 1974: diagnosis; Takeda and Komatsu 2018: comparison of some species).

Neoactumnus Sakai, 1965 Diagnosis. Front entire; carapace entirely covered with fine, short, velvet-like tomentum; surface smooth; posterolateral surface gently convex, not sunken, not accommodating folded pereopod 5; anterolateral margin with 3 teeth, 2 broad, last smallest. Maximum cl. 11 mm. Shelf (25–51 m). Temperate Northern Pacific, Central IndoPacific. W Pacific. 2 species (Garth and Kim 1983: figure; Lee et al. 2008: species comparison).

Peleianus Serène, 1971 Diagnosis. Carapace rounded-subquadrate; front with median and lateral notches; glabrous or at most with few fine scattered setae; upper surface smooth, without ridges; anterolateral margin with 2 anteriorly curved spines. Pereopod 3 longer than twice carapace length. Gonopod 1 straight. Maximum cl. 5 mm. Subtidal, shelf (17–103 m). Central Indo-Pacific (Philippines, Indonesia). 1 species (Serène 1971: diagnosis; Serène and Umali 1972: description, figures).

Pilumnopeus A. Milne-Edwards, 1867 Diagnosis. Carapace rounded-subquadrate; front exceeding onethird carapace width, with anteriorly directed fringe of long setae; anterolateral margin with 4 lobes or teeth. Cheliped palm outer surface of one or both chelipeds smooth or microscopically granular, glabrous. Pereopod 3 longer than twice carapace length. Male pleonite 1 wider than pleonite 2, thoracic sternite 8 not exposed on each side of pleonite 2, pleonite 3 less than 1.5 times as wide as pleonite 4. Male telson not reaching midlength of sternite 4. Maximum cl. 20 mm (Pl. 60d). Intertidal. Tropical E Atlantic, Western and Central IndoPacific, Temperate SE Australia. 9 species (Dai and Yang 1991: key to 3 species; Davie 1989a: rediagnosis, 2 species).

Pilumnus Leach, 1816 Diagnosis. Carapace transversely ovate to subhexagonal; anterolateral margin with anterolateral teeth separated by short intervals, not immediately abutting each other, spiniform or blunt-subconical, occasionally appearing as low angular lobes. Pereopod 3 shorter than twice carapace length. Gonopod 1 sinuous, apex recurved, hooked. Maximum cl. 34 mm (Pl. 60e–k). Intertidal–slope (0–500 m). Cosmopolitan in temperate and tropical seas. 145 species. Synoptic keys are not available for Pilumnus, but regional keys include: Australia (Poore 2004; Rathbun 1923), Americas (Rathbun 1930), South Africa (Barnard 1950), West Africa (Monod 1956), north-east Atlantic (Zariquiey Alvarez 1968), Japan (Sakai 1976), New Zealand (McLay 1988), China (Dai and Yang 1991) and Persian Gulf (Naderloo 2017).

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Priapipilumnus Davie, 1989 Diagnosis. Carapace anterolateral margin with 3 sharp tuberculate teeth behind exorbital angle. Male telson ~1.4 times as long as wide, with concave lateral margins. Gonopod 1 proximal half strongly inflated, swollen, distal half slender, straight, not apically hooked. Maximum cl. 10 mm. Shelf (128–132 m). Central Indo-Pacific (N Australia). 1 species (Davie 1989b).

Rhizopoides Ng, 1987 Diagnosis. Carapace rounded-subquadrate; front with anteriorly directed fringe of long setae; anterolateral margin with 3 broad, closely spaced teeth, bases abutting. Cheliped palm outer surface of major cheliped smooth, glabrous, of minor cheliped granular, setose. Pereopod 3 longer than twice carapace length. Male pleonite 3 ~3 times as wide as pleonite 4. Maximum cl. 5 mm. Intertidal. Central Indo-Pacific (Singapore). 1 species (Ng 1985: figures; 1987: diagnosis).

Serenepilumnus Türkay & Schuhmacher, 1985 Diagnosis. Carapace rounded-subquadrate, widest across fourth (last) anterolateral tooth, fourth tooth equal to or larger than third tooth; front exceeding one-third carapace width, with anteriorly directed fringe of long setae; anterolateral margin with 4 closely spaced teeth, first and fourth triangular, second and third more truncate, all with secondary tubercles. Cheliped palm outer surface both chelipeds tuberculate or spinose, setose. Pereopod 3 longer than twice carapace length. Pereopod 5 ischium, merus unarmed along lower margin. Male pleonite 1 wider than pleonite 2, thoracic sternite 8 not exposed on each side of pleonite 2, pleonite 3 less than 1.5 times as wide as pleonite 4. Maximum cl. 11 mm. Subtidal. Central Indo-Pacific. 5 species (Serène 1971: comparison of 3 species, as Leopoldius)

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propodi upper margins spinose. Pereopod 3 longer than twice carapace length. Male telson longer than wide. Maximum cl. 10 mm. Shelf (54–71 m). Central Indo-Pacific (Australia, New Caledonia). 1 species (Davie 1989b).

Vellumnus Ng, 2010 Diagnosis. Carapace transversely ovate; setation dense, short, forming vermiform or labyrinth-like pattern partially corresponding to underlying curved ridges on carapace surface; anterolateral margin with 3 subequal teeth, sometimes appearing sublobiform. Maximum cl. 15 mm. Subtidal, shelf (1–106 m). Central Indo-Pacific. 5 species (Ng 2010: diagnosis, figures, species list).

Viaderiana Ward, 1942 Diagnosis. Carapace rounded-subquadrate; front with anteriorly directed fringe of long setae; anterolateral margin with 3 acute triangular teeth; exorbital angle forming small spine. Pereopods 2–5 meri upper margin usually spinose. Pereopod 3 longer than twice carapace length. Male pleonite 1 little wider than pleonite 2. Maximum cl. 17 mm. Subtidal, shelf (8–62 m). Western and Central Indo-Pacific. 16 species (Ng and Dai 1997: figure; Serène 1971: diagnosis, 2 species).

Xestopilumnus Ng & Dai, 1997 Diagnosis. Carapace rounded-subquadrate; front entire, without median or lateral notches; glabrous or at most with few fine scattered setae; upper surface smooth, without ridges; anterolateral margin with low, blunt, granular lobes. Pereopod 3 longer than twice carapace length. Gonopod 1 apex strongly recurved, hooked. Maximum cl. 17 mm. Bathyal (2736 m). Central Indo-Pacific (Hong Kong). 1 species (Ng and Dai 1997).

Serenolumnus Galil & Takeda, 1988

Xlumnus Galil & Takeda, 1988

Diagnosis. Carapace glabrous or at most with few fine scattered setae; upper surface smooth, without ridges; suborbital margin straight, inner margin not produced forwards; anterolateral margin with 3 teeth, 2 broad, last smallest. Thoracic sternite 8 of male completely concealed by pleonite 2. Eyestalk as long as cornea. Antennal flagellum half as long as orbit width. Pereopod 3 shorter than twice carapace length. Pereopod 5 ischium, merus unarmed along lower margin. Male telson extending beyond midlength of sternite 4. Maximum cl. 5 mm. Intertidal–shelf (0–78 m). Central Indo-Pacific. 1 species (Galil and Takeda 1988: key to 10 species including Serenolumnus, Lentilumnus and Xlumnus).

Diagnosis. Carapace glabrous or at most with few fine scattered setae; upper surface smooth, without ridges; anterolateral margin with 3 acute triangular teeth. Thoracic sternite 8 of male completely concealed by pleonite 2. Eyestalk pyriform, longer than and with distinct constriction at base of cornea. Cheliped palm outer surface with sharp granules proximally, smooth distally. Pereopod 3 shorter than twice carapace length. Pereopod 5 ischium, merus unarmed along lower margin. Male telson extending beyond midlength of sternite 4. Maximum cl. 5 mm. Subtidal. Central Indo-Pacific (Vietnam). 1 species (Galil and Takeda 1988: key to 10 species including Serenolumnus, Lentilumnus and Xlumnus; Serène 1971: figure of type species).

Takedana Davie, 1989 Diagnosis. Carapace rounded-subquadrate; anterolateral margin with small, sharp anterolateral teeth. Cheliped palm outer surface with long, mushroom- or thorn-like spines. Pereopods 2–5 carpi,

Subfamily Rhizopinae Stimpson, 1858 Rhizopines generally have a rounded-quadrate carapace with obsolescent (or absent) anterolateral teeth, and a more

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proximally widened pleon than pilumnines, albeit with male pleonite 2 narrowed such that sternite 8 is visible on each side. Some rhizopines occur in rocky and coral reef crevices, but most burrow in turbid, muddy habitats. Some have reduced eyes, presumably as a consequence of living in low light environments. Števčić (2005) proposed Xenophthalmodinae for Xenophthalmus, a genus previously considered a rhizopine, a subfamily accepted by Ng et al. (2008) and Davie et al. (2015). The elongated gonopods 1 that extend anteriorly beyond the pleon, believed to be diagnostic of xenophthalmodines, however, are not consistent features of all members of Xenophthalmus, including the type species X. moebii Richters, 1880. We are unable to distinguish Rhizopinae from Xenophthalmodinae and consider the two to be synonymous. Although the subfamily Rhizopinae is probably an unnatural group, it is retained here pending revision of the systematics of the family. Ng (1987) remarked on the definition of several genera but did not illustrate them. The composition of Rhizopinae largely follows Ng et  al. (2008) but several genera placed there are herein pragmatically placed in Pilumninae: Lophoplax and Rhizopoides, with well developed anterolateral teeth on the carapace and moved for

practical reasons; and Itampolus and Peleianus, which are morphologically similar to Glabropilumnus. Heteropilumnus has been variously regarded as a rhizopine (Davie 2002; Ng 1987; Števčić 2005) and pilumnine (Ng et al. 2008), and is acknowledged as heterogeneous and requiring revision (Ng et al. 2018; Ng and Davie 1991). Heteropilumnus is included in Pilumninae but because of morphological heterogeneity, appears twice in the key above, once among the pilumnines and once among the rhizopines. Diagnosis. Carapace anterolateral margin obtusely convex, with or without low teeth, at most weakly indicated. Thoracic sternite 8 of male visible on each side of pleonite 2. Antennules folding transversely. Cheliped fingers without gape, occlusal margins at most sparsely setose. Pereopods 2–5 without dactylus-propodal lock. Implicit rhizopine generic attributes. Carapace transversely ovate. Epistome lower medial margin not lobed. Eyestalk mobile, cornea well developed. Camptoplax Miers, 1884 Diagnosis. Carapace subhexagonal, glabrous; dorsal surface with 2 low transverse ridges; anterolateral margin shorter than posterolateral margin, with 3 low teeth behind exorbital angle;

Fig. 14.112.  Pilumnidae, Rhizopinae and Pilumninae. a, Zehntneriana serrata Ng & Lin, 2015. Carapace (setae not shown): b, Camptoplax; c, Cryptolutea; d, Luteocarcinus; e, Mertonia; f, Paranotonyx; g, Paraselwynia; h, Pronotonyx; i, Lophoplax; j, Pseudolitochira; k, Rhizopa; l, Ser; m, Typhlocarcinops; n, Typhlocarcinus; o, Xenophthalmodes.

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Fig. 14.113.  Pilumnidae, Rhizopinae. Front, eyes, antennules, antennae, epistome (anterior view): a, Luteocarcinus; b, Paraselwynia; c, Typhlocarcinops. Front, eyes, antennules, antennae (ventral view): d, Cryptocoeloma. Left half of pleonites 1–4, thoracic sternites 7, 8, coxa 5 (numbered on f): e, Cryptolutea; f, Heteropilumnus; g, Rhizopa; h, Typhlocarcinops; i, Typhlocarcinus; j, Ser. Maxilliped 3: k, Ceratoplax; l, Luteocarcinus; m, Mertonia; n, Paraselwynia; o, Xenophthalmodes. Chela, outer face: p, Pseudolitochira. Pereopod 5: q, Cryptolutea.

posterolateral margins convergent. Maxilliped 3 merus rectangular distolaterally. Male pleonite 3 reaching coxa 5. Maximum cl. 6 mm (Pl. 59i). Subtidal, shelf (13–52 m). Central Indo-Pacific. 1 species (Davie 1993).

Ceratoplax Stimpson, 1858 Diagnosis. Maxilliped 3 merus produced distolaterally; exopod reaching to distal end of merus; merus wider than ischium at midlength. Male pleonite 1 and pleonite 3 not reaching coxa 5. Maximum cl. 15 mm (Pl. 59j). Subtidal–slope (2–292 m). Western and Central Indo-Pacific, Temperate Australia. 9 species (Ng and Clark 2015: 3 species compared; Poore 2004: key to 3 species).

Cryptocoeloma Miers, 1884 Diagnosis. Orbit below anterior carapace margin, not visible in dorsal view. Maxilliped 3 merus rounded distolaterally. Male pleonite 3 reaching coxa 5. Maximum cl. 7 mm. Intertidal, shelf (0–52 m). Central Indo-Pacific (N Australia, Indonesia). 1 species (Ng 1989).

Cryptolutea Ward, 1936 Diagnosis. Maxilliped 3 merus rectangular distolaterally. Pereopods 2–5 coxae with denticulate plate partially overlapping ischium. Male pleonite 1 broad reaching coxa 5; pleonite 3 not reaching coxa 5. Maximum cl. 16 mm. Intertidal–shelf (0–112 m). Western and Central Indo-Pacific. 5 species (Ng and Davie 1991: rediagnosis, figures; Ng and Huang 2001: figures).

Luteocarcinus Ng, 1990 Diagnosis. Carapace fronto-orbital width two-thirds carapace width; surface glabrous. Epistome lower medial margin cut into 2 narrow rectangular lobes. Maxilliped 3 merus rounded distolaterally; exopod reaching to midlength of merus; merus narrower than ischium at midlength. Male pleonite 1 and pleonite 3 not reaching coxa 5. Maximum cl. 5 mm. Intertidal. Central Indo-Pacific (Peninsular Malaysia). 1 species (Ng 1990).

Mertonia Laurie, 1906 Diagnosis. Carapace semicircular, lateral margins distinctly divergent posteriorly. Antennal flagellum densely plumose. Maxilliped 3 merus produced distolaterally. Maximum cl. 7 mm. Subtidal, shelf (13–109 m). Western and Central Indo-Pacific. 2 species (Laurie 1906).

Paranotonyx Nobili, 1906 Diagnosis. Carapace transversely ovate, margin fully circumscribed by continuous, narrow ridge. Maxilliped 3 merus produced distolaterally. Maximum cl. 5 mm. Subtidal. Western Indo-Pacific (Red Sea). 1 species (Nobili 1906: figure).

Paraselwynia Tesch, 1918 Diagnosis. Carapace anterolateral margins smooth, unarmed, not lined with granules. Maxilliped 3 merus rectangular distolaterally; exopod reaching to midlength of merus; merus narrower

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than ischium at midlength. Male pleonite 1 and pleonite 3 not reaching coxa 5. Maximum cl. 8 mm. Subtidal, shelf (6–31 m). Western and Central Indo-Pacific. 1 species (Tesch 1918: figures).

distolaterally. Male pleonite 1 broad reaching coxa 5; pleonite 3 not reaching coxa 5 almost twice width of pleonite 2. Maximum cl. 15 mm. Intertidal–slope (0–601 m). Western and Central Indo-Pacific. 22 species (Ng and Rahayu 2020b: rediagnosis, key to 22 species).

Pronotonyx Ward, 1936 Diagnosis. Male pleonite 1 broad, reaching coxa 5; pleonite 3 not reaching coxa 5, less than 1.5 times width of pleonite 2. Maximum cl. 10 mm. Subtidal, shelf (12–78 m). Central Indo-Pacific (N Australia). 1 species (Miers 1884a: figures of type species; Ward 1936: diagnosis, poor figures).

Pseudocryptocoeloma Ward, 1936

Typhlocarcinus Stimpson, 1858 Diagnosis. Carapace anterolateral margin irregular, lobate, dentate or lined with granules. Maxilliped 3 merus rounded distolaterally. Male pleonite 1 and pleonite 3 not reaching coxa 5. Maximum cl. 14 mm. Subtidal, shelf (10–146 m). Western and Central Indo-Pacific. 8 species (Ng et al. 2017: discussion of species; Serène 1964: key to 6 species).

Diagnosis. Carapace surface smooth, polished, densely setose anteriorly; anterolateral margin with low obtuse lobes, shorter than posterolateral margin; posterolateral margins convergent. Cheliped palms smooth, densely setose dorsally. Maxilliped 3 merus rounded distolaterally. Male pleonite 3 reaching coxa 5. Maximum cl. 8 mm. Intertidal. Central Indo-West Pacific (N Australia). 1 species (Ward 1936: description and figures).

Diagnosis. Carapace semicircular, lateral margins distinctly divergent posteriorly. Antennal flagellum slender, sparsely setose. Maxilliped 3 merus rounded distolaterally. Maximum cl. 12 mm. Intertidal–shelf (15–28 m). Western and Central Indo-Pacific. 5 species (Hsueh and Ng 2008: discussion of species; Serène and Soh 1976: key to 4 species).

Pseudolitochira Ward, 1942

Zehntneriana Ng & Takeda, 2010

Diagnosis. Carapace surface smooth, regions indistinct or absent, finely setose. Maxilliped 3 merus rectangular distolaterally. Cheliped palm outer surface coarsely granular, setose. Male pleonite 3 reaching coxa 5. Maximum cl. 9 mm. Intertidal, subtidal (0–30 m). Indo-Pacific. 4 species (Miers 1884b: figure of type species; Ng et  al. 2021: redescription; Ward 1942: diagnosis).

Diagnosis. Carapace surface smooth, regions indistinct or absent, glabrous or setose. Maxilliped 3 merus rectangular distolaterally. Male pleonite 3 reaching coxa 5. Cheliped palm outer surface smooth or finely granulate, with or without setae. Maximum cl. 6 mm. Intertidal–shelf (0–123 m). Central Indo-Pacific. 6 species (Lee et  al. 2015: species groups; Ng and Lin 2015: species compared; Takeda 1972: diagnosis as Zehntneria, a preoccupied name).

Rhizopa Stimpson, 1858 Diagnosis. Carapace fronto-orbital width half carapace width; surface finely setose. Epistome lower medial margin cut into 2 narrow rectangular lobes. Maxilliped 3 merus rounded distolaterally; exopod reaching to distal end of merus; merus wider than ischium at midlength. Male pleonite 1 and pleonite 3 not reaching coxa 5. Maximum cl. 7 mm. Subtidal (5–16 m). Central Indo-Pacific. 1 species (Ng 1987: rediagnosis, figures).

Ser Rathbun, 1931 Diagnosis. Carapace quadrate, posterolateral margins subparallel. Male pleonite 3 reaching coxa 5. Maximum cl. 17 mm. Intertidal. Central Indo-Pacific (China). 1 species (Rathbun 1931).

Typhlocarcinops Rathbun, 1909 Diagnosis. Eyestalk usually fixed, cornea reduced, sometimes pigmented. Maxilliped 3 merus rounded or bluntly angular

Xenophthalmodes Richters, 1880

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Ng PKL, Clark PF (2008) A revision of Latopilumnus Türkay & Schuhmacher, 1985, and Aniptumnus Ng, 2000 (Crustacea: Decapoda: Brachyura: Pilumnidae) with larval comparisons. Journal of Natural History 42, 885–912. doi:10.1080/00222930701869422 Ng PKL, Clark PF (2015) Ceratoplax margarita n. sp., a new rhizopine crab (Crustacea: Brachyura: Pilumnidae) from Papua New Guinea, with rediagnoses of C. truncatifrons Rathbun, 1914, and C. fulgida Rathbun, 1914. Zoosystema 37, 323–331. doi:10.5252/z2015n2a2 Ng PKL, Dai AY (1997) The Indo-Pacific Pilumnidae IX. Description of a new genus and new species (Crustacea: Decapoda: Brachyura) from Hong Kong. Asian Marine Biology 13, 37–44. Ng PKL, Davie PJF (1991) The Indo-Pacific Pilumnidae VII. Notes on Heteropilumnus sasekumari (Serène, 1971) and Cryptolutea Ward, 1936 (Crustacea: Decapoda: Brachyura). Memoirs of the Queensland Museum 30, 517–524. Ng PKL, Huang J-F (2001) The Indo-Pacific Pilumnidae XV. On two rare species of Rhizopinae (Decapoda, Brachyura) from Taiwan. Crustaceana 74, 1379–1385. doi:10.1163/15685400152885309 Ng PKL, Lin CW (2015) Zehntneriana serrata n. sp., a new species of pilumnid crab from southern Taiwan (Crustacea, Decapoda, Brachyura). Zootaxa 3915, 263–271. doi:10.11646/zootaxa.3915.2.5 Ng PKL, Rahayu DL (2020a) A new genus and species of pilumnid crab (Decapoda: Brachyura: Pilumnidae) symbiotic with the sponge Callyspongia Duchassaing & Michelotti, 1864 (Porifera: Demospongiae: Callyspongiidae) from Lombok, Indonesia; the identity of Pseudactumnus pestae Balss, 1933; and a review of symbiosis in the Pilumnidae. Journal of Crustacean Biology 40, 918–932. doi:10.1093/ jcbiol/ruaa042 Ng PKL, Rahayu DL (2020b) A synopsis of Typhlocarcinops Rathbun, 1909 (Crustacea: Decapoda: Brachyura: Pilumnidae), with descriptions of nine new species from the Indo-West Pacific. Zootaxa 4788, 1–100. doi:10.11646/zootaxa.4788.1.1 Ng PKL, Tan LWH (1984) The Indo-Pacific Pilumnidae I. Description of four new species of the genus Pilumnus Leach,1815, and definition of a new genus, Bathypilumnus. Journal of the Singapore National Academy of Science 13, 13–19. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Ng PKL, Devi S, Kumar AB (2017) Typhlocarcinus kerala, a new species of rhizopine crab from southwestern India, and the identity of T. craterifer Rathbun, 1914 (Crustacea: Brachyura: Pilumnidae). Zootaxa 4272, 131–141. doi:10.11646/zootaxa.4272.1.7 Ng PKL, Abdelsalam KM, Mona MH, Nour Eldeen MF (2018) A synopsis of Eurycarcinus A. Milne-Edwards, 1867 (Decapoda, Brachyura, Pilumnidae). Crustaceana 91, 471–487. doi:10.1163/15685403-00003775 Ng PKL, Clark PF, Clark B, Kamanli SA (2021) Pseudolitochira integra (Miers, 1884) (Crustacea: Brachyura: Pilumnidae): redescribed and illustrated from micro-ct scanning the type female. Zootaxa 4969, 377–391. doi:10.11646/zootaxa.4969.2.9 Nobili G (1906) Faune carcinologique de la Mer Rouge. Décapodes et stomatopodes. Annales des Sciences Naturelles, 9e série 4, 1–347, pls 1–11. Ozcan T, Katagan T, Ng PKL (2010) First record of Eurycarcinus integrifrons De Man, 1879 (Decapoda, Piluminidae) from the Mediterranean Sea. Crustaceana 83, 507–510. doi:10.1163/001121610X492139 Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Rathbun MJ (1923) Report on the crabs obtained by the F.I.S. “Endeavour” on the coasts of Queensland, New South Wales, Victoria, South Australia and Tasmania. Report on the Brachyrhyncha, Oxystomata

and Dromiacea. Biological Results of the Fishing Experiments carried out by the F.I.S. “Endeavour”, 1909–14 5, 95–156, pls 16–42. Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Rathbun MJ (1931) New and rare Chinese crabs. Lingnan Science Journal 8, 75–104, pls 5–15. Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo. Schmitt WL, McCain JC, Davidson E (1973) Decapoda I. Brachyura I. Fam. Pinnotheridae. Crustaceorum Catalogus 3, 1–160. Serène R (1964) Goneplacidae et Pinnotheridae. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914–1916 80, 181–282, pls 16–24. Serène R (1971) Observations préliminaires sur des brachyoures nouveaux ou mal connus du Sud-Est Asiatique (Crustacea Decapoda). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 42, 903–918. Serène R, Peyrot-Clausade M (1977) Description d’Itampolus peresi gen. et sp. nov. (Decapoda, Brachyura, Xanthidae). Crustaceana 32, 286–289. doi:10.1163/156854077X00764 Serène R, Soh CL (1976) Brachyura collected during the Thai-Danish Expedition (1966). Phuket Marine Biological Center Research Bulletin 12, 1–37, figs 1–28, pls 1–8. Serène R, Umali AF (1972) The family Raninidae and other new and rare species of brachyuran decapods from the Philippines and adjacent regions. Philippine Journal of Science 99, 21–105. Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Števčić Z, Castro P, Gore RH (1988) Re-establishment of the family Eumedonidae Dana, 1853 (Crustacea: Decapoda: Brachyura). Journal of Natural History 22, 1301–1324. doi:10.1080/00222938800770801 Takeda M (1972) Systematic status of Ceratoplax villosa Zehntner and some related species (Crustacea, Decapoda, Brachyura). Proceedings of the Japanese Society of Systematic Zoology 8, 31–41. Takeda M (1974) Pilumnid crabs of the family Xanthidae from the West Pacific. V. Definition of a new genus, with description of its type-species. Bulletin of the National Science Museum, Tokyo 17, 215–219. Takeda M (1977) Rediscovery of Lophoplax sculpta (Stimpson, 1858) (Crustacea, Decapoda, Brachyura). Zoological Magazine, Tokyo 86, 120–123. Takeda M (1989) Shallow-water crabs from the Oshima Passage between Amami-Oshima and Kakeroma-jima Islands, the Northern Ryukyu Islands. Memoirs of The National Science Museum, Tokyo 22, 135–184, pl. 4. Takeda M, Komatsu H (2017) Two new species of the genus Actumnus Dana, 1851 (Decapoda, Brachyura, Pilumnidae) from the Ryukyu Islands, southwest Japan. Crustaceana 90, 1251–1265. doi:10.1163/​ 15685403-00003686 Takeda M, Komatsu H (2018) Offshore crabs of the family Xanthidae and some related families (Crustacea, Decapoda, Brachyura) from the Ogasawara Islands, Japan. Memoirs of the National Museum of Natural Sciences, Tokyo 52, 153–189. Takeda M, Kurata Y (1984) Crabs of the Ogasawara Islands. VII. Third report on the species obtained from stomachs of fishes. Bulletin of the National Science Museum, Ser. Zoology 10, 195–202. Takeda M, Ng PKL (1997) The Indo-Pacific Pilumnidae. XI. An unusual new species of Pilumnus from Japan. Bulletin of the National Science Museum, Ser. Zoology 23, 185–190. Tesch JJ (1918) The Decapoda Brachyura of the Siboga Expedition. II Goneplacidae and Pinnotheridae. Siboga-Expéditie 39c1, 149–296, pls 7–18. Türkay M, Schuhmacher H (1985) Latopilumnus tubicolus n. gen. n. sp., eine neue korallenassoziierte Krabbe, die die Bildung einer Wohnhöhle induziert (Crustacea: Decapoda: Pilumnidae). Senckenbergiana Maritima 17, 55–63.

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Vandenspiegel D, Ovaere A, Massin C (1992) On the association between the crab Hapalonotus reticulatus (Crustacea, Brachyura, Eumedonidae) and the sea cucumber Holothuria (Metriatyla) scabra (Echinodermata, Holothuridae). Bulletin de l’Institut Royal des Sciences Naturelles de Belgique. Biologie 62, 167–177. Vázquez-Bader AR, Gracia A (1995) A new genus and species of pilumnid (Crustacea, Decapoda, Brachyura) from the Gulf of Mexico. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série 17, 259–272. Ward M (1936) Crustacea Brachyura from the coasts of Queensland. Memoirs of Queensland Museum 11, 1–13, pls 1–3. Ward M (1942) Notes on the Crustacea of the Desjardins Museum, Mauritius Institute, with descriptions of new genera and species. Mauritius Institute Bulletin 2, 49–113, pls 5, 6. Williams AB (1984) Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C. Zariquiey Alvarez R (1968) Crustáceos decápodos ibéricos. Investigaciones Pesqueras 32, 1–510.

Tanaochelidae Ng & Clark, 2000 Figure 14.114 The family comprises one genus, Tanaocheles, and two species of Indo-West Pacific coral dwelling crabs (Kropp 1984). Tanaochelids superficially resemble trapeziids and some xanthids, especially species of Chloridiella but are clearly pilumnoids based on larval, male pleonal and gonopod morphology (Ng and Clark 2000). Diagnosis. Carapace transversely ovate, surface smooth, glabrous, regions weakly indicated. At least one cheliped long and slender, twice carapace length or more; fingers spoon-tipped. Male pleon tapering beyond pleonite 3. Tanaocheles Kropp, 1984 Intertidal–shelf (0–27 m; in live coral). Western and Central IndoPacific. Maximum cl. 6 mm. 2 species (Ng and Clark 2000: redescription, species separation). References Kropp RK (1984) Tanaocheles stenochilus, a new genus and species of crab from Guam, Mariana islands (Brachyura: Xanthidae). Proceedings of the Biological Society of Washington 97, 744–747. Ng PKL, Clark PF (2000) The Indo-Pacific Pilumnidae XII. On the familial placement of Chlorodiella bidentata (Nobili, 1901) and Tanaocheles stenochilus Kropp, 1984 using adult and larval characters with the establishment of a new subfamily, Tanaochelinae (Crustacea: Decapoda: Brachyura). Journal of Natural History 34, 207–245. doi:10.1080/002229300299615

Portunoidea Rafinesque, 1815 Several groups of crabs have natatory adaptations, but the most efficient and effective swimmers are among the portunoids (Hartnoll 1971). The portunoids are best known for the large swimming commercial species, particularly those

Fig. 14.114.  Tanaochelidae, Tanaocheles. a, carapace, chelipeds; b, male pleon.

of the genera Scylla, Portunus, Callinectes and Charybdis, all of which have a flattened, paddle-like pereopod 5 well adapted for swimming. These crabs are typically aggressive, agile, and laterally streamlined. The superfamily, however, encompasses not only highly accomplished swimmers, but also basal non-swimming forms with a simple pereopod 5, along with a range of intermediates. Some specialised or commensal portunoids have secondarily lost the ability to swim and do not immediately resemble stereotypical swimming crabs (Evans 2018). Portunoids are most diverse in the shallow tropics, but range into cold-temperate waters, and deep, bathyal habitats. The classification of the swimming crabs at all taxonomic levels has changed considerably in recent decades and the current system is largely an expansion of that of William Stephenson (e.g. Stephenson 1961, 1972; Stephenson and Campbell 1959, 1960; Stephenson and Hudson 1957; Stephenson et al. 1957), Ng et al. (2008) and De Grave et al. (2009). Recent morphological and molecular phylogenetic considerations of Portunoidea (Evans 2018; Karasawa et al. 2008; Schubart and Reuschel 2009; Spiridonov 2020; Spiridonov et al. 2014) clarified the taxonomic scope of the superfamily (with possible ambiguity over whether Brusiniidae should be included; Evans 2018), but views differ on the degree of inclusiveness of some families. Spiridonov et al. (2014) and Davie et al. (2015) recognised nine portunoid families distributed in three broad clades of families (a geryonid group, a carcinid group and a portunid group). Evans (2018) and Spiridonov (2020) then adopted a simplified system by subsuming Ovalipidae into Geryonidae, and subsuming Pirimelidae, Polybiidae, Thiidae, Parathranites and Coelocarcinus in an expanded Carcinidae. We regard these expanded concepts of Geryonidae and Carcinidae,

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however, as having little practical advantage, either taxonomically (because of the difficulty in collectively diagnosing these diverse assemblages, especially the latter) or in harmonizing the classification with phylogeny (owing to current ambiguities in phylogenetic resolution). Therefore, pending robust phylogenetic resolution of portunoid interrelationships, we substantially follow the nine-family classification of Spiridonov et al. (2014) and Davie et al. (2015), but with several modifications: Nautilocorystinae (for Nautilocorystes) is removed from Thiidae and treated as a separate family Nautilocorystidae; Nectocarcinus, Echinolatus and Coenophthalmus are removed from Ovalipidae or Polybiidae and included in Geryonidae. Parathranites and Coelocarcinus, regarded as carcinids by Evans (2018)

and Spiridonov (2020), are herein treated as polybiids and portunids, respectively. Diagnosis. Carapace longer than wide to wider than long. Mouthfield quadrate. Antennules folding transverselyobliquely, rarely longitudinally (Nautilocorystidae). Antennae longer than cornea. Maxilliped 3 merus rounded to subquadrate. Pereopod 5 a walking leg with dactylus slender, lanceolate, styliform, or as a swimming leg, with dactylus and propodus flattened, broadened, ovate to fusiform, forming swimming paddle. Male pleonite 3 usually with transverse ridge; if transverse ridge absent, pereopod 5 paddle-like or narrowly crescentic but flattened (Libystes, Kume). Female gonopores (vulvae) not exposed, fully concealed by pleon.

Key to families of Portunoidea 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. –

Carapace ‘thumb-nail’ shaped, dorsally smooth, surface convex transversely, straight longitudinally; entire margin with continuous fringe of long golden setae (Fig. 14.130; Pl. 62l)�������������������������������������������������������������Thiidae Carapace variously shaped and ornamented but convex both transversely and longitudinally; margins glabrous or variously setose, but without continuous fringe of long golden setae�����������������������������������������������������������2 Carapace longer than wide, pyriform�������������������������������������������������������������������������������������������������������������������������������������������3 Carapace as wide as or wider than long, subcircular to transversely ovate, fusiform or subhexagonal�����������������������������4 Carapace margins with 3 small spines or low angular notches (excluding outer orbital spine); front with 3 rounded lobes. Antennules folding transversely. Antennal flagellum sparsely setose, shorter than one-quarter carapace length. Pereopod 5 dactylus elongate-ovate, distally rounded (Fig. 14.115)������������ Brusiniidae Carapace margins with 4 prominent spines (excluding outer orbital spine); front divided into 2 angular lobes. Antennules folding longitudinally. Antennal flagellum strongly setose, about half carapace length. Pereopod 5 dactylus spatulate, distally sharp (Fig. 14.118)�������������������������������������������������������Nautilocorystidae Male pleonites 1–6 and telson clearly demarcated, even if pleonites 3–5 immovable (Fig. 14.119c)�����������������������������������5 Male pleonites 3–5 fused, sutures absent or only partially indicated (Fig. 14.123x, y)����������������������������������������������������������6 Stridulatory ridges present on lower surface of cheliped propodi or on pterygostomial surface of carapace (Fig. 14.119b). Male pleon narrow, linguiform, margins weakly tapering, anterior width of pleonite 6 about two-thirds width of pleonite 3 (Fig. 14.119c)��������������������������������������������������������������������������������������������� Ovalipidae Stridulatory ridges absent on carapace and chelipeds. Male pleon triangular, margins strongly tapering, anterior width of pleonite 6 half or less that of pleonite 3 (Fig. 14.117o, p)�����������������������������������������������������Geryonidae Chelipeds longer than longest walking legs (pereopods 2 or 3)������������������������������������������������������������������������������Portunidae Chelipeds shorter than longest walking leg���������������������������������������������������������������������������������������������������������������������������������7 Endostomial ridges long, extending to epistome (Fig. 14.121g, h)��������������������������������������������������������������������������� Polybiidae Endostomial ridges absent or short, not extending to epistome (Fig. 14.116d)����������������������������������������������������������������������8 Pereopod 5 dactylus narrowly lanceolate to spatuliform, flattened, always wider than dactylus of other legs (Fig. 14.116a, f)����������������������������������������������������������������������������������������������������������������������������������������� Carcinidae Pereopod 5 dactylus styliform, similar to that of other legs, not flattened (Fig. 14.120e)���������������������������������Pirimelidae

References Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden.

De Grave S, Pentcheff ND, Ahyong ST, Chan T-Y, Crandall KA et  al. (2009) A classification of living and fossil genera of decapod crustaceans. Raffles Bulletin of Zoology Supplement 21, 1–109. Evans N (2018) Molecular phylogenetics of swimming crabs (Portunoidea Rafinesque, 1815) supports a revised family-level classification and suggests a single derived origin of symbiotic taxa. PeerJ 6, e4260. doi:10.7717/peerj.4260

14 – Brachyura – crabs

Hartnoll RG (1971) The occurrence, methods and significance of swimming in the Brachyura. Animal Behaviour 19, 34–50. doi:10.1016/ S0003-3472(71)80132-X Karasawa H, Schweitzer CE, Feldmann RM (2008) Revision of Portunoidea Rafinesque, 1815 (Decapoda: Brachyura) with emphasis on the fossil genera and families. Journal of Crustacean Biology 28, 82–127. doi:10.1651/07-2882R.1 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Schubart CD, Reuschel S (2009) A proposal for a new classification of Portunoidea and Cancroidea (Brachyura: Heterotremata) based on two independent molecular phylogenies. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 533–549. CRC Press, Boca Raton. Spiridonov VA (2020) An update of phylogenetic reconstructions, classification and morphological characters of extant Portunoidea Rafinesque, 1815 (Decapoda, Brachyura, Heterotremata), with a discussion of their relevance to fossil material. Geologija 63, 133–166. doi:10.5474/ geologija.2020.014 Spiridonov VA, Neretina T, Schepetov D (2014) Morphological characterization and molecular phylogeny of Portunoidea Rafinesque,1815 (Crustacea Brachyura): implications for understanding evolution of swimming capacity and revision of the family-level classification. Zoologischer Anzeiger 253, 404–429. doi:10.1016/j.jcz.2014.03.003 Stephenson W (1961) The Australian portunids (Crustacea: Portunidae) V. Recent collections. Australian Journal of Marine and Freshwater Research 12, 92–128, pls 1–5. Stephenson W (1972) An annotated check list and key to the Indo-WestPacific swimming crabs (Crustacea: Decapoda: Portunidae). Royal Society of New Zealand Bulletin 10, 1–64. Stephenson W, Campbell BM (1959) The Australian portunids (Crustacea: Portunidae). III. The genus Portunus. Australian Journal of Marine and Freshwater Research 10, 84–124, pls 1–5. Stephenson W, Campbell BM (1960) The Australian portunids (Crustacea: Portunidae) IV. Remaining genera. Australian Journal of Marine and Freshwater Research 11, 73–122, pls 1–6. Stephenson W, Hudson JJ (1957) The Australian portunids (Crustacea; Portunidae). I. The genus Thalamita. Australian Journal of Marine and Freshwater Research 8, 312–368, pls 1–10. Stephenson W, Hudson JJ, Campbell BM (1957) The Australian portunids (Crustacea; Portunidae). Australian Journal of Marine and Freshwater Research 8, 491–507. doi:10.1071/MF9570491

Brusiniidae Števčić, 1991 Figure 14.115, Plate 61a Brusiniids are small, elongated, presumably fossorial crabs, occurring mostly in outer shelf waters of the south-western Pacific. They are phylogenetically enigmatic. Recent studies have treated brusiniids as a separate family (Spiridonov 2020; Spiridonov et al. 2014) but their phylogenetic position is no clearer now than when Števčić (1991) first delineated the group. Evans (2018) could not determine a stable position for Brusinia and even questioned its identity as a portunoid. Nevertheless, Sakai (1969) recognised numerous similarities to Benthochascon (now Geryonidae) in placing the first discovered species (B. elongata Sakai, 1969) in that genus, and Karasawa et  al. (2008) recovered a close

689

Fig. 14.115.  Brusiniidae. Brusinia brucei Števčić, 1991.

relationship with Ovalipes (Ovalipidae). Thus, brusiniids are possibly in phylogenetic proximity to the geryonid-ovalipid clade. Diagnosis. Carapace longer than wide, pyriform, glabrous; margins with 3 small spines or low angular notches (excluding exorbital spine); front with 3 rounded lobes. Antennules folding transversely. Antennal flagellum sparsely setose, shorter than one-quarter carapace length. Pereopod 5 as a swimming leg, dactylus elongate-ovate, distally rounded. Brusinia Števčić, 1991 Maximum cl. 14 mm (Pl. 61a). Shelf (35–355 m). Central IndoPacific. 4 species (Crosnier and Moosa 2002: key to species). References Crosnier A, Moosa MK (2002) Trois Portunidae (Crustacea, Decapoda, Brachyura) nouveaux de Polynésie française. Zoosystema 24, 385–399. Evans N (2018) Molecular phylogenetics of swimming crabs (Portunoidea Rafinesque, 1815) supports a revised family-level classification and suggests a single derived origin of symbiotic taxa. PeerJ 6, e4260. doi:10.7717/peerj.4260 Karasawa H, Schweitzer CE, Feldmann RM (2008) Revision of Portunoidea Rafinesque, 1815 (Decapoda: Brachyura) with emphasis on the fossil genera and families. Journal of Crustacean Biology 28, 82–127. doi:10.1651/07-2882R.1 Sakai T (1969) Two new genera and twenty-two new species of crabs from Japan. Proceedings of the Biological Society of Washington 82, 243–280. Spiridonov VA (2020) An update of phylogenetic reconstructions, classification and morphological characters of extant Portunoidea Rafinesque, 1815 (Decapoda, Brachyura, Heterotremata), with a discussion of their relevance to fossil material. Geologija 63, 133–166. doi:10.5474/ geologija.2020.014 Spiridonov VA, Neretina T, Schepetov D (2014) Morphological characterization and molecular phylogeny of Portunoidea Rafinesque,1815 (Crustacea Brachyura): implications for understanding evolution of swimming capacity and revision of the family-level classification. Zoologischer Anzeiger 253, 404–429. doi:10.1016/j.jcz.2014.03.003 Števčić Z (1991) Note on some rare and aberrant Australian crabs. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 8, 121–134.

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Carcinidae MacLeay, 1838 Figure 14.116, Plate 61b, c Although Evans (2018) and Spiridonov (2020) expanded Carcinidae to take in several other related families and seemingly unaligned genera, we restrict Carcinidae to three closely related genera: Carcinus, Portumnus and Xaiva, following Spiridonov et al. (2014) and Davie et al. (2015). The three genera and seven species are native to the eastern Atlantic (including the Mediterranean Sea), of which Carcinus maenas (Linnaeus, 1758) is infamously invasive worldwide (Behrens Yamada 2001). With five anterolateral teeth on the carapace, chelipeds shorter than the longest walking legs, and distally pointed pereopod 5 dactyli, carcinids most closely resemble pirimelids and some polybiids. Carcinids, like pirimelids, however,

lack endostomial ridges and have pereopod 5 at most moderately adapted for swimming, unlike that seen in polybiids. Carcinids usually occur in shallow sand and gravel beach habitats or rocky reefs (Christiansen 1969; d’Udekem d’Acoz 1999). The family was rediagnosed by Evans (2018) but based on a much wider suite of genera, most of which are referred here to other families. Diagnosis. Carapace as wide as or wider than long; with 5 anterolateral teeth; endostomial ridges absent; orbit open mesially, antennal peduncle standing in orbit. Antennules folding transversely or obliquely. Male pleonites 3–5 fused, sutures absent or only partially indicated; pleonite 3 with transverse ridge. Chelipeds shorter than longest walking leg; palm without dorsal spines. Pereopod 5 dactylus narrowly lanceolate to spatuliform, flattened, always wider than dactylus of other legs.

Key to genera of Carcinidae 1. – 2. –

Pereopods 2–5 upper margins of carpus and propodus blunt. Pereopod 5 dactylus lanceolate, slender but slightly wider than pereopod 2–4 dactyli (Fig. 14.116a)������������������������������������������������������������������������������������������Carcinus Pereopods 2–5 upper margins of carpus and propodus cristate. Pereopod 5 sharply spatuliform, distinctly wider than pereopod 2–4 dactyli (Fig. 14.116e, f)������������������������������������������������������������������������������������������������������������������2 Carapace ovate, distinctly wider than long; front irregularly triangular; exorbital angle blunt, weakly produced forwards (Fig. 14.116c)������������������������������������������������������������������������������������������������������������������������������������� Xaiva Carapace ovate-pyriform, about as long as wide; front trilobate; exorbital angle strongly pointed forwards, often to level of front (Fig. 14.116b)������������������������������������������������������������������������������������������������������������������������ Portumnus Carcinus Leach, 1814

Diagnosis. Carapace ovate, distinctly wider than long; front trilobate, produced anteriorly beyond level of exorbital angle. Orbit U-shaped. Pereopods 2–5 upper margins of carpus and propodus

blunt. Pereopod 5 dactylus fusiform, slender, but wider than pereopod 4 dactylus. Maximum cl. 60 mm or more (Pl. 61b, c). Estuarine, intertidal–shelf (0–200 m). Temperate Northern Atlantic (native range). C. maenas (Linnaeus, 1958) introduced to north-western Atlantic, South Africa and southern Australia. 2

Fig. 14.116.  Carcinidae. a, Carcinus maenas (Linnaeus, 1958). Carapace: b, Portumnus; c, Xaiva. d, Orbits, antennules, antennae, epistome, endostome (maxillipeds 3 removed), Carcinus. Pereopods 4, 5: e, f, Xaiva; g, male pleon, Portumnus.

14 – Brachyura – crabs

species (Poore 2004: figure; Zariquiey Alvarez 1968: key to species, C. maenas and C. aestuarii Nardo, 1847 (as C. mediterraneus Czerniavsky, 1884)).

Portumnus Leach, 1814 Diagnosis. Carapace ovate-pyriform, about as long as wide; front trilobate; exorbital angle strongly pointed forwards, often to level of front. Orbit U-shaped. Pereopods 2–5 upper margins of carpus and propodus cristate. Pereopod 5 dactylus sharply spatuliform, distinctly wider than pereopod 2–4 dactyli. Maximum cl. 28 mm. Intertidal–shelf (0–28 m). Temperate Northern E Atlantic, including Mediterranean Sea. 2 species (Forest 1967: comparison of 2 species, one as junior synonym).

Xaiva MacLeay, 1838 Diagnosis. Carapace ovate, distinctly wider than long; front irregularly triangular; exorbital angle blunt, weakly produced forwards. Orbit V-shaped. Pereopods 2–5 upper margins of carpus and propodus cristate. Pereopod 5 dactylus sharply spatuliform, distinctly wider than pereopod 2–4 dactyli. Maximum cl. 24 mm. Intertidal–shelf (0–73 m). Temperate Northern and Tropical E Atlantic, Temperate Southern Africa. 3 species (Emmerson 2016: key to 3 species; Macpherson 1989: figures of 2 species). References Behrens Yamada S (2001) Global invader: the European green crab. Corvallis and Oregon Sea Grant, Oregon. Christiansen ME (1969) Crustacea Decapoda Brachyura. Marine Invertebrates of Scandinavia 2, 1–143. d’Udekem d’Acoz C (1999) Inventaire et distribution des crustacés décapodes de l’Atlantique nord-oriental, de la Méditerranée et des eaux continentales adjacentes au nord de 25°N. Patrimoines Naturels 40, i–x, 1–394, 11 figs, 2 maps. Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. Emmerson WD (2016) A guide to, and checklist for, the Decapoda of Namibia, South Africa and Mozambique. Vol. 2. Cambridge Scholars Publishing, Cambridge. Evans N (2018) Molecular phylogenetics of swimming crabs (Portunoidea Rafinesque, 1815) supports a revised family-level classification and suggests a single derived origin of symbiotic taxa. PeerJ 6, e4260. doi:10.7717/peerj.4260 Forest J (1967) Sur une collection de Crustacés Décapodes de la région de Porto Cesareo. Description de Portumnus pestai sp. nov. Thalassia Salentina 2, 3–29, pls 1–4. Macpherson E (1989) The identity of Xaiva pulchella Macleay, 1838 (Decapoda, Portunidae). Crustaceana 57, 107–110. doi:10.1163/​ 156854089X00428 Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Spiridonov VA (2020) An update of phylogenetic reconstructions, classification and morphological characters of extant Portunoidea Rafinesque,

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1815 (Decapoda, Brachyura, Heterotremata), with a discussion of their relevance to fossil material. Geologija 63, 133–166. doi:10.5474/geologija.​ 2020.014 Spiridonov VA, Neretina T, Schepetov D (2014) Morphological characterization and molecular phylogeny of Portunoidea Rafinesque,1815 (Crustacea Brachyura): implications for understanding evolution of swimming capacity and revision of the family-level classification. Zoologischer Anzeiger 253, 404–429. doi:10.1016/j.jcz.2014.03.003 Zariquiey Alvarez R (1968) Crustáceos decápodos ibéricos. Investigaciones Pesqueras 32, 1–510.

Geryonidae Colosi, 1923 Figure 14.117, Plate 61d, e Most geryonids are large deep-water crabs with pereopods 2–5 acting as walking legs (Geryon, Chaceon), some species supporting important fisheries (Davie et al. 2007; Ng 1998). Manning and Holthuis (1981) first recognised the close relationship of these large deep-water crabs to other swimming crabs, noting a particular resemblance to Benthochascon and several other genera at that time placed in Portunidae. The portunoid position of geryonids is now well accepted and Benthochascon (and its relative, Rayminninus) have since been included in Geryonidae (Schubart and Reuschel 2009; Spiridonov et  al. 2014). Several studies observed a close relationship between ovalipids and geryonids, equivocally placing Ovalipes within or as sister to the geryonids (Evans 2018; Schubart and Reuschel 2009; Spiridonov et al. 2014). We follow Spiridonov et al. (2014) in treating Ovalipes (in Ovalipidae) as separate from Geryonidae. The phylogenetic positions of Nectocarcinus and Echinolatus are yet to be satisfactorily determined (Evans 2018; ­Spiridonov et al. 2014) but they share with geryonids, albeit possibly plesiomorphically, a triangular male telson with all pleonites clearly demarcated despite pleonites 3–5 being immovable (Spiridonov et  al. 2020). Additionally, the putatively polybiid genus Coenophthalmus, with a similar male pleon to other geryonids, was found to be closely related to Raymanninus and Nectocarcinus (Karasawa et  al. 2008). Therefore, we herein include Coenophthalmus, Echinolatus and Nectocarcinus in Geryonidae. The family was rediagnosed by Evans (2018) but based on different constituent genera. Diagnosis. Carapace as wide as or wider than long, subcircular to transversely ovate or subhexagonal; stridulatory ridges absent on carapace and chelipeds. Antennules folding transversely or obliquely. Male pleon triangular, margins strongly tapering, anterior width of somite 6 half or less that of somite 3; pleonites 1–6 and telson clearly demarcated, even if pleonites 3–5 immovable; pleonite 3 with transverse ridge. Pereopod 5 as a walking or swimming leg. Implicit generic characters. Branchial region not swollen, level with surrounding carapace surface. Orbit open mesially, antennal peduncle standing in orbit.

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Marine Decapod Crustacea

Key to genera of Geryonidae 1. – 2. – 3. – 4. – 5. – 6. – 7. –

Carapace front broadly rounded with median notch (Fig. 14.117f). Orbit closed mesially, completely excluding antennal peduncle�������������������������������������������������������������������������������������������������������������������������Coenophthalmus Carapace front multilobate, multidentate or arcuate and unbroken. Orbit open mesially, antennal peduncle standing in orbit (Fig. 14.117c)���������������������������������������������������������������������������������������������������������������������������������2 Pereopod 5 a walking leg, dactylus styliform, T-shaped in cross-section (Fig. 14.117m)�����������������������������������������������������3 Pereopod 5 a swimming leg, dactylus ovate to lanceolate, flat (Fig. 14.117a, b, n)�����������������������������������������������������������������5 Carapace with 2 anterolateral teeth behind exorbital angle, reduced to short points in large adults (Fig. 14.117h)���������������������������������������������������������������������������������������������������������������������������������������������������������������������Geryon Carapace with 4 anterolateral teeth behind exorbital angle, reduced to short points in large adults���������������������������������4 Carapace branchial region not swollen, level with surrounding carapace surface (Fig. 14.117d, e)���������������������� Chaceon Carapace branchial region swollen, raised above level of surrounding carapace (Fig. 14.117j)����������������������� Zariquieyon Carapace anterolateral teeth recurved anteriorly, 1 or more with additional slender spines on anterior margin (Fig. 14.117g)��������������������������������������������������������������������������������������������������������������������������������Echinolatus Carapace anterolateral teeth simple, without additional spinules on anterior margin���������������������������������������������������������6 Carapace anterolateral margin with 3 short, usually sharp, equal spines behind exorbital angle (Fig. 14.117b). Cheliped palm outer surface covered with granules, ridged (Fig. 14.117k)����������������������� Nectocarcinus Carapace anterolateral margin with 2 prominent spines, or 2 blunt lobes and 1 sharp spine behind exorbital angle. Cheliped palm outer surface smooth, without covering of granules and ridges (Fig. 14.117l)���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������7 Carapace anterolateral margin with 2 low blunt lobes and 1 sharp spine behind exorbital angle (Fig. 14.117a)�������������������������������������������������������������������������������������������������������������������������������������������������������� Benthochascon Carapace anterolateral margin with 2 prominent spines behind exorbital spine (Fig. 14.117i)�������������������� Raymanninus Benthochascon Alcock & Anderson, 1899

Diagnosis. Carapace front with 3 broad rounded lobes, median lobe occasionally emarginate; anterolateral margin with 2 low blunt lobes and 1 sharp spine behind exorbital angle. Cheliped palm outer surface smooth, without covering of granules and ridges. Pereopod 5 a swimming leg, dactylus ovate, flat. Maximum cl. 66 mm. Shelf, slope (150–950 m). Western and Central Indo-Pacific. 1 species (Alcock and Anderson 1899: diagnosis, figure; Ng 2000: redescription).

Chaceon Manning & Holthuis, 1989 Diagnosis. Carapace front multidentate; anterolateral margin with 4 teeth behind exorbital angle, reduced to short points in large adults. Pereopod 5 a walking leg, dactylus styliform, T-shaped in cross-section. Maximum cl. 200 mm (Pl. 61d). Shelf–bathyal (40–3477 m, usually deeper than 200 m). Cosmopolitan. 34 species (Davie et al. 2007; Manning and Holthuis 1989: diagnosis, 8 species; Tavares and Pinheiro 2011: key to 9 W Atlantic species).

Coenophthalmus A. Milne-Edwards, 1879 Diagnosis. Carapace front broadly rounded with median notch; anterolateral margin with 3 anterolateral teeth. Orbit closed mesially, completely excluding antennal peduncle. Pereopod 5 a swimming leg, dactylus flat, fusiform. Maximum cl. 12 mm.

Subtidal, shelf (19–49 m). Temperate South America (Uruguay to Patagonia). 1 species (de Melo 1996 figure, description).

Echinolatus Davie & Crosnier, 2006 Diagnosis. Carapace front quadrilobate; anterolateral margin with 3 short, usually sharp, equal anteriorly recurved spines behind exorbital angle, 1 or more with additional slender spines on anterior margin. Pereopod 5 a swimming leg, dactylus ovate to lanceolate, flat. Maximum cl. 18 mm. Subtidal–slope (10–580 m). Central and Eastern Indo-Pacific, Tropical Eastern Pacific, Temperate Australasia. 4 species (Davie and Crosnier 2006: diagnosis, key to species).

Geryon Krøyer, 1837 Diagnosis. Carapace front multidentate; anterolateral margin with 2 teeth behind exorbital angle, reduced to short points in large adults. Pereopod 5 a walking leg, dactylus styliform, T-shaped in cross-section. Maximum cl. 80 mm. Shelf–bathyal (32–2220 m, usually deeper than 100 m). Temperate Northern E Atlantic. 2 species (Manning and Holthuis 1987: species; 1989: rediagnosis).

Nectocarcinus A. Milne-Edwards, 1860 Diagnosis. Carapace front multilobate or arcuate, unbroken; anterolateral margin with 3 short, usually sharp, simple, equal spines

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Fig. 14.117.  Geryonidae. a, Benthochascon hemingi Alcock & Anderson, 1899; b, Nectocarcinus integrifrons (Latreille, 1825); c, left ventral orbit, Chaceon. Carapace: d, e, Chaceon; f, Coenophthalmus; g, Echinolatus; h, Geryon; i, Raymanninus; j, Zariquieyon. Cheliped manus, outer face: k, Nectocarcinus; l, Raymanninus. Pereopod 5: m, Chaceon; n, Echinolatus. Male pleonites 3–6, telson: o, Benthochascon; p, Echinolatus.

behind exorbital angle, without additional slender spines on anterior margin. Cheliped palm outer surface covered with granules, ridged. Pereopod 5 a swimming leg, dactylus ovate to lanceolate, flat. Maximum cl. 70 mm (Pl. 61e). Intertidal–slope (0–550 m). Central Indo-Pacific, Temperate Australasia. 6 species (Dell et al. 1970: rediagnosis, key to species; Poore 2004: key to Australian species).

Raymanninus Ng, 2000 Diagnosis. Carapace front with 4 narrow lobes; anterolateral margin with 2 prominent spines behind exorbital spine. Cheliped palm outer surface smooth, without covering of granules and ridges. Pereopod 5 a swimming leg, dactylus flat, fusiform. Maximum cl. 53 mm. Slope (329 m). Tropical W Atlantic. 1 species (Ng 2000).

Zariquieyon Manning & Holthuis, 1989 Diagnosis. Carapace front multidentate; anterolateral margin with 4 teeth behind exorbital angle, reduced to short points in large adults; branchial region swollen, raised above level of surrounding carapace. Pereopod 5 a walking leg, dactylus styliform, T-shaped in cross-section. Maximum cl. 22 mm. Bathyal (2830 m). Mediterranean Sea. 1 species (Manning and Holthuis 1989).

References Alcock A, Anderson ARS (1899) Illustrations of the zoology of the Royal Indian Marine Surveying Steamer “Investigator”. Crustacea, Part VII. Pls XXXVI–XLV. Trustees of the Indian Museum, Calcutta. Davie PJF, Crosnier A (2006) Echinolatus n. gen. (Crustacea, Decapoda, Portunidae) with description of two new species from the South-West Pacific. In: Richer de Forges B, Justine J-L (Eds) Tropical Deep-Sea Benthos vol. 24. Mémoires du Muséum National d’Histoire Naturelle, Paris 193, 393–410. Davie PJF, Ng PKL, Dawson EW (2007) A new species of deep-sea crab of the genus Chaceon Manning & Holthuis, 1989 (Crustacea: Decapoda: Brachyura: Geryonidae) from Western Australia. Zootaxa 1505, 51–62. doi:10.11646/zootaxa.1505.1.4 Dell RK, Griffin DJG, Yaldwyn JC (1970) A new swimming crab from the New Zealand subantarctic and a review of the genus Nectocarcinus A. Milne Edwards. Transactions of the Royal Society of New Zealand 12, 49–68. de Melo GAS (1996) Manual de identifição dos Brachyura (Caranguejos e siris) do litoral Brasileiro. Editora Plêiade Ltda, Sao Paulo. Evans N (2018) Molecular phylogenetics of swimming crabs (Portunoidea Rafinesque, 1815) supports a revised family-level classification and suggests a single derived origin of symbiotic taxa. PeerJ 6, e4260. doi:10.7717/peerj.4260 Karasawa H, Schweitzer CE, Feldmann RM (2008) Revision of Portunoidea Rafinesque, 1815 (Decapoda: Brachyura) with emphasis on the fossil genera and families. Journal of Crustacean Biology 28, 82–127. doi:10.1651/07-2882R.1

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Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Manning RB, Holthuis LB (1987) The status of Geryon trispinosus (Herbst) (Geryonidae). Investigaciones Pesqueras 51, 57–62. Manning RB, Holthuis LB (1989) Two new genera and nine new species of geryonid crabs (Crustacea, Decapoda, Geryonidae). Proceedings of the Biological Society of Washington 102, 50–77. Ng PKL (1998) Crabs. In FAO Species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 2. Cephalopods, crustaceans, holothurians and sharks. (Eds Carpenter KE, Niem VH) pp. 1045–1155. Food and Agriculture Organization, Rome. Ng PKL (2000) The deep-water swimming crabs of the genus Benthochascon (Decapoda: Brachyura: Portunidae), with description of a new genus for the American B. schmitti. Journal of Crustacean Biology 20, 310–324. doi:10.1163/1937240X-90000033 Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Schubart CD, Reuschel S (2009) A proposal for a new classification of Portunoidea and Cancroidea (Brachyura: Heterotremata) based on two independent molecular phylogenies. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 533–549. CRC Press, Boca Raton. Spiridonov VA, Neretina T, Schepetov D (2014) Morphological characterization and molecular phylogeny of Portunoidea Rafinesque,1815 (Crustacea Brachyura): implications for understanding evolution of swimming capacity and revision of the family-level classification. Zoologischer Anzeiger 253, 404–429. doi:10.1016/j.jcz.2014.03.003 Spiridonov VA, Simakova UV, Anosov SE, Zalota AK, Timofeev VA (2020) Review of Macropodia in the Black Sea supported by molecular barcoding data; with the redescription of the type material, observations on ecology and epibiosis of Macropodia czernjawskii (Brandt, 1880) and notes on other Atlanto-Mediterranean species of Macropodia Leach, 1814 (Crustacea, Decapoda, Inachidae). Zoosystematics and Evolution 96, 609–635. doi:10.3897/zse.96.48342 Tavares M, Pinheiro AP (2011) A new species of Chaceon Manning & Holthuis, 1989, from the southwestern Atlantic, with a key to the western Atlantic species (Crustacea, Decapoda, Geryonidae). Zootaxa 3086, 57–68. doi:10.11646/zootaxa.3086.1.3

corystoids, N. ocellatus has similar adaptations for breathing whilst fully buried. The antennal flagella are held together, with the bristly interlocking setae from each flagellum forming a respiratory tube to the surface of the substrate (Emmerson 2016). Diagnosis. Carapace longitudinally pear-shaped, longer than wide; margins with 4 prominent spines (excluding exorbital spine), glabrous or sparsely setose; front divided into 2 or 3 angular lobes. Antennules folding longitudinally. Antennal flagellum about half carapace length, strongly setose, appearing ‘furry’. Pereopod 5 dactylus spatulate, distally acute, ~2.5 times as long as wide. Male pleonite 3 without transverse ridge. Nautilocorystes H. Milne Edwards, 1837 Maximum cl. 35 mm. Intertidal–shelf (0–82 m). Temperate Southern Africa, Western and Central Indo-Pacific. 2 species (Barnard 1950: description, figures; Serène and Umali 1972: key to species). References Barnard KH (1950) Descriptive catalogue of South African decapod Crustacea (crabs and shrimps). Annals of the South African Museum 38, 1–837. Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from

Nautilocorystidae Ortmann, 1893 Figure 14.118 Nautilocorystids are unusual, elongated fossorial crabs with long, setose antennal flagella, superficially resembling that of corystoids, but with portunid-like chelipeds and fused male pleonites 3–5. Nautilocorystes, afforded its own family by Ortmann (1893), has usually been regarded as a corystoid (Barnard 1950), but is now recognised as a highly modified portunoid (Davie et  al. 2015; Spiridonov 2020; Spiridonov et  al. 2014). It has most recently been placed in Nautilocorystinae within either Thiidae (Davie et al. 2015) or tentatively within Carcinidae (Evans 2018; Spiridonov 2020) but its phylogenetic position has not been formally tested. Given its uncertain affinities and strong dissimilarity to thiids or carcinids, we return Nautilocorystes to Nautilocorystidae. The southern African species, N. ocellatus (Gray, 1831) is common on shallow subtidal sandy substrates. Like

Fig. 14.118.  Nautilocorystidae. a, Nautilocorystes ocellatus (Gray, 1831). b, c, pereopods 4, 5.

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the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. Emmerson WD (2016) A guide to, and checklist for, the Decapoda of Namibia, South Africa and Mozambique. Vol. 2. Cambridge Scholars Publishing, Cambridge. Evans N (2018) Molecular phylogenetics of swimming crabs (Portunoidea Rafinesque, 1815) supports a revised family-level classification and suggests a single derived origin of symbiotic taxa. PeerJ 6, e4260. doi:10.7717/peerj.4260 Ortmann AE (1893) Die Decapoden-Krebse des Strassburger Museums, mit besonderer Berücksichtigung der von Herrn Dr. Döderlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und zur Zeit im Strassburger Museum aufbewahrten Formen. VI Theil. Abtheilung: Brachyura (Brachyura genuina Boas), I. Unterabtheilung: Majoidea und Cancroidea, 1: Section Portuninea. Zoologische Jahrbucher. Abteilung fur Systematik, Geographie und Biologie der Tiere 7, 23–88. doi:10.5962/bhl.part.24064 Serène R, Umali AF (1972) The family Raninidae and other new and rare species of brachyuran decapods from the Philippines and adjacent regions. Philippine Journal of Science 99, 21–105. Spiridonov VA (2020) An update of phylogenetic reconstructions, classification and morphological characters of extant Portunoidea Rafinesque, 1815 (Decapoda, Brachyura, Heterotremata), with a discussion of their relevance to fossil material. Geologija 63, 133–166. doi:10.5474/ geologija.2020.014 Spiridonov VA, Neretina T, Schepetov D (2014) Morphological characterization and molecular phylogeny of Portunoidea Rafinesque,1815 (Crustacea Brachyura): implications for understanding evolution of swimming capacity and revision of the family-level classification. Zoologischer Anzeiger 253, 404–429. doi:10.1016/j.jcz.2014.03.003

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water (Parker et  al. 1998). Ovalipids are present in temperate-tropical waters worldwide except the eastern Atlantic. Ovalipes australiensis Stephenson & Rees, 1968, the surf crab, is common on open beaches in southern Australia and is fished recreationally and commercially. Diagnosis. Carapace as wide as or wider than long, subcircular to transversely ovate; with 5 anterolateral teeth. Antennal peduncle standing in orbit. Antennules folding transversely or obliquely. Stridulatory ridges present on lower surface of cheliped propodi or on pterygostomial surface of carapace. Chelipeds shorter than longest walking leg. Male pleon narrow, linguiform, margins weakly tapering, anterior width of pleonite 6 about two-thirds width of somite 3; pleonites 1–6 and telson clearly demarcated, even if pleonites 3–5 immovable; pleonite 3 with transverse ridge. Pereopod 5 as a swimming leg, dactylus ovate, distally rounded. Ovalipes Rathbun, 1898 lady crab, paddle crab, surf crab Maximum cl. 110 mm (Pl. 61f). Intertidal–slope (0–1010 m). Tropical W Atlantic, Indo-West Pacific, Temperate Australasia, Temperate South America. 11 species (Stephenson and Rees 1968: rediagnosis, key to 8 species; Williams 1976: 2 W Atlantic species).

Ovalipidae Spiridonov, Neretina & Schepetov, 2014 Figure 14.119, Plate 61f Ovalipes, has been variously regarded as a carcinid (Karasawa et al. 2008), polybiid (Ng et al. 2008), geryonid (Evans 2018; Schubart and Reuschel 2009), or placed in its own family, either monogenerically (Spiridonov et  al. 2014) or together with Echinolatus and Nectocarcinus (Davie et  al. 2015). Echinolatus and Nectocarcinus are regarded here as geryonids and we follow (Spiridonov et  al. 2014) in recognising a monogeneric Ovalipidae. Ovalipes morphologically resembles some polybiids, especially Polybius, but appears to be phylogenetically close to geryonids (Evans 2018; Schubart and Reuschel 2009; Spiridonov et al. 2014); these relationships are yet to be robustly resolved. Ovalipids can be distinguished from these families by the narrow, linguiform, male pleon and presence of stridulatory structures on the pterygostomial surface of carapace (and in most species, also on the chelipeds). Species of Ovalipes use sound (via stridulation) in conspecific signalling, especially in shallow water, but appear to have evolved towards use of reflected incident blue light (via multilayer cuticular reflectors) in deep

Fig. 14.119.  Ovalipidae. a, Ovalipes australiensis Stephensen & Rees, 1968. b, left front, pterygostomial ridge (arrowed), base of cheliped; c, male pleon.

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References Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. Evans N (2018) Molecular phylogenetics of swimming crabs (Portunoidea Rafinesque, 1815) supports a revised family-level classification and suggests a single derived origin of symbiotic taxa. PeerJ 6, e4260. doi:10.7717/peerj.4260 Karasawa H, Schweitzer CE, Feldmann RM (2008) Revision of Portunoidea Rafinesque, 1815 (Decapoda: Brachyura) with emphasis on the fossil genera and families. Journal of Crustacean Biology 28, 82–127. doi:10.1651/07-2882R.1 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Parker AR, McKenzie DR, Ahyong ST (1998) A unique form of light reflector and the evolution of signalling in Ovalipes (Crustacea: Decapoda: Portunidae). Proceedings of the Royal Society of London 265, 861–867. doi:10.1098/rspb.1998.0371 Schubart CD, Reuschel S (2009) A proposal for a new classification of Portunoidea and Cancroidea (Brachyura: Heterotremata) based on two independent molecular phylogenies. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 533–549. CRC Press, Boca Raton. Spiridonov VA, Neretina T, Schepetov D (2014) Morphological characterization and molecular phylogeny of Portunoidea Rafinesque,1815 (Crustacea Brachyura): implications for understanding evolution of swimming capacity and revision of the family-level classification. Zoologischer Anzeiger 253, 404–429. doi:10.1016/j.jcz.2014.​ 03.003 Stephenson W, Rees M (1968) A revision of the genus Ovalipes Rathbun, 1898 (Crustacea, Decapoda, Portunidae). Records of the Australian Museum 27, 213–261, pls 35–42, figs 1–4. Williams AB (1976) Distinction between a Gulf of Mexico and a Carolinian Atlantic species of the swimming crab Ovalipes (Decapoda: Portunidae). Proceedings of the Biological Society of Washington 89, 205–214.

Pirimelidae Alcock, 1899 Figure 14.120, Plate 61g The pirimelids are a small group of eastern Atlantic crabs closely resembling carcinids, differing chiefly in the styliform rather than flattened lanceolate pereopod 5 dactylus. Pirimelids have been variously allied to portunoids (especially carcinids) (Alcock 1899), atelecyclids (Gordon 1953), cancroids (Martin and Davis 2001), or thiids (Ng et  al. 2008) but their embedded position within the portunoids (along with thiids) has only relatively recently been determined by phylogenetic analysis (Schubart and Reuschel 2009; Spiridonov et  al. 2014). Some recent studies subsumed pirimelids into a wider Carcinidae (Evans 2018 (Evans 2018; Spiridonov 2020) but we treat the two families as separate (Spiridonov 2014 (Davie et al. 2015; Spiridonov et al. 2014). Diagnosis. Carapace as wide as or wider than long; front trilobed; with 4 or 5 anterolateral teeth; endostomial ridges absent; orbit open mesially, antennal peduncle standing in orbit. Antennules folding transversely or obliquely. Male pleonites 3–5 fused, sutures absent or only partially indicated. Chelipeds shorter than longest walking leg. Pereopods 2–5 dactylus styliform, similar, not flattened. Pirimela Leach, 1816 Diagnosis. Carapace frontal lobes on same plane, median lobe longer than laterals; with 5 anterolateral teeth. Orbit complete ventrally. Antennal flagellum sparsely setose (Fig.  14.120 a, c). Maximum cl. 21 mm (Pl. 61g). Intertidal–shelf (0–50 m). Temperate Northern and Tropical E Atlantic, including Mediterranean Sea. 1 species (Adema 1991: description, figure).

Fig. 14.120.  Pirimelidae. a, Pirimela; b; Sirpus. Right orbit (shaded), ventral face: c, Pirimela; d, Sirpus. e, pereopod 5, Sirpus.

14 – Brachyura – crabs

Sirpus Gordon, 1953 Diagnosis. Carapace median lobe as long as or shorter than lateral lobes, on lower plane; with 4 anterolateral teeth. Orbit incomplete ventrally. Antennal flagellum distinctly setose, appearing ‘feathered’ (Fig. 14.120b, d, e). Maximum cl. 7 mm. Intertidal–shelf (0–40 m). Temperate Northern and Tropical E Atlantic, including Mediterranean Sea, Black Sea. 4 species (Gordon 1953: diagnosis, descriptions; Manning and Holthuis 1981: species description; Vereshchaka 1989: species description). References Adema J (1991) De krabben van Nederland en Belgie (Crustacea, Decapoda, Brachyura). Nationaal Natuurhistorisch Museum, Leiden. Alcock A (1899) An account of the deep-sea Brachyura collected by the Royal Indian Marine Survey ship Investigator. Trustees of the Indian Museum, Calcutta. Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. Evans N (2018) Molecular phylogenetics of swimming crabs (Portunoidea Rafinesque, 1815) supports a revised family-level classification and suggests a single derived origin of symbiotic taxa. PeerJ 6, e4260. doi:10.7717/peerj.4260 Gordon I (1953) On Sirpus, a genus of pygmy cancroid crabs. Bulletin of the British Museum, Natural History. Zoology 2, 43–65. doi:10.5962/p.314145 Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Martin JW, Davis GE (2001) An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County. Science Series 39, 1–124. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Schubart CD, Reuschel S (2009) A proposal for a new classification of Portunoidea and Cancroidea (Brachyura: Heterotremata) based on two independent molecular phylogenies. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 533–549. CRC Press, Boca Raton. Spiridonov VA (2020) An update of phylogenetic reconstructions, classification and morphological characters of extant Portunoidea Rafinesque, 1815 (Decapoda, Brachyura, Heterotremata), with a discussion of their relevance to fossil material. Geologija 63, 133–166. doi:10.5474/ geologija.2020.014 Spiridonov VA, Neretina T, Schepetov D (2014) Morphological characterization and molecular phylogeny of Portunoidea Rafinesque,1815 (Crustacea Brachyura): implications for understanding evolution of swimming capacity and revision of the family-level classification. Zoologischer Anzeiger 253, 404–429. doi:10.1016/j.jcz.2014.03.003 Vereshchaka AL (1989) A new crab species Sirpus ponticus sp.n. (Crustacea, Pirimelidae) from the Black Sea. Zoologicheskij Zhurnal 67, 41–47[in Russian].

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Polybiidae Ortmann, 1893 swimming crabs Figure 14.121, Plate 61h–j Polybiidae (sometimes called Macropipidae) include species having a carcinid-like carapace shape, male pleonite fusion and chelipeds that are shorter than the longest walking leg. Unlike carcinids, polybiids have epistomial ridges and pereopod 5 developed as a swimming leg. Distinctions between Liocarcinus, Macropipus and Polybius have been historically fluid and their species can be found under varying generic combinations in synoptic works and guides (e.g. Ashton et  al. 2017; Christiansen 1969; d’Udekem d’Acoz 1999; Ingle 1980; Zariquiey Alvarez 1968). Moreover, some taxonomic and nomenclatural issues remain to be resolved in Liocarcinus and Polybius (Plagge et  al. 2016), so the taxonomy is not yet stable. Here, we treat Liocarcinus, Macropipus and Polybius as separate following Ingle (1980), Manning and Holthuis (1981) and Ng et al. (2008). Several genera formerly regarded as polybiids, have recently been removed to other taxa: Ovalipes to Ovalipidae, Brusinia to Brusiniidae and Benthochascon to Geryonidae (Spiridonov et al. 2014). The little studied south-western Atlantic genus Coenophthalmus, usually associated with other polybiid genera by default (Ng et  al. 2008; Rathbun 1930), is herein transferred to Geryonidae given its geryonid-like male pleon and close relationship to Raymanninus and Nectocarcinus (Karasawa et al. 2008). Polybiidae now comprise five genera and 32 species, most of which are restricted to the eastern Atlantic. Liocarcinus, however, is also represented by two species in the Indo-West Pacific (Plagge et  al. 2016), Bathynectes occurs on both sides of the Atlantic (Manning and Holthuis 1981; Williams 1984) and Parathranites is widespread in the Indo-Pacific (Crosnier 2002). Diagnosis. Carapace as wide as or wider than long, subcircular to transversely ovate or subhexagonal; 5 anterolateral teeth; endostomial ridges long, extending to epistome; orbit open mesially. Antennal peduncle standing in orbit. Antennules folding transversely or obliquely. Male pleonites 3–5 fused, sutures absent or only partially indicated; pleonite 3 with transverse ridge. Chelipeds shorter than longest walking leg. Pereopod 5 as a swimming leg, dactylus ovate to fusiform. Implicit generic characters. Junction of posterior and posterolateral borders forming curved ridge. Dactyli slender, styliform, rounded to polygonal in cross-section. Pereopod 4 longer than pereopod 3.

Key to genera of Polybiidae 1. –

Pereopods 2–4 dactyli lanceolate, flattened (Fig. 14.121f)��������������������������������������������������������������������������������������������Polybius Pereopods 2–4 dactyli slender, styliform, rounded to polygonal in cross-section (Fig. 14.121a, b, e)��������������������������������2

698

Marine Decapod Crustacea

Fig. 14.121.  Polybiidae. Habitus, carapace: a, Bathynectes piperitus Manning & Holthuis, 1981; b; Liocarcinus strigilis (Stimpson, 1858); c, Macropipus; d, Necora puber (Linnaeus, 1767); e, Parathranites orientalis (Miers, 1886); f, Polybius henslowii Leach, 1822. Orbits, antennules, antennae, epistome, endostome (maxillipeds 3 removed): g, Liocarcinus; h, Parathranites. i, pereopod 4, Macropipus. Pereopod 2 propodus, lower margin: j, Liocarcinus; k, Macropipus. l, pleonites 3–6, telson, Parathranites.

2. – 3. – 4. – 5. –

Cheliped merus with spine on anterior and upper margin (Fig. 14.121a, e)���������������������������������������������������������������������������3 Cheliped merus without anterior or upper spine (Fig. 14.121b)������������������������������������������������������������������������������������������������4 Carapace, junction of posterior and posterolateral borders forming curved ridge. Pereopod 4 longer than pereopod 3. Pereopod 5 distally pointed (Fig. 14.121a). Male telson wider than half width of pleonite 6�������������������������������������������������������������������������������������������������������������������������������������������������������������������Bathynectes Carapace, junction posterior and posterolateral borders forming upcurved spine. Pereopod 3 longer than pereopod 4. Pereopod 5 distally rounded (Fig. 14.121e). Male telson half width of pleonite 6 (Fig. 14.121l)�����������������������������������������������������������������������������������������������������������������������������������������Parathranites Carapace front shallowly concave, with pair of submedian lobes flanked by numerous small spines and denticles (Fig. 14.121d)����������������������������������������������������������������������������������������������������������������������������������������������������� Necora Carapace front approximately convex in general outline, either unbroken, sinuous or cut into 3 lobes or teeth (Fig. 14.121b)����������������������������������������������������������������������������������������������������������������������������������������������������������������������5 Pereopods 2–4 propodi lower margin with single longitudinal carina extending full length of margin (Fig. 14.121j); if second carina present, short, never extending full length of margin���������������������������������� Liocarcinus Pereopods 2–4 propodi lower margin with 2 longitudinal carinae, each extending full length of margin (Fig. 14.121i, k)����������������������������������������������������������������������������������������������������������������������������������������������������������Macropipus

14 – Brachyura – crabs

Bathynectes Stimpson, 1871 Diagnosis. Carapace transversely subovate-subhexagonal; front almost straight, cut into 4 teeth. Cheliped merus with spine on anterior and upper margin. Pereopods 2–4 propodi lower margins with 2 longitudinal carinae extending full length of margin. Pereopod 5 distally pointed. Male telson wider than half width of pleonite 6. Maximum cl. 69 mm. Subtidal–slope (0–1455 m). Temperate Northern and Tropical Atlantic, Temperate Southern Africa. 5 species (Manning and Holthuis 1981: comparison of species).

Liocarcinus Stimpson, 1871 Diagnosis. Carapace transversely subovate-subhexagonal; front approximately convex in general outline, unbroken, sinuous or cut into 3 lobes or teeth. Cheliped merus without anterior or upper spine. Pereopods 2–4 propodi lower margins with single longitudinal carina extending full length, if second carina present, short, never extending full length of margin. Maximum cl. 65 mm (Pl. 61h). Intertidal–slope (0–455 m). Temperate Northern Atlantic, Temperate Northern Pacific, Western Indo-Pacific, Central IndoPacific, Temperate Australasia. 13 species (Plagge et al. 2016: discussion of world-wide distribution of Liocarcinus corrugatus (Pennant, 1777); Zariquiey Alvarez 1968: key to 11 European species of Macropipus includes 6 species of Liocarcinus). The North Atlantic, L. navigator (Herbst, 1794) has been recorded from ­Brazil (de Melo and Crivelaro 2002) but it is not clear whether it is established there.

Macropipus Prestandrea, 1833 Diagnosis. Carapace transversely subovate-subhexagonal; front approximately convex in general outline, cut into 3 teeth. Cheliped merus without anterior or upper spine. Pereopods 2–4 propodi lower margins with 2 longitudinal carinae extending full length of margin. Maximum cl. 37 mm. Intertidal–slope (0–834 m). Temperate Northern and Tropical Atlantic, Temperate Southern Africa. 4 species (Guinot 1961: comparison of 3 species).

Necora Holthuis, 1987 Diagnosis. Carapace ovate; front shallowly concave, with pair of submedian lobes flanked by numerous small spines and denticles. Cheliped merus without anterior or upper spine. Pereopods 2–4 propodi lower margins with 2 longitudinal carinae extending full length of margin. Maximum cl. 65 mm (Pl. 61i). Intertidal–shelf (0–60 m). Temperate Northern E Atlantic. 1 species (Holthuis 1987).

Parathranites Miers, 1886 Diagnosis. Carapace transversely subovate-subhexagonal; junction of posterior and posterolateral borders forming upcurved spine. Cheliped merus with spine on anterior and upper margin.

699

Pereopod 3 longer than pereopod 4. Pereopod 5 distally rounded. Male telson half width of pleonite 6. Maximum cl. 25 mm (Pl. 61j). Shelf, slope (73–510 m). Indo-West Pacific, Temperate ­Australasia. 8 species (Crosnier 2002: rediagnosis, key to species).

Polybius Leach, 1820 Diagnosis. Carapace subcircular; front approximately convex in general outline, cut into 4 teeth. Dactyli lanceolate, flattened. Maximum cl. 40 mm. Usually pelagic, often swarming offshore (2–1245 m). Temperate Northeast Atlantic. 1 species (Magalhães et al. 2014: significant wasted by-catch of prawn fisheries; Zariquiey Alvarez 1968: rediagnosis). References Ashton GV, Brandt A, Isaac MJ, Makings P, Moyse J, et al. (2017) Crustaceans. In Handbook of the marine fauna of north-west Europe. Second edition. (Eds Hayward PJ, Ryland JS) pp. 283–440. Oxford University Press, New York. Christiansen ME (1969) Crustacea Decapoda Brachyura. Marine Invertebrates of Scandinavia 2, 1–143. Crosnier A (2002) Révision du genre Parathranites Miers, 1886 (Crustacea, Brachyura, Portunidae). Zoosystema 24, 799–825. de Melo GAS, Crivelaro TB (2002) Primeira ocorrência de Polybius navigator (Herbst) (Decapoda, Brachyura, Portunidae) no Atlântico ocidental. Revista Brasileira de Zoologia 19, 233–238. doi:10.1590/ S0101-81752002000100022 d’Udekem d’Acoz C (1999) Inventaire et distribution des crustacés décapodes de l’Atlantique nord-oriental, de la Méditerranée et des eaux continentales adjacentes au nord de 25°N. Patrimoines Naturels 40, i–x, 1–394, 11 figs, 2 maps. Guinot D (1961) Caractères et affinités de Macropipus australis sp. nov., Crustacé Décapode Brachyoure de la côte sud-ouest Africaine. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique 37, (26) 1 –13, pls 1, 2. Holthuis LB (1987) Necora, a new genus of European swimming crabs (Crustacea Decapoda, Portunidae) and its type species, Cancer puber L., 1767. Zoölogische Mededeelingen 61, 1–14. Ingle RW (1980) British crabs. Oxford University Press, Inc., New York. Karasawa H, Schweitzer CE, Feldmann RM (2008) Revision of Portunoidea Rafinesque, 1815 (Decapoda: Brachyura) with emphasis on the fossil genera and families. Journal of Crustacean Biology 28, 82–127. doi:10.1651/07-2882R.1 Magalhães L, Felício M, Gaspar MB, Quintino V, Rodrigues AM (2014) Reproductive biology of Polybius henslowii Leach, 1820 (Decapoda, Polybiidae): a discard species from the northwestern Portuguese beam trawl fishery. Crustaceana 87, 784–800. doi:10.1163/15685403-00003330 Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Plagge C, Nguyen TS, Ng PKL, Türkay M, Streit B, et al. (2016) Liocarcinus corrugatus (Pennant, 1777) (Crustacea: Brachyura: Portunidae): a cosmopolitan brachyuran species? Raffles Bulletin of Zoology 64, 374–388. Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Spiridonov VA, Neretina T, Schepetov D (2014) Morphological characterization and molecular phylogeny of Portunoidea Rafinesque,1815

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(Crustacea Brachyura): implications for understanding evolution of swimming capacity and revision of the family-level classification. Zoologischer Anzeiger 253, 404–429. doi:10.1016/j.jcz.2014.03.003 Williams AB (1984) Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C. Zariquiey Alvarez R (1968) Crustáceos decápodos ibéricos. Investigaciones Pesqueras 32, 1–510.

Portunidae Rafinesque, 1815 swimming crabs Figures 14.122–14.129, Plates 61k–o, 62a–k The portunids are the ‘classic’ swimming crabs, best exemplified by commercial genera such as Callinectes, Charybdis, Portunus and Scylla (Ng 1998; Williams 1984). Species in these and related genera are active, agile, free-living predators; all have flattened walking legs and a well developed, paddle-like swimming leg, enabling rapid and effective swimming, as well as burying in soft substrates. The portunids, however, include not only the most capable swimmers, but also highly modified genera in which the swimming adaptations are secondarily reduced or lost, as in most Carupinae and some commensal thalamitines (Lissocarcinus, Caphyra) (Evans 2018; Spiridonov and Neumann 2008). Portunids occur worldwide in tropical to temperate waters but are most diverse in the tropical Indo-west Pacific (Stephenson 1972). In the past two decades, five main groupings have generally been recognised for genera now placed in Portunidae:

Caphyrinae, Carupinae (or Catoptrinae), Podophthalminae, Portuninae and Thalamitinae (Apel and Spiridonov 1998; Davie 2002; Davie et al. 2015; Ng et al. 2008). New schemes based on results of phylogenetic analysis and consideration of fossils (Davie et al. 2015; Evans 2018; De Grave et al. 2009; Karasawa et al. 2008; Spiridonov 2020) introduced additional subfamilies, mainly for genera formerly included in Portuninae (Evans 2018; Spiridonov 2020). Significantly, Evans (2018) showed that Caphyrinae is nested within Thalamitinae and synonymised the two groups. For portunid subfamilies, we herein largely follow Evans (2018) and Spiridonov (2020), but regard Coelocarcininae as portunid rather than carcinid (the chelipeds being longer instead of shorter than the longest walking legs), and regard Sanquerus as a portunine rather than necronectine (because of its prismatic and ridged, rather than inflated, smooth chelae). The poorly known Indo-West Pacific portunine genus Carupella Lenz & Strunck, 1914, is not keyed here given that it is based on early juveniles. The type species, C. natalensis Lenz & Strunck, 1914, is most likely a portunine (possibly lupocycline) and others referred to the genus are based on Portunus and probably other portunines (Spiridonov 2020). Diagnosis. Carapace as wide as or wider than long, subcircular to transversely ovate or subhexagonal. Antennules folding transversely or obliquely. Chelipeds longer than longest walking legs (pereopods 2 or 3). Male pleonites 3–5 fused, sutures absent or only partially indicated. Pereopod 5 usually as swimming leg with ovate dactylus ovate; rarely narrowly crescentic, flattened (Libystes). Male pleonite 3 usually with transverse ridge.

Key to subfamilies and genera of Portunidae The exorbital, or outer orbital, tooth is counted as the first anterolateral tooth.

1. – 2. – 3. – 4. – 5. –

Eyestalks extremely long, slender, exceeding half carapace length (Fig. 14.127)���������������������������Podophthalminae … 2 Eyestalks short, stout, shorter than one-fourth carapace length����������������������������������������������������������������������������������������������4 Carapace with 3 or 4 anterolateral spines. Eyestalk about half carapace length (Fig. 14.127a)����������������������������� Euphylax Carapace with 2 anterolateral spines. Eyestalk as long as or longer than carapace length (Fig. 14.127b, c)����������������������3 Eyestalk simple, without wing-like expansions next to cornea (Fig. 14.127b; Pl. 62a)�������������������������������� Podophthalmus Eyestalk with wing-like expansions next to cornea (Fig. 14.127c)�������������������������������������������������������������Vojmirophthalmus Cheliped carpus with 0–2 spines on outer face (Fig. 14.124b). Antenna flagellum standing in orbit; basal article narrow, as long as or longer than wide������������������������������������������������������������������������������������������������������������������������5 Cheliped carpus with 0–4 (usually 3) spines on outer face. Antenna flagellum excluded from orbit, at least in adults (except Charybdis (Gonioneptunus), but 3 outer carpal spines present); basal article usually distinctly wider than long (if longer than wide, flagellum always excluded from orbit)����������������� Thalamitinae … 31 Carapace front undivided, prominently protruding beyond general anterior outline (Fig. 14.124a). Pereopods 2–4 shorter than carapace. Pereopod 5 dactylus and propodus circular (Fig. 14.124c)����������������������������������������������������������������������������������������������������� Coelocarcininae … Coelocarcinus Carapace front usually bi- or multilobate, not markedly protruding beyond general anterior outline, lobes, if present, forming more-or-less even arc. Pereopods 2–4 longer than carapace. Pereopod 5 dactylus and propodus elongate-ovate to lanceolate, never circular����������������������������������������������������������������������������������������������������������6

14 – Brachyura – crabs

6.

701

Pereopod 5 merus linear, twice or more as long as wide, no wider than pereopod 4 merus; merus lower margin sometimes distally acute, never with subdistal spine (Fig. 14.123q–s)���������������������������������������Carupinae … 7 – Pereopod 5 merus usually ovate, less than twice as long as wide, wider than pereopod 4 merus, if linear and more than twice as long as wide (some Lupocyclus), merus lower margin with subdistal spine����������������������������������14 7. Cheliped fingers slender, occlusal margins lined with obliquely directed needle-like spines (Fig. 14.123k, l)�������������������������������������������������������������������������������������������������������������������������������������������������������� Atoportunus – Cheliped fingers slender or robust, occlusal margins variously armed�����������������������������������������������������������������������������������8 8. Carapace pterygostomial region with stridulatory ridge (Fig. 14.123j); last anterolateral spine much the longest, directed laterally; junction of posterior and posterolateral borders forming upcurved spine (Fig. 14.123e, f). Cheliped palm with spines, ridges and granules (Fig. 14.123m); merus upper-proximal surface with stridulatory ridges����������������������������������������������������������������������������������������������������������������������������� Laleonectes – Carapace pterygostomial region without stridulatory ridges; junction of posterior and posterolateral borders forming curved ridge (Fig. 14.123a–d, g–i). Cheliped palm unarmed, surface smooth, without spines, granules or carinae (Fig. 14.123n o, p); merus upper-proximal surface without stridulatory ridges���������������9 9. Cheliped carpus inner margin with distinct spine (Fig. 14.123o)������������������������������������������������������������������������������������������10 – Cheliped carpus inner margin rounded to bluntly right-angled��������������������������������������������������������������������������������������������11 10. Carapace front bilobed; with 4 anterolateral teeth (Fig. 14.123i)���������������������������������������������������������������������������� Richerellus – Carapace front quadrilobed; with 7 anterolateral teeth (Fig. 14.123b)������������������������������������������������������������������������ Carupa 11. Maxilliped 3 merus distolateral margin produced, bluntly angular to auriculiform (Fig. 14.123u, v). Cheliped merus without spines on anterior margin����������������������������������������������������������������������������������������������������������� 12 – Maxilliped 3 merus distolateral margin weakly produced, evenly rounded (Fig. 14.123t). Cheliped merus usually with 1 or more spines on anterior margin (Fig. 14.123n)��������������������������������������������������������������������������������������13 12. Carapace transversely subrectangular; straight posterior margin between coxae of pereopods 5 as wide as or wider than fronto-orbital width (Fig. 14.123g). Maxilliped 3 merus distolateral margin strongly produced, auriculiform (Fig. 14.123v)��������������������������������������������������������������������������������������������������������������������������Libystes – Carapace transversely broad, twice as wide as long; straight posterior margin between coxae of pereopods 5 narrower than fronto-orbital width (Fig. 14.123d). Maxilliped 3 merus distolaterally angular, margins prominently granular (Fig. 14.123u)������������������������������������������������������������������������������������������������������������������������������� Kume 13. Carapace transversely ovate; with 6 distinct anterolateral teeth (Fig. 14.123c). Male telson length little longer than wide (Fig. 14.123w)�������������������������������������������������������������������������������������������������������������������������������� Catoptrus – Carapace transversely broad, twice as wide as long; with 6 low anterolateral granules (Fig. 14.123h). Male telson twice as long as wide (Fig. 14.123x)��������������������������������������������������������������������������������������������������������������������������Pele 14. Junction of carapace posterior and posterolateral borders acutely angular or spiniform, without curved ridge (Fig. 14.128c, h, i, l, m)����������������������������������������������������������������������������������������������������������� Portuninae (part1) … 15 – Junction of carapace posterior and posterolateral borders forming curved ridge (Figs 14.122a, b, 14.128a, b, d–g, j, k)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������19 15. Maxilliped 3 merus strongly produced laterally as angular lobe (Fig. 14.128u)�������������������������������������������������������Eodemus – Maxilliped 3 merus rounded or produced distally as blunt lobe, not laterally directed (Fig. 14.128s, x)������������������������16 16. Maxilliped 3 merus distally rounded, not produced distally beyond palp (Fig. 14.128s)�������������������������������������Alionectes – Maxilliped 3 merus with long distal lobe produced well beyond palp (Fig. 14.128x)���������������������������������������������������������17 17. Front with 3 teeth, median tooth as long as or longer than lateral teeth (Fig. 14.128l)�����������������������������������������Trionectes – Front with 3 or 4 teeth, if 3 teeth present, median shorter than laterals�������������������������������������������������������������������������������18 18. Carapace with anterolateral teeth lobiform, blunt (Fig. 14.128i). Cheliped fingers shorter than dorsal margin of palm (Fig. 14.128o)�������������������������������������������������������������������������������������������������������������������������������������� Incultus – Carapace anterolateral teeth sharp, triangular or spiniform (Fig. 14.128m). Cheliped fingers about as long as dorsal margin of palm (Fig. 14.128q)���������������������������������������������������������������������������������������������������������������Xiphonectes 19. Maxilliped 3 merus elongated distally, palp placed near mid-length of inner meral margin (Fig. 14.122d)����� Achelouinae … 20

702

– 20. – 21. – 22. – 23. – 24. – 25. – 26. – 27. – 28. – 29. – 30. – 31. – 32. –

Marine Decapod Crustacea

Maxilliped 3 merus not elongated, palp placed near inner distal meral margin, or at least in distal half, such that upper margin of carpus of folded palp in line or above distal margin of merus (Fig. 14.125d, 14.128v, w)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������21 Cheliped fingers greatly elongated, more than 3 times as long as palm in adult males, 1.5 times in adult females. Pereopod 5 merus upper margin with distal spine (Fig. 14.122b)���������������������������������������������������Lupella Cheliped fingers as long as or shorter than palm in both sexes. Pereopod 5 merus upper margin without distal spine (Fig. 14.122a)������������������������������������������������������������������������������������������������������������������������������Achelous Pereopod 5 merus lower margin with distal spine (Fig. 14.125c)������������������������������������������������������������Lupocyclinae … 22 Pereopod 5 merus unarmed�������������������������������������������������������������������������������������������������������������������������������������������������������� 23 Carapace subcircular to narrowly ovate, 1.1–1.3 times as wide as long; with 5 equal anterolateral teeth or 9 teeth of alternating size (Fig. 14.125b, c). Antenna basal article without distolateral lobule������������������������ Lupocyclus Carapace transversely ovate, 1.5–1.7 times as wide as long; with 8 equal anterolateral teeth, ninth tooth distinctly longer (Fig. 14.125a). Antenna basal article with distolateral lobule���������������������������� Lupocycloporus Cheliped palm inflated, ovate in cross-section, surface smooth, without ridges or granules (Fig. 14.126b; Pl. 61o)�������������������������������������������������������������������������������������������������������������������������Necronectinae … Scylla Cheliped palm prismatic, ridged, variously granulate (Fig. 14.128n–q)�����������������������������������������Portuninae (part) … 24 Cheliped carpus inner spine absent (Fig. 14.128n)�����������������������������������������������������������������������������������������������������Callinectes Cheliped carpus inner spine present (Fig. 14.128a, b)������������������������������������������������������������������������������������������������������������� 25 Carapace surface evenly granulate. Maxilliped 3 merus distolaterally rounded (Fig. 14.128w)�������������������������������������� 26 Carapace surface with granules distributed in groups or patches (Fig. 14.128d, f, j). Maxilliped 3 merus distolaterally produced as angular or auriculiform lobe (Fig. 14.128t, v)����������������������������������������������������������������������� 28 Carapace dorsal orbital fissures open, narrowly V-shaped (Fig. 14.128a). Endostomial ridges absent��������������� Arenaeus Carapace dorsal orbital fissures closed, margins appressed (Fig. 14.128b, k). Endostomial ridges present (Fig. 14.128r)������������������������������������������������������������������������������������������������������������������������������������������������������������������������������ 27 Carapace dorsal surface distinctly granular, rugose; metagastric, mesogastric and epibranchial ridges distinct (Fig. 14.128k)���������������������������������������������������������������������������������������������������������������������������������������������������Portunus Carapace dorsal surface microscopically granulate, appearing smooth; metagastric, mesogastric and epibranchial ridges scarcely discernible (Fig. 14.128b)���������������������������������������������������������������������������������������� Sanquerus Carapace broad, 1.5 or more times as long as wide, transversely ovate-subhexagonal, last anterolateral tooth distinctly larger than other teeth (Fig. 14.128d, j)���������������������������������������������������������������������������������������������������� 29 Carapace narrow, 1.4 or less times as long as wide, with subcircular-hexagonal outline, last anterolateral tooth smaller than to only slightly larger other teeth (Fig. 14.128f, g)���������������������������������������������������������������������������� 30 Carapace broadly ovate-subhexagonal, 1.7 times as wide as long; front with 4 low teeth, median pair shorter, projecting little beyond inner orbital angle (Fig. 14.128j). Cheliped merus posterior margin with 2 distal spines (Fig. 14.128p)���������������������������������������������������������������������������������������������������������������������������������������Monomia Carapace narrowly ovate, 1.5 times as wide as long; front with 4 projecting subequal teeth projecting distinctly beyond inner orbital angle (Fig. 14.128d). Cheliped merus posterior margin with 1 distal spine�������������������������������������������������������������������������������������������������������������������������������������������������������������Allomonomia Male sternopleonal cavity with pair of hemispherical cavities to accommodate large lateral subdistal lobe of gonopod 1 (Fig. 14.128y)���������������������������������������������������������������������������������������������������������������������������������Cavoportunus Male sternopleonal cavity evenly V-shaped. Gonopod 1 without large lateral subdistal lobes (Fig. 14.128z)���������������������������������������������������������������������������������������������������������������������������������������������������������Cycloachelous Cheliped carpus outer surface smooth or granular (Fig. 14.129p)�����������������������������������������������������������������������������������������32 Cheliped carpus outer surface with 3 or 4 large spines (Fig. 14.129o)�����������������������������������������������������������������������������������33 Pereopod 5 a walking leg, dactylus slender, claw-like or lanceolate (Fig. 14.129t). Pereopods 2–4 longer than carapace������������������������������������������������������������������������������������������������������������������������������������������������������������������Caphyra Pereopod 5 a swimming leg, dactylus flattened, ovate, apex rounded or pointed (Fig. 14.129u). Pereopods 2–4 as long as or shorter than carapace������������������������������������������������������������������������������������������������������������� Lissocarcinus

14 – Brachyura – crabs

703

33. – 34. – 35.

Carapace with 9 anterolateral teeth, alternating in size, all prominent (Fig. 14.129e)����������������������������������������������Cronius Carapace with 7 or fewer anterolateral teeth, some minute��������������������������������������������������������������������������������������������������� 34 Carapace with 4 or 5 (rarely 3) anterolateral teeth, size variable��������������������������������������������������������������������������������������������35 Carapace with 6 or 7 anterolateral teeth (second, fourth, sixth teeth may be minute)�������������������������������������������������������37 Carapace with spinous anterolateral margin continuous with unarmed posterolateral margin, forming more-or-less straight or broadly curved margin (Fig. 14.129k). Cheliped dactylus spoon-tipped (Fig. 14.129q)�������������������������������������������������������������������������������������������������������������������������������������������������������� Thalamitoides – Carapace with spinous anterolateral margin demarcated from unarmed posterolateral margin by obtuse angle (Fig. 14.129b–d, f, g, i–l). Cheliped dactylus tip usually simple (spoon-tipped only in Thalamita stephensoni Crosnier, 1962)���������������������������������������������������������������������������������������������������������������������������������������������������� 36 36. Cheliped palm outer surface with granular ridge terminating in small spine near base of pollex (Fig. 14.129r). Pereopods 2–4 meri lower margin with distal spine (Fig. 14.129s)���������������������������������������������������Zygita – Cheliped palm outer surface without spine near base of pollex, granular ridge present or absent. Pereopod 2–4 meri lower margin usually without distal spine���������������������������������������������������������������������������������Thalamita (part) 37. Junction of carapace posterior and posterolateral borders bluntly angular, without broadly curved ridge (Fig. 14.129c, d). Cheliped merus with posterodistal spine����������������������������������������������������������������������������������������������� 38 – Junction of carapace posterior and posterolateral borders forming broadly curved ridge (Fig. 14.129f). Cheliped merus without posterodistal spine����������������������������������������������������������������������������������������������������������������������� 39 38. Antennal flagellum excluded from orbit (Fig. 14.129n)����������������������������������������������������������������Charybdis (Goniohellenus) – Antennal flagellum standing in orbit (Fig. 14.129m)������������������������������������������������������������������ Charybdis (Gonioneptunus) 39. Carapace anterolateral margin with 4 large teeth and 2 or 3 minute teeth, one each behind first, second and sometimes third large teeth (Fig. 14.129f)����������������������������������������������������������������������������������������������������� Gonioinfradens – Carapace anterolateral margin with at least 5 large teeth������������������������������������������������������������������������������������������������������� 40 40. Carapace anterolateral margin with 6 large teeth or 5 large teeth and smaller tooth behind first large tooth, length of smaller tooth exceeding half-length of posterior margin of first tooth (Fig. 14.129b)������������������������������������������������������������������������������������������������������������������������������������������Charybdis (Charybdis) – Carapace anterolateral margin with 5 large teeth and minute tooth behind first large tooth, length of minute teeth less than half length of posterior margin of adjacent large tooth; with or without minute tooth behind second large teeth (Fig. 14.129g, i)������������������������������������������������������������������������������������������������������������������41 41. Carapace with third and fourth large anterolateral teeth equal; minute tooth behind first large tooth, usually also behind second large tooth, sometimes behind third large tooth (Fig. 14.129g)��������������� Goniosupradens – Carapace with third large anterolateral tooth slightly larger than fourth large tooth; minute tooth behind first large tooth only (Fig. 14.129i)��������������������������������������������������������������� Thalamita (part, T. exetastica species group) 1 Achelous iridescens (Rathbun, 1893) and A. tuberculatus Stimpson, 1860 will key out here, but can be distinguished from Xiphonectes and allies by the wider distal half of the maxilliped 3 merus – little narrower than the proximal half in Achelous (Fig. 14.122d), about half in the Xiphonectes (Fig. 14.128x).

Subfamily Achelouinae Spiridonov, 2020 Spiridonov (2020) established Achelouinae for a clade of Eastern Pacific and Atlantic portunines first identified by molecular phylogenetics (Evans 2018; Mantelatto et  al. 2009), which included Lupella, a group of species of Portunus for which the synonymised genus Achelous was resurrected and reconstituted, and several other species of ‘Portunus’. Achelouinae as diagnosed by Spiridonov (2020), however, was indistinguishable from Portuninae, and diagnostic features of the reinstated and much constrained Achelous were not identified by Mantelatto et  al. (2009;

2018), obscuring its taxonomic limits (and by extension, that also of Portunus) (Evans 2018; Rodrigues et al. 2017). Our re-evaluation of diagnostic features of the achelouinae clade emphasises the shape of the maxilliped 3 merus, a feature noted by Stimpson (1860) for species he referred to Achelous, but overlooked by subsequent workers. In achelouines, the maxilliped 3 merus is distally elongated and suboblong, so that the palp is placed near the midlength of the merus, with the distal half little narrower than the proximal half. In most portunines (except some species of Xiphonectes), however, the maxilliped 3 merus is not distally elongated, with the palp placed near inner distal meral

704

Marine Decapod Crustacea

margin, or at least in the distal half, such that upper margin of carpus of the folded palp is in line with or above the distal margin of merus. In Xiphonectes, the maxilliped 3 merus is usually also elongated distally, but the distal portion is distinctly narrower, being about half the width of the proximal portion. Notably, two species of Achelous (A. iridescens (Rathbun, 1893), A. tuberculatus Stimpson, 1860) will key to Xiphonectes because of the acute-spiniform rather than rounded junction of the posterior and posterolateral borders of the carapace, but they remain separable from Xiphonectes by the much wider distal half of the maxilliped 3 merus. Spiridonov (2020) included in Achelouinae, Achelous, Lupella and the several species of Portunus forming a clade with Lupella in Evans’ (2018) phylogram, and treated as ‘Portunus 1’ by Mantelatto et  al. (2018), viz: P. anceps (de Saussure, 1857), P. floridanus (Rathbun, 1930), P. hastatus (Linnaeus, 1767), P. inequalis (Miers, 1881), P. ventralis (A. Milne-Edwards, 1879)). Koch et al. (2022) included these species and the monotypic Lupella in Achelous. Both genera are provisionally retained here but in the aforementioned phylogenetic studies, Lupella is nested among the species of Achelous suggesting only a single genus may be warranted. Diagnosis. Carapace subcircular to transversely ovate; front multilobate, in transverse or convex row; anterolateral margin with 9 teeth, last markedly longer than others, directed laterally; junction of posterior and posterolateral borders forming curved ridge (usual), or acutely angular, or spiniform. Eyestalk short, stout, shorter than one-fourth carapace length. Antennal basal article narrow; flagellum standing in orbit. Maxilliped 3 merus flat, oblong, elongated distally, distal half little narrower than proximal half; palp placed near midlength of inner meral margin. Cheliped

carpus with inner and outer spine; palm prismatic, ridged, surface variously granular. Pereopods 2–4 longer than carapace. Pereopod 5 a swimming leg; merus ovate, less than twice as long as wide, wider than pereopod 4 merus; propodus and dactylus elongate-ovate. Achelous De Haan, 1833 Diagnosis. Carapace dorsal surface with granules distributed in patches. Cheliped fingers as long as or shorter than palm in both sexes. Pereopod 5 merus upper margin without distal spine, lower margin with or without distal spine. Maximum cl. 52 mm (Pl. 61k). Intertidal–slope (0–640 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific, Temperate South America. 27 species (Koch et al. 2022: diagnosis, species list; Rathbun 1930: key to 15 American species as subgenus of Portunus; Rodrigues et al. 2017: key to 9 Brazilian species).

Lupella Rathbun, 1897 Diagnosis. Carapace dorsal surface evenly granular. Cheliped fingers more than 3 times palm length in adult males, 1.5 times in adult females. Pereopod 5 merus upper and lower margins with distal spines. Maximum cl. 33 mm (Pl. 61l). Subtidal, shelf (4–46 m). Tropical Atlantic (West Indies). 1 species (Mantelatto et al. 2009: relationship to Portunus s.l.; Rathbun 1930: rediagnosis, figure). Lupella was synonymised with Achelous by Koch et al. (2022).

Subfamily Carupinae Paulson, 1875 The carupines (catoptrines of earlier literature, e.g. ­Stephenson 1972) are a morphologically diverse assemblage of genera exhibiting varying degrees of secondary modification of pereopod 5 from a swimming (e.g. Carupa)

Fig. 14.122.  Portunidae, Achelouinae. Carapace, pereopods: a, Achelous; b, Lupella (male). Pereopod 1, carpus, manus: c, Achelous. Maxilliped 3: d, Achelous.

14 – Brachyura – crabs

to walking leg, presumably in adaptation to cavernicolous habitats such as deep subterranean rubble or anchialine karst crevices (Fujita and Naruse 2011; Ng 2011; Ng and Takeda 2003); some of these genera also exhibit corresponding eye reduction. A common feature of carupines is the slender pereopod 5 with an elongated slender merus, in contrast to the stout, swollen, more muscular merus of most other portunids. Karasawa et  al. (2008) regarded the carupines (as composed here) as polyphyletic, but Spiridonov 2014 Spiridonov et al. (2014) and Evans (2018) found a monophyletic Carupinae as sister to Scylla (Necronectinae). Separating Catoptrus from Libystes has never been straightforward and their current genus-level taxonomy is widely acknowledged as inadequate (Apel and Spiridonov 1998; Ng 2011; Serène 1966; Stephenson 1972) or even discounted (Stephenson and Campbell 1960). Distinctions between Catoptrus and Libystes identified here are subtle and based on our assessment of constituent species within their current generic placement. Diagnosis. Carapace front not markedly protruding beyond general outline of anterior margin, lobes forming

705

more-or-less even arc; junction of posterior and posterolateral borders forming curved ridge. Eyestalk short, stout, shorter than one-fourth carapace length. Antennal basal article as long as or longer than wide; flagellum standing in orbit. Cheliped carpus with 0–2 spines on outer face. Pereopods 2–4 longer than carapace. Pereopod 5 a swimming leg; merus slender, twice as long as wide or more; propodus and dactylus ovate-lanceolate. Implicit generic attributes, Carupinae. Junction of posterior and posterolateral borders forming curved ridge. Cheliped palm smooth; dactylus tip simple. Pereopod 5 a swimming leg; merus without distolateral spine; dactylus ovate with distal point. Atoportunus Ng & Takeda, 2003 Diagnosis. Carapace transversely ovate; front broadly, sinuously bilobed, separated by median notch; anterolateral margin with 6 teeth. Antennal basal article longer than wide. Maxilliped 3 merus distolateral margin weakly produced, evenly rounded. Cheliped carpus outer face without spines; merus with spines on anterior margin; fingers slender, occlusal margins lined with obliquely

Fig. 14.123.  Portunidae, Carupinae. Carapace: a, Atoportunus; b, Carupa; c, Catoptrus; d, Kume; e, f, Laleonectes (with posterior view); g, Libystes; h, Pele; i, Richerellus. Pterygostomium, stridulatory ridge: j, Laleonectes. Cheliped; k, l, Atoportunus; m, Carupa; n, Catoptrus; o, p, Richerellus. Pereopod 5: q, Atoportunus; r, Catoptrus; s, Richerellus. Maxilliped 3: t, Catoptrus; u, Kume; v, Libystes. Male pleonites 3/4–6, telson: w, Catoptrus; x, Pele; y, Richerellus.

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Marine Decapod Crustacea

directed needle-like spines. Pereopod 5 propodus and dactylus ovate-lanceolate. Maximum cl. 19 mm. Subtidal, shelf (2–50 m; deep rubble crevices and caves). IndoWest Pacific. 3 species (Ng and Takeda 2003; Takeda 2003).

Carupa Dana, 1851 Diagnosis. Carapace transversely ovate; front with 4 lobes, not markedly protruding beyond general outline of anterior margin; anterolateral margin with 7 teeth. Antennal basal article longer than wide. Cheliped carpus inner margin with distinct spine; merus with spines on anterior margin; fingers robust, occlusal margins with obtuse teeth. Pereopod 5 propodus and dactylus ovate-lanceolate. Maximum cl. 38 mm (Pl. 61m). Intertidal–shelf (0–100 m; mainly in crevices in coral reefs and rubble, submarine caves). Indo-West Pacific. Carupa tenuipes Dana, 1852 introduced to Mediterranean Sea (Galil 2004). 2 species (Apel and Spiridonov 1998: rediagnosis; Leene 1940: figures; Takeda 1993).

Catoptrus A. Milne-Edwards, 1870 Diagnosis. Carapace transversely ovate, ~1.6 times as wide as long; front transverse; anterolateral margin sharply defined, with teeth; posterior margin broadly convex between coxae of pereopods 5, narrower than fronto-orbital width. Antennal basal article longer than wide, or as long as wide. Maxilliped 3 merus distolateral margin weakly produced, evenly rounded. Cheliped carpus inner and outer margins unarmed; merus usually with 2 spines on anterior margin; fingers slender or robust, occlusal margins irregularly toothed. Pereopod 5 propodus and dactylus flattened, dactylus narrowly lanceolate to spatulate. Male telson little longer than wide. Maximum cl. 22 mm. Intertidal–slope (0–326 m; submarine caves or reef crevices). Indo-West Pacific. 8 species (Fujita and Naruse 2011; Naruse and Uyeno 2021: differences between species; Serène 1966: rediagnosis).

Kume Naruse & Ng, 2012 Diagnosis. Carapace transversely broad, twice as wide as long; front transverse; anterolateral margin with 5 blunt teeth. Maxilliped 3 merus distolaterally angular, produced, margins prominently granular. Cheliped carpus inner and outer margins unarmed; merus anterior margin without spines; fingers slender, occlusal margins regularly toothed. Pereopod 2 about twice carapace length. Pereopod 5 propodus and dactylus slender, dactylus lanceolate. Male telson little longer than wide. Maximum cl. 3 mm. Shelf (55 m). Central Indo-Pacific. (Ryukyu Is). 1 species (Naruse and Ng 2012).

Laleonectes Manning & Chace, 1990 Diagnosis. Carapace transversely broad, twice as wide as long; front lobes in transverse row; anterolateral margin with 9 teeth, last longest, directed laterally; junction of posterior and posterolateral borders forming upcurved spine. Pterygostomial region

with stridulatory ridges. Antennal basal article longer than wide. Maxilliped 3 merus distolateral margin strongly produced, auriculiform. Cheliped carpus with 2 spines and several acute granules on outer face; merus with spines on anterior margin; palm with small spines, granules and transverse carinae; fingers robust, occlusal margins with triangular teeth. Pereopod 5 propodus and dactylus elongate-ovate. Maximum cl. 38 mm (Pl. 61n). Subtidal–slope (6–380 m). Tropical Atlantic, Indo-West Pacific. 4 species (Mendoza and Devi 2017: species compared).

Libystes A. Milne-Edwards, 1867 Diagnosis. Carapace transversely subrectangular, 1.5–1.9 (usually 1.6) times as wide as long; front transverse; anterolateral margin bluntly defined, teeth present or absent; posterior margin straight between coxae of pereopods 5, as wide or wider than as frontoorbital width. Antennal basal article longer than wide, or as long as wide. Maxilliped 3 merus distolateral margin strongly produced, auriculiform. Cheliped carpus inner and outer margins unarmed; merus anterior margin without spines. Pereopod 5 propodus and dactylus flattened, dactylus narrowly arcuate to spatulate. Male telson about as long as wide. Maximum cl. 12 mm. Intertidal–shelf (0–200 m). Indo-West Pacific, Temperate Southern Australia (S Qld). 9 species (Spiridonov et al. 2021: rediagnosis, species compared; Serène 1966: rediagnosis).

Pele Ng, 2011 Diagnosis. Carapace transversely broad, twice as wide as long; front weakly biconvex, almost transverse; anterolateral margin weakly or not toothed. Antennal basal article longer than wide. Maxilliped 3 merus weakly produced, evenly rounded. Cheliped carpus inner and outer margins unarmed; merus with distal anterior spine; fingers slender or robust, occlusal margins irregularly toothed. Pereopod 2 ~3 times carapace length. Pereopod 5 propodus and dactylus slender, dactylus lanceolate. Male telson twice as long as wide. Maximum cl. 6 mm. Marine anchialine caves (0–1 m). Eastern Indo-Pacific. (Hawaii). 1 species (Ng 2011).

Richerellus Crosnier, 2003 Diagnosis. Carapace transversely ovate, ~1.4 times as wide as long; front broadly bilobed, separated by median notch; anterolateral margin with 4 teeth. Antennal basal article longer than wide. Maxilliped 3 merus distolateral margin weakly produced, evenly rounded. Cheliped carpus inner margin with distinct spine; merus anterior margin without spines; fingers robust, occlusal margins with obtuse teeth. Pereopod 5 propodus and dactylus elongate-ovate. Maximum cl. 12 mm. Subtidal, shelf (15–50 m). Central Indo-Pacific. (New Caledonia). 1 species (Crosnier 2003).

Subfamily Coelocarcininae Števčić, 2005 Coelocarcinines are small coral reef crabs whose carapace shape and surface texture provide good camouflage

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Coelocarcinus Edmondson, 1930 Maximum cl. 9 mm. Intertidal, subtidal (0–15 m; coral reefs). 2 species (Edmondson 1930).

Subfamily Lupocyclinae Alcock, 1899

Fig. 14.124.  Portunidae, Coelocarcininae, Coelocarcinus. a, carapace; b, cheliped; c, pereopod 5.

amongst rubble and Halimeda fragments on coral reefs. They uniquely have near-circular dactyli and propodi of pereopod 5. Coelocarcinus has been variously regarded as a caphyrine (now Thalamitinae) (Edmondson 1930; Ng 2002; Števčić 2005) or carcinid (Evans 2018). The antennal structure, however, excludes Coelocarcinus from Thalamitinae, and a carcinid placement received weak support in phylogenetic analysis (Evans 2018). Given its ambiguous phylogenetic position, and because the antennal structure and cheliped length (longer than the walking legs) are consistent with Portunidae, Coelocarcininae is treated here as an independent portunid subfamily. Diagnosis. Carapace ovate; dorsal surface irregularly rugose; front prominently protruding beyond general outline of anterior margin; junction of posterior and posterolateral borders forming curved ridge. Eyestalk short, stout, ~0.1 carapace length. Cheliped carpus with 0–2 spines on outer face. Pereopods 2–4 shorter than carapace. Pereopod 5 a swimming leg; propodus and dactylus subcircular.

Lupocyclinae include two Indo-West Pacific genera formerly placed in Portuninae (Spiridonov et  al. 2014). Although Karasawa et  al. (2008) included Carupella Lenz & Strunck, 1914 in Lupocyclinae, that genus is based on early juveniles probably of existing portunine genera (Spiridonov 2020) and are therefore excluded here. Koch et  al. (2022) placed the genus Alionectes (containing two former species of Xiphonectes) in Lupocyclinae based on the posteriorly spined pereopod 5 merus and the results of phylogenetic analysis, albeit equivocally supported. Given the otherwise strong similarities to other portunine genera, we tentatively place Alionectes in Portuninae. Further study may even show that Lupocyclinae itself should be subsumed into Portuninae. Diagnosis. Carapace subcircular to transversely ovate; front multilobate, in transverse or convex row; junction of posterior and posterolateral borders forming curved ridge. Eyestalk short, stout, shorter than one-fourth carapace length. Antennal basal article narrow; flagellum standing in orbit. Maxilliped 3 merus not elongated distally; palp placed near inner distal meral margin. Cheliped carpus with 1 or 2 spines on outer face. Pereopods 2–4 longer than carapace. Pereopod 5 a swimming leg; merus ovate to subrectangular, length usually less than twice width; lower margin with subdistal spine; propodus and dactylus ovate. Lupocycloporus Alcock, 1899 Diagnosis. Carapace transversely ovate, width 1.5–1.7 times as wide as long; anterolateral margin with 8 equal teeth, ninth tooth distinctly longer. Antennal basal article with distolateral lobule.

Fig. 14.125.  Portunidae, Lupocyclinae. Carapace: a, Lupocycloporus; b, Lupocyclus. c, Lupocyclus rotundatus Adams & White, 1849. d, maxilliped 3, Lupocycloporus.

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Marine Decapod Crustacea

Cheliped carpus with inner and outer spine. Pereopod 5 merus ovate, twice as long as wide. Maximum cl. 28 mm. Subtidal, shelf (12–140 m). Western and Central Indo-Pacific, Temperate Australasia. 8 species (Moosa 1981b: species differences).

Lupocyclus Adams & White, 1849 Diagnosis. Carapace subcircular to narrowly ovate, 1.1–1.3 times as wide as long; anterolateral margin with 5 equal teeth or 9 teeth of alternating size. Antennal basal article without distolateral lobule. Cheliped carpus with inner spine and 1 or 2 spines on outer face. Pereopod 5 merus ovate to subrectangular, usually less than twice as long as wide. Maximum cl. 37 mm. Subtidal, shelf (13–174 m). Western and Central Indo-Pacific, Temperate Australasia. 7 species (Leene 1940: figures of 4 species; Moosa 1981a: figures of 2 species).

chelipeds and other features. The diagnosis of Necronectinae below is based largely on Scylla. Diagnosis. Carapace transversely ovate; dorsal surface smooth, at most with faintly indicated epibranchial and gastric ridges; front with 4 teeth in transverse row in addition to supraorbital teeth; junction of posterior and posterolateral borders forming curved ridge. Eyestalk stout, shorter than 0.1 carapace length. Antennal basal article narrow; flagellum standing in orbit. Maxilliped 3 merus not elongated distally; palp placed near inner distal meral margin. Cheliped carpus with 0–2 spines on outer face; palm inflated, ovate in cross-section, surface smooth. Pereopods 2–4 longer than carapace. Pereopod 5 a swimming leg; merus ovate, unarmed, ~1.5 times as long as wide; propodus and dactylus ovate. Scylla De Haan, 1833

Subfamily Necronectinae Glaesnner, 1928 The subfamily includes the fossil genus Necronectes A. Milne-Edwards, 1881, and the Recent Indo-West Pacific genus, Scylla, united by the smooth and inflated chela, and very smooth carapace with similar marginal spination (Schweitzer et al. 2006). Necronectines resemble portunines in most respects, but are reminiscent of some carupines in their smooth, inflated, cheliped palm (rather than prismatic and carinate) and smooth carapace surface. Indeed, the similarity may reflect common ancestry given sister relationship between Necronectinae and Carupinae (Evans 2018; Spiridonov et  al. 2014). Species of Scylla reach large sizes and are commercially important wherever they occur (Keenan et al. 1998). The West African genus Sanquerus has most recently been placed in Necronectinae (e.g. Karasawa et  al. 2008; Spiridonov 2020) although its chelipeds are prismatic and carinate. Although the molecular phylogenetic position of Sanquerus is currently unclear, a close relationship to Scylla is not supported (Mantelatto et al. 2018). Sanquerus is herein returned to Portuninae on the basis of its portunine-like

mud crab, mangrove crab Diagnosis. Carapace transversely ovate; dorsal surface smooth, epibranchial ridges short, indistinct; anterolateral margin with 9 equal teeth. Antennal basal article with distolateral lobule. Cheliped carpus with inner spine, outer face with 1 or 2 spines. Maximum cl. 200 mm (Pl. 61o). Estuarine, intertidal, subtidal. Indo-West Pacific, Temperate Australasia. 4 species (Keenan et al. 1998: key to species). All species are fished commercially throughout their range (Dumas et al. 2012; Fondo et al. 2010; Jumawan et al. 2021; Le Vay 2001; Rahman et al. 2020). Scylla serrata (Forsskål, 1775) has been introduced to the Caribbean coast of Colombia (Lemaitre et al. 2013).

Subfamily Podophthalminae Dana, 1851 Podophthalmines are easily recognised by their long, stalked eyes. Three genera are known, but it is not clear whether the eastern Pacific genus Euphylax is closely related to the IndoWest Pacific genera, or whether the ocular similarities are convergent (Evans 2018; Garth and Stephenson 1966). Molecular phylogenetic analyses are currently ambiguous about podophthalmine monophyly but the latter study some results point to a possible close relationship between the American Euphylax and the West African portunine, Sanquerus (Evans 2018; Mantelatto et  al. 2018). Species of Euphylax and Podophthalmus are fished commercially. Diagnosis. Junction of posterior and posterolateral borders forming curved ridge. Eyestalk extremely long, slender, exceeding half carapace length. Cheliped carpus with 0–2 spines on outer face. Pereopod 5 a swimming leg. Euphylax Stimpson, 1860

Fig. 14.126.  Portunidae, Necronectinae, Scylla. a, carapace; b, cheliped.

Diagnosis. Anterolateral margin with 3–5 spines. Eyestalk slightly exceeding half carapace length. Maximum cl. 61 mm.

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Fig. 14.127.  Portunidae, Podophthalminae. a, Euphylax dovii Stimpson, 1860; b, Podophthalmus nacreus Alcock, 1899; c, Vojmirophthalmus minabensis (Sakai, 1961).

Intertidal–shelf (0–64 m). Tropical Eastern Pacific, Temperate South America (E Pacific coast). 2 species (Rathbun 1930: key to species). Euphylax dovii Stimpson, 1860 is pelagic and can occur in dense swarms (Hearn et al. 2020).

Podophthalmus Lamarck, 1801 Diagnosis. Anterolateral margin with 2 spines. Eyestalk as long as or longer than carapace, simple, without wing-like expansion next to cornea. Maximum cl. 60 mm (Pl. 62a). Subtidal, shelf (10–127 m). Western and Central Indo-Pacific, Eastern Indo-Pacific, Temperate Australasia. 2 species (Apel and Spiridonov 1998: rediagnosis).

Vojmirophthalmus Števčić, 2011 petal-eyed swimmer crab Diagnosis. Anterolateral margin with 2 spines. Eyestalk as long as or longer than carapace, with wing-like expansions next to cornea. Maximum cl. 23 mm. Subtidal, shelf (15–49 m). Central Indo-Pacific. 1 species (Bruce 1965: redescription, figure; Števčić 2011: diagnosis; Yosuva et al. 2020: figure).

Subfamily Portuninae Rafinesque, 1815 Ongoing phylogeny-based studies of the portunids (e.g. Evans 2018; Karasawa et  al. 2008; Mantelatto et  al. 2018; Spiridonov 2020; Spiridonov et  al. 2014) have refined the

composition of Portuninae by progressive removal of the achelouines, lupocyclines and thalamitines. With present re-inclusion of Sanquerus (most recently considered a necronectine; Spiridonov 2020), Portuninae now includes eight genera. All have slender, prismatic chelae and many have long, laterally directed last anterolateral spines on the carapace. All species now included in Portuninae are freeliving, ‘true’ swimming crabs in which pereopod 5 is natatory. Many species are comparatively small, with carapace length less than 20 mm, but the portunines also include some of the most important fishery species globally, such as the Blue Crab (Callinectes sapidus Rathbun, 1896) (Kennedy and Cronin 2007) in the Atlantic, and the Flower Crab (Portunus pelagicus (Linnaeus, 1758)) and Blue Swimmer Crab (Portunus armatus (A. Milne-Edwards, 1861)) in the IndoPacific (Lai et al. 2010). Phylogenetic knowledge of the portunine genera is far from complete. Arenaeus, Callinectes and Portunus form a core group (Evans 2018; Mantelatto et  al. 2018), but how these relate to the remaining genera is currently unclear given equivocal results in studies to date (Evans 2018; Mantelatto et al. 2018). Monomia, Cycloachelous and Xiphonectes (formerly subgenera of Portunus) (Stephenson 1972) are now considered independent genera (Davie 2002; Spiridonov et al. 2014), and Sanquerus (formerly in Portunus) and Cavoportunus (formerly in Cycloachelous) were relatively recently proposed (Manning 1989; Nguyen and Ng 2010). Koch et al. (2022) split the heterogeneous and polyphyletic

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Marine Decapod Crustacea

Fig. 14.128.  Portunidae, Portuninae. a, Arenaeus mexicanus (Gerstaecker, 1856); b, Sanquerus validus (Herklots, 1851). Carapace: c, Alionectes; d, Allomonomia; e, Callinectes; f, Cavoportunus; g, Cycloachelous; h, Eodemus; i, Incultus; j, Monomia; k, Portunus; l, Trionectes; m, Xiphonectes. Cheliped: n, Callinectes; o, Incultus; p, Monomonia; q, Xiphonectes. r, orbits, antennules, antennae, epistome, endostome (maxillipeds 3 removed), Portunus. Maxilliped 3: s, Alionectes; t, Cycloachelous; u, Eodemus; v, Monomia; w, Portunus; x, Xiphonectes. y, male thoracic sternum, left gonopod 1 in situ, Cavoportunus. z, gonopod 1, Cycloachelous.

genera Monomia and Xiphonectes (Koch et al. 2015a; Spiridonov et  al. 2014), creating four new genera. Of these, Alionectes was referred to Lupocyclinae based on the posteriorly spined pereopod 5 merus and the results of phylogenetic analysis, albeit equivocal. Pending more decisive phylogenetic analyses, we include Alionectes in Portuninae because of its strong similarities to other genera placed there, especially Xiphonectes and allies.

Diagnosis. Carapace subcircular to transversely ovate; front multilobate, in transverse or convex row; junction of posterior and posterolateral borders forming curved ridge, or acutely angular, or spiniform. Eyestalk short, stout, shorter than one-fourth carapace length. Antennal basal article narrow; flagellum standing in orbit. Cheliped carpus with 0–2 spines on outer face; palm prismatic, ridged, surface variously granular. Pereopods 2–4 longer than

14 – Brachyura – crabs

carapace. Pereopod 5 a swimming leg; merus ovate, less than twice as long as wide, wider than pereopod 4 merus, lower margin distally unarmed, or if subdistal spine present carapace with acute junction of posterolateral and posterior borders; propodus and dactylus elongate-ovate. Implicit generic attributes, Portuninae. Junction of posterior and posterolateral borders forming curved ridge. Maxilliped 3 palp placed near inner distal meral margin. Cheliped carpus with inner spine. Alionectes Koch, Spiridonov & Ďuriš, 2022

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Maxilliped 3 merus distolaterally rounded. Cheliped carpus without inner spine. Pereopod 5 merus without distolateral spine. Maximum cl. 99 mm (Pl. 62c). Estuarine, intertidal–shelf (0–92 m). Temperate Northern and Tropical Atlantic, Tropical Eastern Pacific. 15 species (Williams 1974: key to 14 species). The blue crab Callinectes sapidus (Rathbun, 1896) is the basis of a valuable commercial fishery along the estuaries of the eastern Americas. The same species has occurred sporadically on the W Atlantic coast but is considered an invasive species in the Mediterranean Sea (Fuentes et al. 2019).

Cavoportunus Nguyen & Ng, 2010

Diagnosis. Carapace front with 4 teeth, median/s shorter than laterals; dorsal surface with granules distributed in patches; anterolateral margin with 8 low angular teeth, and long, spiniform last tooth, directed posterolaterally; junction of posterior and posterolateral borders acutely angular or spiniform. Maxilliped 3 merus distally rounded, not produced distally beyond palp. Cheliped merus posterior distal margin with 2 spines. Pereopod 5 merus without distolateral spine. Maximum cl. 12 mm. Subtidal–slope (3–834 m). Western and Central Indo-West Pacific, 2 species (Koch et  al. 2022). The species were formerly placed in Xiphonectes.

Diagnosis. Carapace subcircular-hexagonal, ~1.3 or less times as wide as long; dorsal surface with granules distributed in patches; anterolateral margin with 9 teeth, last subequal to others. Male sternopleonal cavity with pair of hemispherical cavities to accommodate large lateral subdistal lobe of gonopod 1. Maxilliped 3 merus distolaterally produced as angular or auriculiform lobe, not elongated distally. Pereopod 5 merus without distolateral spine. Maximum cl. 20 mm. Shelf, slope (20–245 m). Indo-West Pacific. 1 species (Nguyen and Ng 2010).

Allomonomia Koch, Spiridonov & Ďuriš, 2022

Cycloachelous Ward, 1942

Diagnosis. Carapace narrowly ovate, 1.5 times as wide as long; dorsal surface with granules distributed in patches; front with 4 projecting subequal teeth projecting distinctly beyond inner orbital angle; anterolateral margin with 9 teeth, last longer than others, directed anterolaterally to slightly posterolaterally. Maxilliped 3 merus distolaterally produced as angular or auriculiform lobe, not elongated distally. Cheliped merus posterior margin with 1 distal spine. Pereopod 5 merus without distolateral spine. Maximum cl. 25 mm. Shelf (50–150 m). Indo-West Pacific, Temperate Australasia. 2 species (Koch et  al. 2015a: comparison of species as Monomia; Koch et al. 2022).

Diagnosis. Carapace subcircular-hexagonal, ~1.4 or less times as wide as long; dorsal surface with granules distributed in patches; anterolateral margin with 9 teeth, last smaller to only slightly larger than others. Male sternopleonal cavity evenly V-shaped; gonopod 1 without large lateral distal lobes. Maxilliped 3 merus distolaterally produced as angular or auriculiform lobe, not elongated distally. Pereopod 5 merus without distolateral spine. Maximum cl. 24 mm. Intertidal–shelf (0–120 m). Indo-West Pacific, Temperate Australasia. 9 species (Koch et al. 2022: figures of 2 species; Koch 2021; Ward 1942: diagnosis).

Arenaeus Dana, 1851

Eodemus Koch, Spiridonov & Ďuriš, 2022

Diagnosis. Carapace dorsal surface microscopically and evenly granular; epibranchial ridges faintly indicated; dorsal orbital fissures open, narrowly V-shaped; anterolateral margin with 9 teeth, last longest, directed laterally. Endostomial ridges absent. Maxilliped 3 merus distolaterally rounded. Pereopod 5 merus without distolateral spine. Maximum cl. 65 mm (Pl. 62b). Intertidal–shelf (0–68 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Zupolini et al. 2017: key to species).

Diagnosis. Carapace front with 3 or 4 teeth, median/s shorter than laterals; dorsal surface with granules distributed in patches; anterolateral margin with 8 sharp teeth, and long, spiniform last tooth, directed laterally or posterolaterally; junction of posterior and posterolateral borders acutely angular or spiniform. Maxilliped 3 merus strongly produced laterally as angular pointed lobe. Cheliped merus posterior distal margin with 2 spines. Pereopod 5 merus without distolateral spine. Maximum cl. 23 mm. Intertidal–shelf (0–205 m). Western and Central Indo-West Pacific, Temperate Australasia. 6 species (Koch et al. 2022). The species were formerly placed in Xiphonectes.

speckled crab

Callinectes Stimpson, 1860 blue crab Diagnosis. Carapace dorsal surface distinctly and evenly granular; anterolateral margin with 9 teeth, last longest, directed laterally.

Incultus Koch, Spiridonov & Ďuriš, 2022 Diagnosis. Carapace front with 3 or 4 teeth, if 3 teeth present, median shorter than laterals; dorsal surface with granules

712

Marine Decapod Crustacea

distributed in patches; anterolateral margin with 8 blunt lobiform teeth, and long, spiniform last tooth, directed laterally or posterolaterally; junction of posterior and posterolateral borders acutely angular or spiniform. Maxilliped 3 merus distally produced well beyond palp as blunt lobe, about half-width of proximal half. Cheliped merus posterior distal margin with 1 spine; fingers shorter than dorsal margin of palm. Pereopod 5 merus without distolateral spine. Maximum cl. 29 mm. Intertidal–slope (0–580 m). Indo-West Pacific. 3 species (Koch et  al. 2022). The species were formerly placed in Xiphonectes.

Monomia Gistel, 1848 Diagnosis. Carapace transversely ovate to subhexagonal, ~1.7 times as wide as long; dorsal surface with granules distributed in patches; front with 4 low teeth, median pair shorter, projecting little beyond inner orbital angle; anterolateral margin with 9 teeth, last longer (sometimes markedly longer) than others, directed anterolaterally to slightly posterolaterally. Maxilliped 3 merus distolaterally produced as angular or auriculiform lobe, not elongated distally. Cheliped merus posterior margin with 2 distal spines. Pereopod 5 merus without distolateral spine. Maximum cl. 51 mm. Intertidal–slope (0–402 m). Indo-West Pacific, Temperate Australasia. 10 species (Koch and Ďuriš 2018; recent species: Koch et al. 2015b).

Portunus Weber, 1795 Diagnosis. Carapace dorsal surface distinctly and evenly granulate, rugose; metagastric, mesogastric and epibranchial ridges distinct; dorsal orbital fissures closed, margins appressed; anterolateral margin with 9 teeth, last longer (sometimes markedly longer) than others, directed anterolaterally to slightly posterolaterally. Endostomial ridges present. Maxilliped 3 merus distolaterally rounded or bluntly obtuse, not produced laterally. Pereopod 5 merus without distolateral spine. Maximum cl. 95 mm (Pl. 62d). Intertidal–shelf (0–65 m). Indo-West Pacific, Temperate ­Australasia. 13 species. The genus Portunus has been widely used for swimming crabs throughout the world but most Atlantic species have been transferred to other genera (Mantelatto et al. 2018 Koch et al. 2022). Portunus serratifrons (Montrouzier, 1865) is a juvenile of Scylla serrata (fide Milne-Edwards 1873). Keys to species from the Indo-West Pacific (e.g. Crosnier 1962; Stephenson and Campbell 1959) include few species now retained in this genus.

Sanquerus Manning, 1989 Diagnosis. Carapace dorsal surface microscopically, evenly granulate, appearing smooth; metagastric, mesogastric and epibranchial ridges scarcely discernible; dorsal orbital fissures closed, margins appressed; anterolateral margin with 9 teeth, last markedly longer than others, directed laterally. Endostomial ridges present. Maxilliped 3 merus distolaterally rounded or

bluntly obtuse, not produced laterally. Pereopod 5 merus without distolateral spine. Maximum cl. 93 mm (Pl. 62e). Intertidal–shelf (0–55 m). Tropical E Atlantic. 1 species (Capart 1951: figure; Manning 1989). Sanquerus validus (Herklots, 1851) is an important fishery species off West Africa (Fransen 2014).

Trionectes Koch, Spiridonov & Ďuriš, 2022 Diagnosis. Carapace front with 3 teeth, median as long as or longer than laterals; dorsal surface with granules distributed in patches; anterolateral margin with 8 spiniform or lobiform teeth, and long, spiniform last tooth, directed laterally or posterolaterally; junction of posterior and posterolateral borders acutely angular or spiniform. Maxilliped 3 merus distally produced well beyond palp as blunt lobe, about half-width of proximal half. Cheliped merus posterior distal margin with 1 or 2 spines. Pereopod 5 merus without distolateral spine; fingers as long as dorsal margin of palm. Maximum cl. 25 mm. Subtidal–shelf (3–71 m). Western and Central Indo-West Pacific, Temperate Australasia. 7 species (Koch et al. 2022). The species were formerly placed in Xiphonectes.

Xiphonectes A. Milne-Edwards, 1873 Diagnosis. Carapace front with 3 or 4 teeth, if 3 teeth present, median shorter than laterals; dorsal surface with granules distributed in patches; anterolateral margin with 6–9 sharp, spiniform or triangular teeth, last markedly longer than others, directed laterally or posterolaterally; junction of posterior and posterolateral borders acutely angular or spiniform. Maxilliped 3 merus distally produced well beyond palp as blunt lobe, about half-width of proximal half. Cheliped merus posterior distal margin with 1 spine; fingers as long as dorsal margin of palm. Pereopod 5 merus with or without distolateral spine. Maximum cl. 16 mm. Intertidal–slope (0–320 m). Indo-West Pacific, Temperate Australasia. 14 species (Spiridonov 2016: 2 species compared; Stephenson and Campbell 1959: key to 28 species of Portunus includes 8 now in Xiphonectes). Most species were described as species of Portunus.

Subfamily Thalamitinae Paulson, 1875 Thalamitines differ from portunines by features of the antenna, chiefly by the exclusion of the peduncle from the orbit (except in Charybdis (Gonioneptunus)) and the wide basal article (except in Caphyra and Lissocarcinus). Most thalamitines are aggressive, active and agile, and many reach large sizes, especially species of Charybdis that are commercially fished, such as C. feriata (Linnaeus, 1758) and C. natator (Herbst, 1794). The generic composition of Thalamitinae has been largely stable but Evans (2018) confirmed the enigmatic genus Cronius as a thalamitine, rather than portunine (Manning 1989; Mantelatto et al. 2009) and that Caphyra and Lissocarcinus (formerly comprising

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Fig. 14.129.  Portunidae, Thalamitinae. Carapace: a, Caphyra; b, Charybdis (Charybdis); c, Charybdis (Goniohellenus); d, Charybdis (Gonioneptunus); e, Cronius; f, Gonioinfradens; g, Goniosupradens; h, Lissocarcinus; i, Thalamita exetastica Alcock, 1899: j, Thalamita investigatoris Alcock, 1899; k, Thalamitoides; l, Zygita. Antennal basal article (b), free articles, orbit (o), antennular fossa (dashed line): m, Charybdis (Gonioneptunus); n, Charybdis (Charybdis). Cheliped, carpus–fingers, upper view: o, Goniosupradens; p, Lissocarcinus. Cheliped palm, inner view: q, Thalamitoides. Cheliped palm, outer view: r, Zygita. Pereopod 4: s, Zygita. Pereopod 5: t, Caphyra; u, Lissocarcinus.

Caphyrinae), were derived from within Thalamitinae. Unlike their free-living relatives, Caphyra and Lissocarcinus are variously associated with cnidarians, echinoderms and the tropical green alga, Chlorodesmis. Moreover, in Caphyra, pereopod 5 is not a swimming leg, but secondarily modified for clinging to the host. Charybdis and Thalamita are the most diverse portunid genera, with the latter particularly common on coral reef flats of the Indo-West Pacific. Thalamita is often recognisable by the wide subtrapezoid carapace and wide-set eyes, being placed at the lateral extremities. Conversely, many species of Charybdis have a more ovate carapace with the eyes placed much further inward of the lateral margins. Distinctions between Thalamita and Charybdis, however, are not always obvious, because of intergrading of carapace shape and sometimes confusing patterns of carapace spination (Apel and Spiridonov 1998; Stephenson et al. 1957; Wee and Ng 1995). Although both genera are reciprocally monophyletic (Evans 2018), accurate identification requires careful observation of the anterolateral carapace spination. Unless otherwise stated in the keys and diagnoses, the counts for the anterolateral teeth include both the prominent teeth (‘large’) and much smaller teeth (‘minute’) that may lie

between the larger teeth or arise from the base of the posterior margin of the larger teeth. These ‘minute’ teeth can be concealed by detritus, which should be carefully removed with a soft paint brush or similar. In initiating revision of the diverse genus Thalamita, Evans (2018) resurrected the genus Thalamonyx A. MilneEdwards, 1873, and diagnosed three new genera for several clades within the genus: Zygita, Thranita, Trierarchus. Zygita is included here, but, unfortunately, Thalamonyx, Thranita and Trierarchus do not appear to be effectively recognisable when considered in the context of character suites present in the remaining species of Thalamita, and as such are not used here. Nevertheless, Thalamita is morphologically diverse and we acknowledge that Evans’ (2018) subdivisions may be vindicated when the genus can be evaluated synoptically and other species appropriately re-assigned. Diagnosis. Carapace subhexagonal; junction of posterior and posterolateral borders bluntly angular, without curved ridge. Eyestalk short, stout, shorter than one-fourth carapace length. Antennal basal article wider than long; flagellum excluded from orbit, at least in adults, or wider than long; flagellum standing in orbit (Charybdis

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(Gonioneptunus)). Cheliped carpus with 0–4 (usually 3) spines on outer face. Pereopods 2–4 longer than carapace, or shorter than carapace (Lissocarcinus). Pereopod 5 a swimming leg, or a walking leg. Implicit generic attributes, Thalamitinae. Junction of posterior and posterolateral borders forming curved ridge. Antennal basal article broad; flagellum excluded from orbit. Cheliped carpus with inner spine; merus without posterodistal spine; dactylus tip simple. Pereopod 5 a swimming leg; merus without distolateral spine; dactylus ovate with distal point. Caphyra Guérin, 1832 turtle-weed crab Diagnosis. Carapace subhexagonal; anterolateral margin weakly or not toothed. Antennal basal article narrow; flagellum excluded from orbit. Cheliped carpus outer face without spines. Pereopods 2–4 longer than carapace. Pereopod 5 a walking leg; dactylus slender, claw-like or lanceolate. Maximum cl. 14 mm. Intertidal, subtidal (0–18 m). Obligate commensals of alcyonarian corals and the green alga, Chlorodesmis. Indo-West Pacific, Temperate Australasia. 16 species (Stephenson and Campbell 1960: key to species; Stephenson and Rees 1968a: review, 4 species; Takeda and Webber 2006: recent species).

Subtidal–slope (10–420 m). Western and Central Indo-Pacific, Temperate Australasia. 11 species (Apel and Spiridonov 1998: rediagnosis, 6 species; Leene 1938: diagnosis, key to 8 species). Charybdis smithii periodically swarms near the surface over deep water (Türkay and Spiridonov 2006).

Charybdis (Gonioneptunus) Ortmann, 1893 Diagnosis. Carapace transversely subhexagonal; anterolateral margin with 6 large teeth, subequal or last longest; junction of posterior and posterolateral borders bluntly angular, without curved ridge. Antennal basal article broad; flagellum standing in orbit. Cheliped carpus outer surface with 3 spines; merus with posterodistal spine. Maximum cl. 27 mm. Subtidal–slope (6–439 m). Western and Central Indo-Pacific, Temperate Australasia. 5 species (Leene 1938: diagnosis, key to 4 species).

Cronius Stimpson, 1860 Diagnosis. Carapace transversely ovate; anterolateral margin with 9 teeth, alternating in size but all prominent. Cheliped carpus outer surface with 3 spines. Maximum cl. 63 mm. Intertidal–slope (0–300 m). Temperate Northern and Tropical Atlantic (Canaries and Madeira Island), Tropical Eastern Pacific. 1 species (Mantelatto et al. 2009: phylogeny; de Melo 1996: identification; Schäfer et al. 2019: distribution).

Charybdis (Charybdis) De Haan, 1833

Gonioinfradens Leene, 1938

Diagnosis. Carapace transversely ovate, subhexagonal; anterolateral margin with 6 large teeth or 5 large teeth and smaller tooth behind first large tooth, length of smaller tooth exceeding halflength of posterior margin of first tooth. Cheliped carpus outer surface with 3 spines. Maximum cl. 110 mm (Pl. 62f). Intertidal–slope (0–1203 m). Indo-West Pacific, Temperate Australasia. 44 species (Apel and Spiridonov 1998: rediagnosis, key to subgenera and 17 species; Leene 1938: diagnosis, key to subgenera, 33 species and varieties; Spiridonov and Türkay 2001: comparison of 6 species; Stephenson et al. 1957: key to 12 Australian species; Stephenson and Rees 1968b; Wee and Ng 1995: key to 20 species from Malayan Peninsula and Singapore). Several species have been introduced worldwide (Dessouassi et  al. 2019; Froglia 2012): C. feriata (Linnaeus, 1758), C. longicollis (Leene, 1938), C. lucifera (Fabricius, 1798) to the Mediterranean Sea; C. hellerii (A. Milne-Edwards, 1867) to the Mediterranean Sea, Gulf of Guinea, Brazil, Caribbean Sea and Hawaii; C. ­japonica (A. Milne-Edwards, 1867) to the Mediterranean Sea and New Zealand.

Diagnosis. Carapace transversely ovate, subhexagonal; anterolateral margin with 4 large teeth and 2 or 3 minute teeth, one each behind first, second and sometimes third large teeth. Cheliped carpus outer surface with 3 spines. Maximum cl. 53 mm. Intertidal–slope (0–300 m). Temperate Northern Atlantic, Indo-West Pacific. 2 species (Leene 1938: diagnosis, key to species). Gonioinfradens giardia (Nobili, 1906) has been introduced to the Mediterranean Sea (Galil et al. 2018).

Charybdis (Goniohellenus) Alcock, 1899 Diagnosis. Carapace transversely ovate, subhexagonal; anterolateral margin with 6 large teeth, subequal or last longest; junction of posterior and posterolateral borders bluntly angular, without curved ridge. Cheliped carpus outer surface with 3 spines; merus with posterodistal spine. Maximum cl. 53 mm.

Goniosupradens Davie, 2002 Diagnosis. Carapace transversely ovate; anterolateral margin with 5 large teeth and 1–3 minute teeth, one each behind first, usually second and sometimes third large teeth. Cheliped carpus outer surface with 3 or more spines. Maximum cl. 61 mm (Pl. 62g). Intertidal–shelf (0–30 m). Indo-West Pacific. 4 species (Apel and Spiridonov 1998: rediagnosis; Evans 2018: rediagnosis; Leene 1938: diagnosis, key to 3 species).

Lissocarcinus Adams & White, 1849 harlequin crab Diagnosis. Carapace subhexagonal, irregularly subcircular; anterolateral margin with 5 blunt teeth. Antennal basal article narrow; flagellum excluded from orbit. Cheliped carpus outer face without spines. Pereopods 2–4 longer than carapace, or shorter than

14 – Brachyura – crabs

carapace. Pereopod 5 a swimming leg; dactylus ovate, apex rounded or pointed. Maximum cl. 22 mm (Pl. 62h). Intertidal–shelf (0–205 m). Obligate commensals of holothurians, echinoderms and scleractinian corals (Caulier et  al. 2014). Indo-West Pacific, Temperate Australasia. 9 species (Apel and Spiridonov 1998: rediagnosis; Sakai 1976: key to 4 Japanese species; Stephenson and Campbell 1960: key to 2 Australian species).

Thalamita Latreille, 1829 Diagnosis. Carapace subhexagonal, subtrapezoid; anterolateral margin with 3–5 large teeth and sometimes with minute tooth behind first or third large tooth, length of minute teeth less than half length of posterior margin of adjacent large tooth, spinous margin demarcated from unarmed posterolateral margin by obtuse angle (if 5 large anterolateral teeth present, third large tooth slightly larger than fourth large tooth). Cheliped carpus outer surface with 3 or 4 spines; palm outer surface without spine near base of pollex, granular ridge present or absent; dactylus tip simple, or spoonshaped (in T. stephensoni Crosnier, 1962)). Pereopods 2–4 meri usually without ventrodistal spine. Maximum cl. 67 mm (Pl. 62i). Intertidal–slope (0–433 m). Tropical E Atlantic, Indo-West Pacific. 88 species (Apel and Spiridonov 1998: rediagnosis, key to 15 Arabian Gulf species; species-groups; Stephenson and Hudson 1957: diagnosis, key to 53 Indo-West Pacific species; Wee and Ng 1995: key to 17 species from Malayan Peninsular and Singapore). We treat Thalamonyx, Thranita and Trierarchus as junior synonyms of Thalamita.

Thalamitoides A. Milne-Edwards, 1869 Diagnosis. Carapace subtriangular to subtrapezoid; anterolateral margin with 3 or 4 slender teeth, spinous margin continuous with unarmed posterolateral margin, forming more-or-less straight or broadly curved margin. Cheliped carpus outer surface with 3 spines; dactylus tip spoon-shaped. Maximum cl. 15 mm (Pl. 62j). Intertidal–shelf (0–65 m). Indo-West Pacific. 3 species (­Crosnier 1962: key, figures of 2 species; Spiridonov and Neumann 2008: descriptions of 2 species).

Zygita Evans, 2018 Diagnosis. Carapace subhexagonal; anterolateral margin with 5 teeth, third large tooth slightly larger than fourth large tooth, spinous margin demarcated from unarmed posterolateral margin by obtuse angle. Cheliped carpus outer surface with 4 spines; palm outer surface with granular ridge terminating in small spine near base of pollex. Pereopods 2–4 meri with ventrodistal spine. Maximum cl. 14 mm (Pl. 62k). Subtidal, shelf (1–50 m). Usually associated with nephtheid soft corals. Indo-West Pacific. 2 species (Evans 2018). References Apel M, Spiridonov VA (1998) Taxonomy and zoogeography of the portunid crabs (Crustacea: Decapoda: Brachyura: Portunidae) of the Arabian Gulf. Fauna Arabica 17, 159–331.

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Mantelatto FL, Robles R, Wehrtmann IS, Schubart CD, Felder DL (2018) New insights into the molecular phylogeny of the swimming crabs of the genera Portunus Weber, 1795 and Achelous De Haan, 1833 (Brachyura: Portunidae) of the Americas. Journal of Crustacean Biology 38, 190–197. doi:10.1093/jcbiol/rux119 Mendoza JCE, Devi SS (2017) A new species of the swimming crab genus, Laleonectes Manning & Chace, 1990 (Crustacea: Brachyura: Portunidae), from the western Indian Ocean. Zootaxa 4323, 219–228. doi:10.11646/zootaxa.4323.2.5 Milne-Edwards A (1873) Recherches sur la faune carcinologique de la Nouvelle-Calédonie. Nouvelles Archives du Muséum d’Histoire Naturelle, Paris 9, 155–322, pls 4–18. Moosa K (1981a) Crustacés Décapodes: Portunidae. In: Forest J (Ed.) Résultats des campagnes MUSORSTOM: I. Philippines (18–28 Mars 1976). Mémoires du Muséum National d’Histoire Naturelle (Série A, Zoologie) 91, 141–150. Moosa MK (1981b) Portunus (Lupocycloporus) wilsoni, a new portunid from the Indo-Australian region (Brachyura, Portunidae). Crustaceana 40, 26–30. doi:10.1163/156854081X00363 Naruse T, Ng PKL (2012) Kume tigra, a new genus and new species of carupine swimming crab (Crustacea: Brachyura: Portunidae) from Kume Island, Ryukyu Islands, Japan. Zootaxa 3367, 204–210. doi:10.11646/zootaxa.3367.1.19 Naruse T, Uyeno D (2021) Catoptrus lavicolus, a new species of swimming crab (Crustacea, Brachyura, Portunidae) from shallow subtidal lava rock field at Sakurajima, an active volcano in Kagoshima, southern Japan. Crustacean Research 50, 107–118. Ng PKL (1998) Crabs. In FAO Species identification guide for fishery purposes. The living marine resources of the Western Central Pacific. Volume 2. Cephalopods, crustaceans, holothurians and sharks. (Eds Carpenter KE, Niem VH) pp. 1045–1155. Food and Agriculture Organization, Rome. Ng PKL (2002) On the unusual swimming crab, Coelocarcinus foliatus Edmondson, 1930, with description of a new species from the Indian Ocean (Decapoda, Brachyura, Portunidae). Crustaceana 75, 51–60. doi:10.1163/156854002317373519 Ng PKL (2011) Pele ramseyi, a new genus and new species of anchialine swimming crab (Crustacea: Brachyura: Portunidae) from the Hawaiian Islands. Zootaxa 2737, 34–48. doi:10.11646/zootaxa.2737.1.3 Ng PKL, Takeda M (2003) Atoportunus, a remarkable new genus of cryptic swimming crab (Crustacea; Decapoda; Brachyura: Portunidae), with descriptions of two new species from the Indo-West Pacific. Micronesica 35–36, 417–430. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Nguyen TS, Ng PKL (2010) A new genus of the family Portunidae (Crustacea: Decapoda: Brachyura) and the identity of Portunus (Cycloachelous) yoronensis Sakai, 1974. Zootaxa 2677, 38–48. doi:10.11646/zootaxa.​ 2677.1.4 Rahman MM, Haque SM, Galib SM, Islam MA, Parvez MT, et al. (2020) Mud crab fishery in climate vulnerable coastal Bangladesh: an analysis towards sustainable development. Aquaculture International 28, 1243–1268. doi:10.1007/s10499-020-00523-2 Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Rodrigues IB, Cardoso IA, Serejo CS (2017) Catalogue and illustrated key of Achelous De Haan, 1833 and Portunus Weber, 1795 (Brachyura: Portunidae: Portuninae) species occurring in Brazilian waters. Nauplius 25, e2017005. doi:10.1590/2358-2936e2017005 Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo.

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Schäfer S, Monteiro J, Castro N, Rilov G, Canning-Clode J (2019) Cronius ruber (Lamarck, 1818) arrives to Madeira Island: a new indication of the ongoing tropicalization of the northeastern Atlantic. Marine Biodiversity 49, 2699–2707. doi:10.1007/s12526-019-00999-z Schweitzer CE, Iturralde-Vinent M, Hetler JL, Velez-Juarbe J (2006) Oligocene and Miocene decapods (Thalassinidea and Brachyura) from the Caribbean. Annals of the Carnegie Museum 75, 111–136. doi:10.2992/0097-4463(2006)75[111:OAMDTA]2.0.CO;2 Serène R (1966) Note sur les genres Catoptrus et Libystes et les Catoptrinae (Decapoda, Brachyura). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 37, 989–1000, pl. 1. Spiridonov VA (2016) Xiphonectes tuerkayi sp. n. from the Indian Ocean with notes on Xiphonectes longispinosus Dana, 1852 (Crustacea: Decapoda: Portunidae). Arthropoda Selecta 25, 357–372. doi:10.15298/ arthsel.25.4.03 Spiridonov VA (2020) An update of phylogenetic reconstructions, classification and morphological characters of extant Portunoidea Rafinesque, 1815 (Decapoda, Brachyura, Heterotremata), with a discussion of their relevance to fossil material. Geologija 63, 133–166. doi:10.5474/ geologija.2020.014 Spiridonov VA, Neumann V (2008) Coral-inhabiting swimming crabs (Crustacea, Decapoda, Portunidae) of the Sudanese Red Sea. Organisms, Diversity & Evolution 8, 170.e1–170.e19. doi:10.1016/j. ode.2007.06.005 Spiridonov VA, Türkay M (2001) Deep sea swimming crabs of the Charybdis miles species group in the western Indian Ocean (Crustacea: Decapoda: Portunidae). Journal of Natural History 35, 439–469. doi:10.1080/002229301300009649 Spiridonov VA, Neretina T, Schepetov D (2014) Morphological characterization and molecular phylogeny of Portunoidea Rafinesque,1815 (Crustacea Brachyura): implications for understanding evolution of swimming capacity and revision of the family-level classification. Zoologischer Anzeiger 253, 404–429. doi:10.1016/j.jcz.2014.03.003 Spiridonov VA, Kamanli SA, Naruse T, Clark PF (2021) Libystes A. MilneEdwards, 1867 (Crustacea: Decapoda: Portunidae): reestablishment of L. nitidus A. Milne-Edwards, 1867, reinstatement of L. alphonsi Alcock, 1900 and a description of a new species from the Red Sea. Arthropoda Selecta 30, 267–284. Stephenson W (1972) An annotated check list and key to the Indo-WestPacific swimming crabs (Crustacea: Decapoda: Portunidae). Royal Society of New Zealand Bulletin 10, 1–64. Stephenson W, Campbell BM (1959) The Australian portunids (Crustacea: Portunidae). III. The genus Portunus. Australian Journal of Marine and Freshwater Research 10, 84–124, pls 1–5. Stephenson W, Campbell BM (1960) The Australian portunids (Crustacea: Portunidae) IV. Remaining genera. Australian Journal of Marine and Freshwater Research 11, 73–122, pls 1–6. Stephenson W, Hudson JJ (1957) The Australian portunids (Crustacea; Portunidae). I. The genus Thalamita. Australian Journal of Marine and Freshwater Research 8, 312–368, pls 1–10. Stephenson W, Rees M (1968a) The Endeavour and other Australian Museum collections of portunid crabs (Crustacea, Decapoda, Portunidae). Records of the Australian Museum 27, 285–198, pl. 43. Stephenson W, Rees M (1968b) A revision of the Charybdis miles “group” of species (Crustacea: Portunidae), with description of a new species from Queensland waters. Memoirs of the Queensland Museum 15, 91–109. Stephenson W, Hudson JJ, Campbell B (1957) The Australian portunids (Crustacea: Portunidae). II. The genus Charybdis. Marine and Freshwater Research 8, 491–508. doi:10.1071/MF9570491 Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159.

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Števčić Z (2011) Addition to the reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Part I. New taxa. Natura Croatica 20, 125–139. Stimpson W (1860) Notes on North American Crustacea, in the Museum of the Smithsonian Institution, No. II. Annals of the Lyceum of Natural History of New York 7, 176–246, pls 2, 5. Takeda M (1993) A new swimming crab of the genus Carupa from submarine caves in the Ryukyu Islands. Bulletin of the National Science Museum, Ser. Zoology 19, 145–150. Takeda M (2003) Atoportunus dolichopus, a new cavernicolous crab of the family Portunidae (Crustacea: Decapoda) from the Ryukyu Islands. Bulletin of the National Science Museum, Ser. Zoology 29, 141–146. Takeda M, Webber R (2006) Crabs from the Kermadec Islands of the South Pacific. National Science Museum Monographs 34, 191–237. Türkay M, Spiridonov V (2006) Deep water swimming crabs of the subgenus Charybdis (Goniohellenus) of the western Indian Ocean (Crustacea: Decapoda: Portunidae). Fauna of Arabia 22, 199–223. Ward M (1942) Notes on the Crustacea of the Desjardins Museum, Mauritius Institute, with descriptions of new genera and species. Mauritius Institute Bulletin 2, 49–113, pls 5, 6. Wee DPC, Ng PKL (1995) Swimming crabs of the genera Charybdis De Haan, 1833, and Thalamita Latreille, 1829 (Crustacea: Decapoda: Brachyura: Portunidae) from Peninsular Malaysia and Singapore. Raffles Bulletin of Zoology Supplement 1, 1–128. Williams AB (1974) The swimming crabs of the genus Callinectes (Decapoda: Portunidae). Fishery Bulletin 72, 685–798. Williams AB (1984) Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C. Yosuva M, Viswanathan C, Samuel VKD, Chamundeeswari K, Purvaja R, et  al. (2020) New distributional record of petal-eyed swimmer crab, Vojmirophthalmus minabensis (Sakai 1961) (Portunidae: Podophthalminae) from Indian Waters. Thalassas: An International Journal of Marine Sciences 36, 535–538. doi:10.1007/s41208-020-00226-8 Zupolini LL, Magalhães T, Pileggi LG, Mantelatto FL (2017) Taxonomic revision of the speckled crabs, genus Arenaeus Dana, 1851 (Brachyura: Portunidae) based on morphological and molecular data. Zootaxa 4273, 362–380. doi:10.11646/zootaxa.4273.3.3

Thiidae Dana, 1852 Figure 14.130, Plate 62l Thiids, with only the genus Thia, are small, fossorial crabs with an unusual cordiform carapace lined with a continuous fringe of setae around the entire margin. Thiidae have usually been associated with atelecylids, but recent studies place them among the portunoids, usually close to the polybiids (Ng et  al. 2008; Schubart and Reuschel 2009; Spiridonov et  al. 2014). The single known species, Thia scutellata (­Fabricius, 1793), commonly called the thumbnail crab, is relatively common in the North Sea on nearshore sandy and shelly habitats (Christiansen 1969). Diagnosis. Carapace cordiform, dorsally smooth, surface convex transversely, straight longitudinally; front entire, broadly curved, without spines or lobes; lateral margins with up to 3 minute spines; entire margin with continuous fringe of long golden setae. Antennules folding obliquely. Male pleonite 3 without transverse ridge.

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Fig. 14.130.  Thiidae, Thia scutellata (Fabricius, 1793).

Thia Leach, 1815 thumbnail crab Maximum cl. 22 mm (Pl. 62l). Intertidal–shelf (0–25 m). Temperate North Atlantic (including Mediterranean Sea), Tropical E Atlantic. 1 species. References Christiansen ME (1969) Crustacea Decapoda Brachyura. Marine Invertebrates of Scandinavia 2, 1–143. Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Schubart CD, Reuschel S (2009) A proposal for a new classification of Portunoidea and Cancroidea (Brachyura: Heterotremata) based on two independent molecular phylogenies. In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 533–549. CRC Press, Boca Raton. Spiridonov VA, Neretina T, Schepetov D (2014) Morphological characterization and molecular phylogeny of Portunoidea Rafinesque,1815 (Crustacea Brachyura): implications for understanding evolution of swimming capacity and revision of the family-level classification. Zoologischer Anzeiger 253, 404–429. doi:10.1016/j.jcz.2014.03.003

Pseudocarcinoidea Ng & Davie, 2020 The superfamily includes a single family and species, the Tasmanian Giant Crab, Pseudocarcinus gigas (Lamarck, 1818). Pseudocarcinus was formerly considered a menippid (Eriphioidea) but was removed to its own superfamily by Ng and Davie (2020) on the basis of eriphioid polyphyly demonstrated by molecular phylogenetics (Lai et al. 2014). Adult and larval pseudocarcinoids are morphologically similar to eriphioids, especially oziids and menippids, but are

distinguished by subtle differences in features of the anterior cephalothorax and thoracic sternum (Ng and Davie 2020), and more obvious adult features of the carapace and pereopods as used in the keys. Adult Pseudocarcinus can be distinguished from all eriphioids, not only by their massive size, but also by the closely set orbits and length of greatly enlarged major male chela, which distinctly exceeds, rather than not exceeding carapace width. Owing to allometric changes in carapace and cheliped proportions, however, juveniles of Pseudocarcinus will resemble oziids and menippids. In such instances, Pseudocarcinus can still be recognised by the spine row on the extensor margin of the walking leg meri, always absent in oziids and menippids. Diagnosis. Carapace transversely ovate, wider than long; frontal width of adults 0.15 carapace width or greater. Antennules folding obliquely. Antenna longer than cornea width; flagellum well developed. Eyes on anterior margin, only upper margin of orbit visible in dorsal view when eyes retracted. Cheliped palm ovate in cross-section; adult male major chela length distinctly exceeding carapace width. Pereopods 2–5 similar, as walking legs; dactylus slender, pointed; unarmed except for row of extensor meral spines. Mouthfield approximately quadrate. Maxilliped 3 merus subquadrate. Male pleonites 1–6 freely articulating; pleonite 3 width less than twice telson width, without transverse ridge; narrowly linguiform, with lateral margins of pleonites 3–6 gradually converging towards telson. Gonopod 1 straight. Gonopod 2 about as long as gonopod 1, distally looped.

Pseudocarcinidae Ng & Davie, 2020 Figure 14.131, Plate 63a Pseudocarcinus H. Milne Edwards, 1834 Tasmanian giant crab Diagnosis. Carapace smooth in adults, tuberculate in juveniles; frontal margin with 4 short teeth lobes; distance between antennae oneseventh carapace width; anterolateral margin about as long as posterolateral margin, with exorbital tooth plus 4 obscure lobes, each with 2 or 3 short apices, epibranchial tooth blunt; orbit open, supra- and suborbital angles not in contact. Antennal basal article very short, not in contact with front. Maximum cl. 290 mm (Pl. 63a). Shelf, slope (30–820 m). Temperate Australia. 1 species (Ng and Davie 2020: relationships, ecology, figures; Poore 2004: figures). The Tasmanian giant crab, Pseudocarcinus gigas (Lamarck, 1818), the only species of its genus is so familiar to most Australian carcinologists that neither it nor its genus have ever been diagnosed since Henri Milne Edwards described it on the basis of specimens collected at the end of the eighteenth century. The species occurs on rocky reefs across southern Australia and may be the heaviest brachyuran crab, reaching at least 17.6 kg but possibly more. This weight is exceeded only by estimates of fishers who have reported

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Fig. 14.131.  Pseudocarcinidae. Pseudocarcinus gigas (Lamarck, 1818), adult male, female (not to same scale, from McCoy 1890).

even larger animals falling back into the sea from crab pots! It is only males with huge chelipeds that reach this weight; the heaviest females are ~6 kg. Males can reach 40 cm across the carapace and the larger cheliped is longer than this. For comparison, the majoid spider crab, Macrocheira kaempferi, reaches more than 18 kg and its legs have a 3 m span. A commercial giant crab pot fishery commenced in 1992 after a 20-year period when crabs were taken as a by-catch with rock lobsters. Much has been written on fisheries biology (Levings et al. 2001; McGarvey et al. 2002). References Lai JCY, Thoma BP, Clark PF, Felder DL, Ng PKL (2014) Phylogeny of eriphioid crabs (Brachyura, Eriphioidea) inferred from molecular and morphological studies. Zoologica Scripta 43, 52–64. doi:10.1111/zsc.12030 Levings A, Mitchell BD, McGarvey R, Mathews J, Laurenson L, et al. (2001) Fisheries biology of the giant crab, Pseudocarcinus gigas. Fisheries Research and Development Corporation Final Report 93/220 & 97/132. McCoy F (1890) Prodromus of the Zoology of Victoria: or figures and descriptions of the living species of all classes of the Victorian indigenous animals. Vol. 2. Decade 18. Government Printer, Melbourne. McGarvey R, Levings A, Matthews JM (2002) Moulting growth of the Australian giant crab, Pseudocarcinus gigas. Marine and Freshwater Research 53, 869–881. doi:10.1071/MF00074 Ng PKL, Davie PJF (2020) A new family and superfamily for the southern giant crab of Australia, Pseudocarcinus gigas (Lamarck, 1818) (Decapoda: Brachyura). Journal of Crustacean Biology 40, 607–626. doi:10.1093/jcbiol/ruaa058

Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne.

Pseudozioidea Alcock, 1898 The composition of Pseudozioidea has been one of the most fluid within Brachyura. The 13 genera now included have been variously entangled in the flux between goneplacoid, pilumnoid and xanthoid families. They are now unified by having gonopod 2 much shorter than gonopod 1, the penis emerging from the pereopod 5 coxa anterior to the proximal portion of the condyle, and vulvae without an operculum positioned relatively close to each other (Naruse and Ng 2014; Ng 2010; Ng et al. 2008). Four families of Pseudozioidea are recognised (Naruse and Ng 2014) Diagnosis. Carapace usually ovate to rounded, or subhexagonal to subquadrate; wider than long, flat, regions moderately or not defined. Antennule folding obliquely. Buccal cavity approximately quadrate. Endostomial ridges present. Maxilliped 3 irregularly subquadrate. Male pleonites 1–6 and telson free. Gonopod 2 about half gonopod 1 length, or slightly less. Penis emerging from the pereopod 5 coxa anterior to the proximal portion of condyle.

Key to families of Pseudozioidea 1. –

Carapace oval-subcircular (Fig. 14.133a, b). Male thoracic sternite 3 wider than anterior width of sternite 4 (Fig. 14.133d, e)��������������������������������������������������������������������������������������������������������������������������������������������������� Pilumnoididae Carapace transversely ovate to subquadrate (Figs 14.132a, b, 14.135a, b, 14.134a–e). Male thoracic sternite 3 as wide as or narrower than anterior margin of sternite 4 (Figs 14.132e, 14.135e, f, 14.134f–i)��������������������������������������2

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Male thoracic sternite 3 width subequal to combined length of thoracic sternites 3 and 4 (Figs 14.132e, 14.135e, f)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Male thoracic sternite 3 width distinctly greater than combined length of thoracic sternites 3 and 4 (Fig. 14.134f–h)�����������������������������������������������������������������������������������������������������������������������������������������������Planopilumnidae Carapace narrowly subovate, 1.4 times as wide as long; with 2 sharp, anteriorly recurved spines near lateral extremities (Fig. 14.132a, b). Male thoracic sternites 1–2 forming approximately right-angled triangle, margins straight, width almost twice length (Fig. 14.132e). Antennae, when folded, not completely retracting into fossa (Fig. 14.132c)����������������������������������������������������������������������������������� Christmaplacidae Carapace transversely ovate, 1.6–1.7 times as wide as long; margins unarmed or with very shallow notches (Fig. 14.135a, b). Male thoracic sternites 1–2 forming broad triangle, margins concave, divergence angle obtuse although apex acute, width exceeding twice length (Fig. 14.135e, f). Antennae, when folded, completely retracting into fossa (Fig. 14.135c, d)�������������������������������������������������� Pseudoziidae

References Naruse T, Ng PKL (2014) A new family, genus and species of cavernicolous crab (Crustacea: Decapoda: Brachura: Pseudozioidea) from Christmas Island, Australia. Raffles Bulletin of Zoology Supplement 30, 263–273. Ng PKL (2010) On the Planopilumnidae Serène, 1984 (Crustacea: Brachyura: Pseudozioidea), with diagnoses of two new pilumnoid genera for species previously assigned to Planopilumnus Balss, 1933. Zootaxa 2392, 33–61. doi:10.11646/zootaxa.2392.1.2 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286.

Christmaplacidae Naruse & Ng, 2014 Figure 14.132 Christmaplacids have relatively longer and thinner legs than other pseudozioids, are more or less blind, and lack dorsal carapace ornamentation or dense setae. One species inhabits anchialine caves, the other is from coral rubble. Diagnosis. Carapace narrowly subovate, 1.4 times as long as wide; with 2 sharp, anteriorly recurved spines near lateral extremities. Male thoracic sternites 1, 2 forming rightangled triangle; sternite 4 wider than sternite 3, about as wide as combined length of sternites 3 and 4. Male pleon broad, pleonite 3 width about equal to combined length of pleonite 4–6 and telson. Antennular article 3 as long as article 2, not or partially retracting into fossa. Cheliped carpus with strong lamellar tooth on inner margin. Gonopod 1 slender, long, gently curved, tapering; lateral margins without spinules. Christmaplax Naruse & Ng, 2014 Diagnosis. Carapace front scarcely produced. Pereopod 4 longest, 2.3 times carapace width; dactylus about as long as propodus (Fig. 14.132a). Maximum cl. 8 mm. Marine anchialine caves (1–2 m). Western Indo-Pacific (Christmas I.). 1 species (Naruse and Ng 2014).

Harryplax Mendoza & Ng, 2017 Diagnosis. Carapace front produced as 2 lobes. Pereopod 4 longest, 1.6 times carapace width; dactylus less than two-thirds as long as propodus (Fig. 14.132b, h). Maximum cl. 8.3 mm. Subtidal (1–1.5 m). Coral rubble. Central Indo-Pacific (Guam). 1 species (Mendoza and Ng 2017). References Mendoza JC, Ng PKL (2017) Harryplax severus, a new genus and species of an unusual coral rubble-inhabiting crab from Guam (Crustacea, Brachyura, Christmaplacidae). ZooKeys 647, 23–35. doi:10.3897/ zookeys.647.11455 Naruse T, Ng PKL (2014) A new family, genus and species of cavernicolous crab (Crustacea: Decapoda: Brachura: Pseudozioidea) from Christmas Island, Australia. Raffles Bulletin of Zoology Supplement 30, 263–273.

Pilumnoididae Guinot & Macpherson, 1987 Figure 14.133 The two pilumnoidid genera are notable for their subcircular carapace, a male pleon that is more slender than in other families and the male thoracic sternite 3 being wider than the anterior margin of sternite 4. In general habitus and strong overall setation, pilumnoidids superfacially resemble pilumnids and acidopsids. Diagnosis. Carapace oval-subcircular, ~1.2 times as wide as long. Male thoracic sternites 1, 2 forming broad triangle with concave lateral margins and pointed tip, or forming right-angled triangle; sternite 4 anteriorly narrower than sternite 3, sternite 3 narrower than combined length of sternites 3 and 4, together as long as wide. Male pleon narrow, pleonite 3 width two-thirds length of pleonite 4 to telson. Antennular article 3 much shorter than article 2, completely retracting into fossa. Cheliped carpus with weak tooth on inner margin. Gonopod 1 slender, long, gently curved, tapering, or stout, slightly sinuous, tip bluntly pointed, simple; lateral margins lined with spinules.

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Fig. 14.132.  Christmaplacidae. a, Christmaplax mirabilis Naruse & Ng, 2014. b, carapace, Harryplax. Left antennule, antenna, orbit: c, Christmaplax; d, Harryplax. e, male thoracic sternites 1–4, Christmaplax. f, right male thoracic sternites 7, 8, coxa 5, penis (shaded), Christmaplax. g, male pleon, Christmaplax. h, pereopod 4, Harryplax. i, j, gonopods 1, 2, Christmaplax.

Fig. 14.133.  Pilumnoididae. a, Pilumnoides rubus Guinot & Macpherson, 1987. b, carapace, Setozius incertus Ng & Ahyong, 2013; c, left antennule, antenna, orbit, Setozius. Male thoracic sternites 1–4: d, Pilumnoides; e, Setozius. Male pleon: f, Pilumnoides; g, Setozius. h, i, gonopod 1, with detail of tip, Pilumnoides. j, gonopod 2, Pilumnoides.

Pilumnoides Lucas, 1844 Diagnosis. Carapace dorsal surface with setae, particularly around margin; anterolateral margin with irregular teeth (Fig.  14.133a). Male thoracic sternite 3–4 as wide as long (excluding episternites) (Fig. 14.133d). Maximum cl. 23 mm. Intertidal–slope (0–500 m). Temperate Northern and Tropical Atlantic, Tropical Eastern Pacific, Temperate South America, Temperate Southern Africa. 8 species (Guinot and Macpherson 1987: key to species).

Setozius Ng & Ahyong, 2013 Diagnosis. Carapace dorsal surface with stiff setae evenly distributed; anterolateral margin with 5 well spaced short spines

(Fig. 14.133b). Male thoracic sternite 3–4 wider than long (excluding episternites) (Fig. 14.133e). Maximum cl. 13 mm. Subtidal (probable). Central Indo-Pacific. 2 species (Naruse and Maenosono 2019; Ng and Ahyong 2013). References Guinot D, Macpherson E (1987) Révision du genre Pilumnoides Lucas, 1844, avec description de quatre espèces nouvelles et création de Pilumnoidinae subfam. nov. (Crustacea Decapoda Brachyura). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série 9, 211–247, pls 1, 2. Naruse T, Maenosono T (2019) Setozius ikeharai, a new species of Pilumnoididae (Crustacea: Decapoda: Brachyura: Pseudozioidea) from Okinawa Island, Ryukyu Islands, Japan. Biological Magazine Okinawa 57, 169–179.

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Ng PKL, Ahyong ST (2013) Discovery of a new genus and new species of Indo-West Pacific pilumnoidid crab from a semisubmersible oil platform (Crustacea: Brachyura: Pseudozioidea). Zootaxa 3682, 513–520. doi:10.11646/zootaxa.3682.4.2

Planopilumnidae Serène, 1984 Figure 14.134 Most planopilumnids have a subquadrate-trapezoid carapace, some more or less ornamented, but the enigmatic Flindersoplax, once thought to be a goneplacid, is transversely ovate. The five genera are distinguished from

other pseudozioids by the wide rectangular male thoracic sternite 4. Diagnosis. Carapace transversely ovate to subquadrate. Male thoracic sternites 1, 2 forming obtuse or right-angled triangle; sternite 4 wider than sternite 3, wider than combined length of sternites 3 and 4, transversely trapezoid. Male pleon broad, pleonite 3 width about equal to combined length of pleonites 4–6 and telson. Antennular article 3 much shorter than article 2, completely retracting into fossa. Cheliped carpus with weak tooth on inner margin. Gonopod 1 slender, long, gently curved, tapering, or stout, slightly sinuous, tip bluntly pointed, simple or flared; lateral margins lined with spinules.

Key to genera of Planopilumnidae 1. – 2. – 3. –

Carapace transversely ovate, ~1.6 times as wide as long; anterolateral margin with 4 low, blunt teeth (Fig. 14.134a). Gonopod 2 distal part very short (Fig. 14.134o)���������������������������������������������������������������������Flindersoplax Carapace subquadrate–trapezoid, 1.3–1.4 times as wide as long; anterolateral margin with 3 or 4 spines or teeth (Fig. 14.134b–e). Gonopod 2 distal part about one-quarter length (Fig. 14.134q)��������������������������������������������������2 Carapace anterolateral margin with 3 short well spaced spines (Fig. 14.134b). Pereopods 2–4 meri upper margins with spines (Fig. 14.134k)������������������������������������������������������������������������������������������������������������������������Haemicinus Carapace anterolateral margin with 3 or 4 broad teeth or lobes (Fig. 14.134c–e). Pereopods 2–4 meri upper margins smooth or spinulose (Fig. 14.134l, m)�����������������������������������������������������������������������������������������������������������������������3 Carapace flat, with few setae; anterolateral margin with 3 truncate teeth, 1 triangular tooth (Fig. 14.134d)��������������������������������������������������������������������������������������������������������������������������������������������������������Platychelonion Carapace convex, densely pubescent; anterolateral margin with 3 or 4 regular or uneven teeth or lobes (Fig. 14.134c, e)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������4

Fig. 14.134.  Planopilumnidae. Carapace: a, Flindersoplax; b, Haemocinus; c, Planopilumnus; d, Platychelonion; e, Rathbunaria. Male thoracic sternites 1–4: f, Flindersoplax; g, Haemocinus; h, Rathbunaria. i, female thoracic sternum (vulvae shaded), Planopilumnus. j, male pleon, Flindersoplax. Pereopod 4 merus: k, Haemocinus; l, Platychelonion; m, Rathbunaria. Gonopods 1, 2: n, o, Flindersoplax; p, q, Planopilumnus

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Carapace with distinct transverse gastric cristae, without posterolateral and posterior submarginal groove extending to sub-branchial area; anterolateral margin with exorbital plus 3 distinct angular or subquadrate teeth (Fig. 14.134c). Pereopods 2–4 without prominent longitudinal ridge���������������������� Planopilumnus Carapace without distinct transverse gastric cristae, with posterolateral and posterior submarginal groove extending to sub-branchial area; anterolateral margin with 3 or 4 irregular teeth, some flattened (Fig. 14.134e). Pereopods 2–4 meri eroded, with prominent longitudinal ridge (Fig. 14.134m)��������������� Rathbunaria Flindersoplax Davie, 1989

Diagnosis. Carapace transversely ovate, ~1.5 times as wide as long; dorsal surface with regions poorly defined, anteriorly granulate; anterolateral margin slightly convex, shorter than posterolateral margin, with 4 low blunt teeth. Pereopods 2–5 slender, meri ~4 times as long as wide. Gonopod 1 stout, tapering. Gonopod 2 about half length of gonopod 1, distal part very short. Maximum cl. 16 mm. Subtidal. Temperate Australia. 1 species (Davie 1989).

Haemocinus Ng, 2003 Diagnosis. Carapace rounded-subquadrate, ~1.3 times as wide as long; dorsal surface with distinct transverse gastric cristae, pubescent; anterolateral margin slightly convex, as long as lateral margin, with 3 short spines. Pereopods 2–4 meri upper margins spinose. Gonopod 1 slender, curved. Gonopod 2 half as long as gonopod 1, distal part one-quarter length. Maximum cl. 18 mm. Intertidal. Temperate Northern W Pacific, Central IndoPacific. 1 species (Ng 2003).

Planopilumnus Balss, 1933 Diagnosis. Carapace subquadrate, ~1.3 times as wide as long; dorsal surface with distinct transverse gastric cristae, pubescent; anterolateral margin weakly convex, with exorbital plus 3 distinct angular or subquadrate teeth. Pereopods 2–4 meri with low subdistal tooth. Gonopod 1 stout, with flared apex. Gonopod 2 half as long as gonopod 1, distal part one-quarter length. Maximum cl. 28 mm. Intertidal, subtidal. Western Indo-Pacific. 2 species (Ng 2010; Ng and Kazmi 2011).

Platychelonion Crosnier & Guinot, 1969 Diagnosis. Carapace subquadrate, ~1.4 times as wide as long; dorsal surface flat, weakly pubescent; anterolateral margin with 3 truncate teeth, 1 triangular tooth. Pereopods 2–5 meri margins with spinules. Gonopod 1 stout, tapering. Gonopod 2 half as long as gonopod 1, distal part one-quarter length. Maximum cl. 21 mm. Subtidal (10–20 m). Tropical E Atlantic. 1 species (Ng 2010: rediagnosis, compared with Planopilumnus).

Rathbunaria Ward, 1933 Diagnosis. Carapace subquadrate, ~1.3 times as wide as long; dorsal surface with regions poorly defined; with posterolateral and

posterior submarginal groove extending to sub-branchial area, pubescent; anterolateral margin with 3 or 4 irregular teeth, some flattened. Pereopods 2–4 meri eroded, with prominent longitudinal ridge. Gonopod 1 slender, curved. Gonopod 2 half as long as gonopod 1, distal part one-quarter length. Maximum cl. 13 mm. Intertidal, subtidal. Central Indo-Pacific. 1 species (Ng 2010: rediagnosis, compared with Planopilumnus). References Davie PJF (1989) A re-appraisal of Heteropanope Stimpson, and Pilumnopeus A. Milne Edwards (Crustacea: Decapoda: Pilumnidae) with descriptions of new species and genera. Memoirs of the Queensland Museum 27, 129–156. Ng PKL (2003) The Indo-Pacific Pilumnidae XVII. On the identity of Pilumnus elatus A. MIlne_Edwards, 1873 (Crustacea, Decapoda, Brachyura). Journal of Taiwan Museum 56, 15–28. Ng PKL (2010) On the Planopilumnidae Serène, 1984 (Crustacea: Brachyura: Pseudozioidea), with diagnoses of two new pilumnoid genera for species previously assigned to Planopilumnus Balss, 1933. Zootaxa 2392, 33–61. doi:10.11646/zootaxa.2392.1.2 Ng PKL, Kazmi QB (2011) Planopilumnus holthuisi, a new species of planopilumnid crab (Decapoda, Brachyura, Pseudozioidea) from Pakistan. In: Fransen C, De Grave S, Ng P (Eds) Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume. Crustaceana Monographs 14, 585–596.

Pseudoziidae Alcock, 1898 Figure 14.135 Pseudoziid crabs are transversely ovate, distinguished from other pseudozioids by the male thoracic sternites 1 and 2 forming a broad triangle with concave margins converging on a point and a broad sternite 4. Euryozius possesses a submarginal stridulating ridge along the anterolateral margin, Pseudozius does not. Diagnosis. Carapace transversely ovate, 1.6–1.7 times as wide as long. Male thoracic sternites 1, 2 forming broad triangle with concave lateral margins and pointed tip; sternite 4 wider than sternite 3, about as wide as combined length of sternites 3 and 4. Male pleon narrow, pleonite 3 width less than combined length of pleonites 4–6 and telson. Antennular article 3 much shorter than article 2, completely retracting into fossa. Cheliped carpus with weak tooth on inner margin. Gonopod 1 slender, long, gently curved, tapering, or stout, slightly sinuous, tip bluntly pointed, simple; lateral margins lined with spinules.

724

Marine Decapod Crustacea

Fig. 14.135.  Pseudoziidae. Carapace: a, Euryozius; b, Pseudozius. Left antennule, antenna, orbit, anterolateral margin: c, Euryozius; d, Pseudozius. Male thoracic sternites 1–4: e, Euryozius; f, Pseudozius. g, male pleon, Euryozius. h, i, gonopods 1, 2, Euryozius. Pereopod 5: j, Euryozius; k, Pseudozius.

Euryozius Miers, 1886

Retroplumoidea Gill, 1894

Diagnosis. Carapace transversely ovate, ~1.6 times as wide as long; anterolateral margin evenly convex, longer than posterolateral margin, with submarginal row of stridulating ridges (Fig. 14.135a, c). Pereopods 2–5 slender, meri ~5 times as long as wide (Fig.  14.135j). Gonopod 2 half as long as gonopod 1, distal part one-quarter length (Fig. 14.135h, j). Maximum cl. 30 mm. Intertidal–slope (0–300 m). Tropical Atlantic, Indo-West Pacific. 6 species (Davie 1993: rediagnosis; Manning and Holthuis 1981: descriptions; Ng and Liao 2002: species description, discussion).

Retroplumoid crabs have a flat almost rectangular carapace, long pereopods 2–4 and shorter thinner plumose fifth pereopods. All are from muddy environments in deep water from the mid-shelf to upper slope. The superfamily was introduced for a single family of two genera and comprehensively revised by de Saint Laurent (1989). Further records and illustrations were provided by McLay (2006) who also provided a key to all ten species. Diagnosis. Pereopod 5 reduced, with long marginal setae, appearing feather-like (Fig. 14.136d).

Pseudozius Dana, 1851 Diagnosis. Carapace transversely ovate, ~1.6 times as wide as long; anterolateral margin strongly convex, weakly demarcated from converging posterolateral margin, with scarcely defined lobes (Fig. 14.135b, d). Pereopods 2–5 stout, meri about twice as long as wide (Fig.  14.135k). Gonopod 2 about half length of gonopod 1, distal part very short. Maximum cl. 13 mm. Intertidal, subtidal (0–6 m). Indo-West Pacific. 3 species (Guinot 1968; Ng and Wang 1994). References Davie PJF (1993) Deepwater xanthid crabs from French Polynesia (Crustacea, Decapoda, Xanthoidea). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 14, 501–561. Guinot D (1968) Recherches préliminaires sur les groupements naturels chez les crustacés décapodes brachyoures. VI. Les Carpilinae. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 40, 320–334. Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Ng PKL, Liao LM (2002) On a new species of Euryozius Miers, 1886 (Crustacea: Decapoda: Brachyura: Pseudoziidae) from the Philippines, with notes on the taxonomy of the genus. Proceedings of the Biological Society of Washington 115, 585–593. Ng PKL, Wang C (1994) Notes on the enigmatic genus Pseudozius Dana, 1851 (Crustacea, Decapoda, Brachyura). Journal of Taiwan Museum 47, 83–99.

Retroplumidae Gill, 1894 Figure 14.136, Plate 63b Bathypluma de Saint Laurent, 1989 Diagnosis. Carapace lateral border with teeth. Eyestalks not reaching supraorbital teeth (Fig.  14.136a). Maximum cl. 22 mm (Pl. 63b). Slope (300–614 m). Western and Central Indo-Pacific. 3 species (McLay 2006: distribution, key to species; de Saint Laurent 1989: diagnosis, key).

Retropluma Gill, 1894 Diagnosis. Carapace lateral border with or without blunt lobes. Eyestalks reaching almost anterolateral angle (Fig.  14.136b, c). Maximum cl. 21 mm. Shelf, slope (70–440 m). Western and Central Indo-Pacific. 7 species (McLay 2006: figures, distribution, key to species; de Saint Laurent 1989: rediagnosis).

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Fig. 14.136.  Retroplumidae. Carapace: a, Bathypluma; b, Retropluma solomonensis McLay, 2006 (with pereopods); c, Retropluma planiforma Kensley, 1969. Pereopod 5: d, Retropluma.

References de Saint Laurent M (1989) La nouvelle superfamille des Retroplumoidea Gill, 1894 (Decapoda, Brachyura): systématique, affinités et évolution. In: Forest J (Ed.), Résultats des Campagnes Musorstom, vol. 5. Mémoires du Muséum National d’Histoire Naturelle, Paris 144, 103–179. McLay CL (2006) Retroplumidae (Crustacea, Decapoda) from the IndoMalayan archipelago (Indonesia, Philippme) and the Melanesian Arc islands (Solomon Islands, Fiji and New Caledonia), and paleogeographical comments. In: Richer de Forges B and Justine J-L (Eds) Tropical Deep-Sea Benthos vol. 24. Mémoires du Muséum National d’Histoire Naturelle, Paris 193, 375–391.

Trapezioidea Miers, 1886 Trapezioids are symbionts of tropical corals. With few exceptions this symbiosis is obligatory (Castro 1988, 2015). Most genera are confined to the Indo-West Pacific but a few representative species occur on tropical American reefs. The crabs are recognised by their large chelipeds relative to the

carapace. The carapace is relatively flat (Castro et al. 2004). Many are colourful, colour patterns being species-specific. The monophyly of the three families was debated by Ng et  al. (2008) and by Clark and Ng (2010) after comparing zoeal morphologies. The monophyly of each family was confirmed using molecular data by Lai et  al. (2009) but their results were equivocal regarding monophyly of Trapezioidea. Diagnosis. Carapace usually ovate to rounded, or subhexagonal to subquadrate; wider than long, flat, regions moderately or not defined. Antennule folding transversely. Buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular. Male pleonites 1–6 and telson free, or male pleonites 3–5 fused, immovable, even if some or all sutures visible. Gonopod 1 slender. Gonopod 2 usually half as long as gonopod 1, rarely one-third or as long as gonopod 1. Penis emerging from gonopore on pereopod 5 coxa to directly enter gonopod 1.

Key to families of Trapezioidea 1. – 2.



Male pleonites and telson freely articulating (Fig. 14.138d, e). Chelipeds conspicuously dissimilar in size (Fig. 14.138a–c)���������������������������������������������������������������������������������������������������������������������������������������������� Tetraliidae Male pleonites 3–5 fused, immovable, even if some or all sutures visible (Fig. 14.139j, k). Chelipeds usually equal or slightly unequal in size (Figs 14.137a–d, 14.139b, f, g, i)����������������������������������������������������������������������������������������2 Carapace rounded, dorsal surface covered with small granules and spines; anterolateral regions lined with numerous spines and granules (Fig. 14.137b). Cheliped propodus with prominent round or pointed tubercles along outer surface; merus short, without a row of teeth along anterior margin (Fig. 14.137a, b, e, f)������������������������������������������������������������������������������������������������������������������������������������������������� Domeciidae Carapace trapezoidal or transversely ovate, dorsal surface smooth or faintly rugose at best; anterolateral margin usually entire or with low teeth, sometimes with spines (Fig. 14.139a–i). Cheliped propodus smooth, without tubercles along outer surface; merus long, usually with a row of conspicuous teeth along one-third or more of anterior margin (Fig. 14.139f, g, i)�������������������������������������������������������������������������Trapeziidae

References Castro P (1988) Animal symbioses in coral reef communities: a review. Symbiosis 5, 161–184. Castro P (2015) Symbiotic Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes

translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 543–981. Brill, Leiden. Castro P, Ng PKL, Ahyong ST (2004) Phylogeny and systematics of the Trapeziidae Miers, 1886 (Crustacea: Brachyura), with the description of a new family. Zootaxa 643, 1–70. doi:10.11646/zootaxa.643.1.1

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Clark PF, Ng PKL (2010) Description of the first zoea of Domecia glabra Alcock, 1899 (Crustacea: Brachyura, Domeciidae) and implications for the systematics of the Trapezioidea. Proceedings of the Biological Society of Washington 123, 258–273. doi:10.2988/10-17.1 Lai JCY, Ahyong ST, Jeng M-S, Ng PKL (2009) Are coral-dwelling crabs monophyletic? A phylogeny of the Trapezioidea (Crustacea: Decapoda:Brachyura). Invertebrate Systematics 23, 402–408. doi:10.1071/​ IS09012 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286.

Domeciidae Ortmann, 1893 Figure 14.137, Plate 63c Domeciids are small crabs that are facultative or obligate symbionts with shallow-water scleractinian corals.

Species of Domecia may form pits in the coral skeleton where they become trapped and must resort to filterfeeding (Hoeksema and García-Hernández 2020; Patton 1967). The carapace is oval and covered with few granules but the chelae have conspicuous tubercles or short spines. The merus of the cheliped is barely visible under the anterior carapace. The taxonomy of the family was reviewed briefly by Castro et al. (2004). Their key is modified here. Diagnosis. Carapace rounded, smooth or with scattered small granules and spines. Chelipeds equal or unequal; propodus with prominent round or pointed tubercles on outer surface; merus short, without row of teeth along anterior margin. Male pleonites 3–5 fused, immovable, even if some or all sutures visible.

Key to genera of Domeciidae 1. – 2.

Carapace frontal and anterolateral margins with numerous acute teeth (Fig. 14.137c). Maxilliped 3 merus about twice as wide as long (Fig. 14.137h)����������������������������������������������������������������������������������������������������������������� Domecia Carapace frontal margin smooth or with denticles or tubercles, anterolateral margin with 3 teeth or small tubercles (Fig. 14.137a, b, d). Maxilliped 3 merus 1.3–1.8 times as wide as long��������������������������������������������������������������2 Chelipeds equal (Fig. 14.137b)������������������������������������������������������������������������������������������������������������������������������������������Maldivia

Fig. 14.137.  Domeciidae. a, Cherusius triunguiculatus (Borradaile, 1902); b, Maldivia symbiotica Borradaile, 1902. Carapace: c, Domecia glabra Alcock, 1899; d, Palmyria palmyrensis (Rathbun in Edmondson, 1923). Major chela: e, Cherusius; f, Palmyria. Maxilliped 3: g, Cherusius; h, Domecia; i, Palmyria.

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Chelipeds unequal (Fig. 14.137a)����������������������������������������������������������������������������������������������������������������������������������������������������3 Cheliped propodus with round tubercles (Fig. 14.137e). Maxilliped 3 merus 1.3 times as wide as long (Fig. 14.137g)����������������������������������������������������������������������������������������������������������������������������������������������������������������Cherusius Cheliped propodus with pointed tubercles (Fig. 14.137f). Maxilliped 3 merus 1.8 times as wide as long (Fig. 14.137i)������������������������������������������������������������������������������������������������������������������������������������������������������������������ Palmyria Cherusius Low & Ng, 2012

Diagnosis. Carapace frontal margin smooth, anterolateral margin with short, blunt teeth. Maxilliped 3 merus ~1.3 times as wide as long. Chelipeds unequal; cheliped propodus with round tubercles. Maximum cl. 3.7 mm. Intertidal–shelf (0–95 m; symbionts of many genera of ­shallow-water scleractinian corals). Indo-West Pacific, Tropical Eastern Pacific. 1 species (Castro et  al. 2004: rediagnosis; Garth 1939: figure; Serène 1984: figures). The only species has several synonyms. Cherusius is a replacement name for Jonesius Sankarankutty, 1962, the genus name used by Castro et al. (2004).

Domecia Eydoux & Souleyet, 1842 Diagnosis. Carapace frontal and anterolateral margins with numerous acute teeth. Maxilliped 3 merus about twice as wide as long. Chelipeds unequal; cheliped propodus spinose. Maximum cl. 4 mm (Pl. 63c). Intertidal–shelf (0–146 m; symbionts of shallow-water scleractinian corals, especially Pocillopora and Acropora, but also free-living on dead corals and rocks). Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific. 4 species (Castro et al. 2004: rediagnosis, key to species; Guinot 1964: redescriptions, figures, distribution).

Maldivia Borradaile, 1902 Diagnosis. Carapace frontal margin with microscopic teeth, anterolateral margin with short teeth. Maxilliped 3 merus as wide as long. Chelipeds equal; cheliped propodus with short teeth. Maximum cl. 1.5 mm. Subtidal (15 m; obligate symbiont of gorgonians). Western Indo-Pacific (Maldives only). 1 species (Borradaile 1902: description, figure; Castro et al. 2004: rediagnosis).

Palmyria Galil & Takeda, 1986 Diagnosis. Carapace frontal margin smooth, anterolateral margin with acute teeth. Maxilliped 3 merus ~1.8 times as wide as long. Chelipeds unequal; cheliped propodus with pointed tubercles. Maximum cl. 3.4 mm. Intertidal–shelf (0–30 m; obligate symbiont of shallow-water scleractinian and hydrozoan corals). Indo-West Pacific. 1 species (Castro et al. 2004: rediagnosis; Galil and Takeda 1986: diagnosis, figures; Serène 1984: figures). References Borradaile LA (1902) Marine crustaceans. III. The Xanthidae and some other crabs. In The Fauna and Geography of the Maldive and Laccadive

Archipelagoes; Being the Account of the Work carried on and of the Collections made by an Expedition during the years 1899 and 1900. (Ed. Gardiner JS) pp. 237–271. University Press, Cambridge. Castro P, Ng PKL, Ahyong ST (2004) Phylogeny and systematics of the Trapeziidae Miers, 1886 (Crustacea: Brachyura), with the description of a new family. Zootaxa 643, 1–70. doi:10.11646/zootaxa.643.1.1 Galil BS, Takeda M (1986) Resurrection of the genus Jonesius and establishment of a new genus: commensal crabs associated with corals from the Indo-Pacific Ocean. Bulletin of the National Science Museum, Ser. Zoology 12, 163–171. Garth JS (1939) New brachyuran crabs from the Galapagos Islands. Allan Hancock Pacific Expeditions 5, 9–29, pls 1–10. Guinot D (1964) Les trois espèces du genre Domecia (Decapoda: Brachyura): D. hispida Eydoux & Souleyet, D. glabra Alcock et D. acanthophora (Desbonne & Schram). Crustaceana 7, 267–283. doi:10.1163/​ 156854064X00470 Hoeksema BW, García-Hernández JE (2020) Host-related morphological variation of dwellings inhabited by the crab Domecia acanthophora in the corals Acropora palmata and Millepora complanata (Southern Caribbean). Diversity 12, 143. doi:10.3390/d12040143 Patton WK (1967) Studies on Domecia acanthophora, a commensal crab from Puerto Rico, with particular reference to modifications of the coral host and feeding habits. Biological Bulletin 132, 56–67. doi:10.2307/​ 1539878 Serène R (1984) Crustacés Décapodes Brachyoures de l’Océan Indien Occidental et de la Mer Rouge, Xanthoidea: Xanthidae et Trapeziidae. Avec un addendum par Crosnier, A.: Carpiliidae et Menippidae. Faune Tropicale 24, 1–400.

Tetraliidae Castro, Ng & Ahyong, 2004 Figure 14.138, Plate 63d Tetraliids are obligate symbionts of shallow-water scleractinian corals of the genus Acropora (Castro 1976, 2015; Castro et  al. 2004; Trautwein 2007). The crabs inhabit spaces between the coral branches feeding on mucus secreted by the coral (Castro 2015). Species can be hostspecific, distinguishing between species of Acropora (Sin 1999). All species are colourful, colour patterns being a useful adjunct to morphology distinguish species (Trautwein 2007). Diagnosis. Carapace widest across anterior; posterolateral margins strongly converging, posterior margin strongly narrowed, distance between coxae of pereopods 5 less than one-fifth carapace width. Carapace rounded, smooth or with scattered small granules and spines. Chelipeds equal or conspicuously unequal; propodus with prominent round or pointed tubercles on outer surface smooth or finely granulate; merus short, without row of small teeth along anterior margin. Male pleonites 1–6 and telson free.

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Marine Decapod Crustacea

Fig. 14.138.  Tetraliidae. Carapace, chelipeds: a, Tetralia. Minor, major chelipeds: b, c, Tetraloides. Male thoracic sternites 3–7, pleonites 3–6, telson: d, Tetralia; e, Tetraloides.

Tetralia Dana, 1851 Diagnosis. Larger cheliped propodus with proximal seta-filled depression on upper face (Fig. 14.138a). Thoracic sternites 4 and 5 with median longitudinal groove (Fig.  14.138d). Maximum cl. 9 mm (Pl. 63d). Intertidal–shelf (0–66 m). Indo-West Pacific. 10 species (Castro et al. 2004: rediagnosis, key to 6 species; Trautwein 2007: description of 4 species).

Tetraloides Galil, 1986 Diagnosis. Larger cheliped propodus without seta-filled depression (Fig.  14.138c). Thoracic sternites 4 and 5 without median longitudinal groove (Fig. 14.138e). Maximum cl. 9 mm. Intertidal–shelf (0–94 m). Indo-West Pacific. 2 species (Castro et al. 2004: rediagnosis, key to species; Galil 1986: diagnosis). References Castro P (1976) Brachyuran crabs symbiotic with scleractinian corals: a review of their biology. Micronesica 12, 99–110. Castro P (2015) Symbiotic Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 543–981. Brill, Leiden. Castro P, Ng PKL, Ahyong ST (2004) Phylogeny and systematics of the Trapeziidae Miers, 1886 (Crustacea: Brachyura), with the description of a new family. Zootaxa 643, 1–70. doi:10.11646/zootaxa.643.1.1 Galil BS (1986) Tetraloides - a new genus of coral-inhabiting crabs. Crustaceana 50, 68–77. doi:10.1163/156854085X00080 Sin T (1999) Distribution and host specialization in Tetralia crabs (Crustacea: Brachyura) symbiotic with corals in the Great Barrier Reef, Australia. Bulletin of Marine Science 65, 839–850. Trautwein SE (2007) Four new species of coral crabs belonging to the genus Tetralia Dana, 1851 (Crustacea, Decapoda, Brachyura, Tetraliidae). Zootaxa 1450, 1–20. doi:10.11646/zootaxa.1450.1.1

Trapeziidae Miers, 1886 Figure 14.139, Plate 63e–i Trapeziids are small obligate symbiotic crabs associated with a range of hard and soft corals. All are smooth or

slightly rugose with long chelipeds. The crabs usually occur as heterosexual pairs, feeding on the mucus generated by the coral and providing some protection for the host against predators such as crown-of-thorns starfish (Castro 1988, 2015; Huber and Coles 1986). The genera and species can be separated on morphology but colour pattern is a useful adjunct character, particularly in the largest genus, Trapezia (Castro 1996). The taxonomy of the trapeziids has been extensively studied primarily by Peter Castro, Bella Galil and colleagues (e.g. Castro 1996, 1997, 1998, 1999; Castro et  al. 2004; Galil 1986, 1997; Galil and Clark 1990). Števčić (2005) reworked the higher taxonomy and treated Trapeziidae (as understood here) as a subfamily that he divided into four tribes without specifying included ­genera. The tribes were discussed by Ng et  al. (2008) who recognised three subfamilies based on carapace shape, cheliped heterochely and setation of the dactyli of pereopods 2–4: Calocarcininae for Calocarcinus, ­Philippicarcinus and Sphenomerides; Quadrellinae for Quadrella and Hexagonalia; and Trapeziinae for Trapezia. Calocarcinines are associated with deep-sea soft and precious corals and sponges, Quadrellinae with antipatharians, gorgonians or soft corals in moderately deep water, and Trapezia is associated with stony corals. A molecular phylogeny of nine species in four genera does not contradict this subfamily arrangement (Lai et  al. 2009). One genus erected later, Hexagonaloides, was provisionally placed in Quadrellinae on the similar shape of the carapace and elongate pereopods by Komai et al. (2010) but it lacks the transverse rows of setae said to characterise this subfamily. Ectaesthesius was likened by Rathbun (1930) to Trapezia but has been placed hitherto in Xanthidae. It is currently poorly known but resembles members of Trapeziidae and perhaps Tanaochelidae. We tentatively include Ectaesthesius in Trapeziidae; the male pleonites 3–5 are likely fused but with sutures visible as in Philippicarcinus.

14 – Brachyura – crabs

Diagnosis. Carapace subtrapezoid to transversely ovate, smooth or at most faintly rugose. Chelipeds equal or unequal; propodus smooth, without tubercles along outer surface;

729

merus long, anterior margin usually with row of conspicuous teeth along one-third or more of length. Male pleonites 3–5 fused, immovable, even if some or all sutures visible.

Key to genera of Trapeziidae 1. – 2. – 3. – 4. –

Pereopods 2–5 dactyli with hollowed, hoof-like tip, transverse rows of spiniform setae on distal flexor margin (Fig. 14.139p)���������������������������������������������������������������������������������������������������������������������������������������������������� Trapezia Pereopods 2–5 dactyli with acute tip, without transverse rows of spiniform setae on flexor margin (Fig. 14.139l–o)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Carapace frontal margin with deep median notch. Pereopods 2–5 dactyli with facial transverse rows of setae (Fig. 14.139l, o)�������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Carapace frontal margin straight or obscurely bilobed. Pereopods 2–5 dactyli without facial transverse rows of thick setae (Fig. 14.139m, n) or setae not in rows�����������������������������������������������������������������������������������������������������4 Carapace anterolateral margin with 1 triangular epibranchial tooth, rarely second small intermediate tooth; front cut into 4 triangular lobes (Fig. 14.139f)��������������������������������������������������������������������������������������������Quadrella Carapace anterolateral margin with 2 acute anteriorly-directed teeth; front bilobed (Fig. 14.139c)������������� Hexagonalia Carapace anterolateral margin with 2 similar triangular teeth in anterior half of carapace (Fig. 14.139b)���������������������������������������������������������������������������������������������������������������������������������������������������������� Ectaesthesius Carapace anterolateral margin with 1 or 2 blunt or small anterolateral teeth, second if present posterior to midlength (Fig. 14.139a, d, e, g)������������������������������������������������������������������������������������������������������������������������������������������������5

Fig. 14.139.  Trapeziidae. Carapace, chelipeds: a, Calocarcinus; b, Ectaesthesius; c, Hexagonalia; d, Hexagonaloides; e, Philippicarcinus; f, Quadrella; g, Sphenomerides; h, i, Trapezia. Male pleon, telson: j, Hexagonaloides; k, Quadrella. Pereopod 5, dactylus: l, Hexagonalia; m, Hexagonaloides; n, Philippicarcinus; o, Quadrella; p, Trapezia.

730

5. – 6. – 7. –

Marine Decapod Crustacea

Carapace hexagonal, with 1 epibranchial tooth (Fig. 14.139d)�����������������������������������������������������������������������Hexagonaloides Carapace transversely octagonal (Fig. 14.139a) or ovate (Fig. 14.139e, g)�������������������������������������������������������������������������������6 Carapace octagonal, anterolateral margin with 2 teeth, anterior and posterior to subparallel sides of middle section of carapace (Fig. 14.139a)�������������������������������������������������������������������������������������������������������������������������Calocarcinus Carapace transversely ovate, anterolateral margin each with 1 or 2 teeth (Fig. 14.139e, g)�������������������������������������������������7 Carapace frontal margin straight. Cheliped merus anterior margin smooth. Carapace widest anterior to midpoint (Fig. 14.139e)������������������������������������������������������������������������������������������������������������������������������������ Philippicarcinus Carapace frontal margin gently biconvex. Cheliped merus anterior margin spinose. Carapace widest posterior to midpoint (Fig. 14.139g)���������������������������������������������������������������������������������������������������������������� Sphenomerides Calocarcinus Calman, 1909

Diagnosis. Carapace octagonal; frontal margin straight; anterolateral margin with 2 teeth, anterior and posterior to subparallel sides of middle section of carapace. Chelipeds dissimilar in size; merus anterior margin without teeth. Pereopods 2–5 dactyli with acute tip, without facial transverse rows of thick setae. Maximum cl. 8 mm (Pl. 63e). Shelf, slope (183–1098 m; symbionts of deep-water corals, Corallium, azooxanthellate or ahermatypic scleractinian corals, and probably antipatharian black corals and alcyonacean soft corals). Western and Central Indo-Pacific, Temperate Australasia (Kermadec Is). 4 species (Castro et al. 2004: rediagnosis, key to 4 species; Galil and Clark 1990: 4th species; Takeda and Galil 1980: figures of 3 species).

Ectaesthesius Rathbun, 1898 Diagnosis. Carapace ovate; frontal margin obscurely sinuous; anterolateral margin with 2 similar triangular teeth in anterior half of carapace. Chelipeds dissimilar in size. Cheliped merus anterior margin without teeth. Pereopods 2–5 dactyli with acute tip, with fine setae not in rows. Maximum cl. 7 mm. Shelf (37–110 mm, possibly associated with soft corals). Tropical Eastern Pacific (Galápagos). 1 species (Garth 1946: redescription, figures; Rathbun 1930: rediagnosis, description of only species).

Hexagonalia Galil, 1986 Diagnosis. Carapace hexagonal; frontal margin with conspicuous V-shaped median notch between 2 broad lobes; anterolateral margin with 2 acute anteriorly-directed teeth. Chelipeds dissimilar in size; merus anterior margin with teeth. Pereopods 2–5 dactyli with acute tip, with facial transverse rows of thick setae. Maximum cl. 6 mm. Intertidal, subtidal (0–5 m; symbionts on stylasterid hydrozoan corals and possibly gorgonians). Western and Central IndoPacific. 3 species (Castro et al. 2004: rediagnosis, key to species; Galil 1986: diagnosis; 1997: figures).

Hexagonaloides Komai, Higashiji & Castro, 2010 Diagnosis. Carapace hexagonal; frontal margin straight; anterolateral margin with 1 triangular epibranchial tooth at about midlength. Chelipeds dissimilar in size; merus anterior margin

without teeth. Pereopods 2–5 dactyli with acute tip, without facial transverse rows of thick setae. Maximum cl. 3.7 mm. Slope (240–247 m; unidentified anthozoan host). Central Indo-Pacific (Ryuku Is, Japan; New Caledonia). 1 species (Komai et al. 2010).

Philippicarcinus Garth & Kim, 1983 Diagnosis. Carapace transversely ovate; frontal margin straight; anterolateral margin with 1 or 2 blunt teeth, second set anterior to midpoint. Chelipeds dissimilar in size. Cheliped merus anterior margin without teeth. Pereopods 2–5 dactyli with acute tip, without facial transverse rows of thick setae. Maximum cl. 11.4 mm. Slope (333–510 m; host unknown). Central Indo-Pacific (Philippines). 2 species (Castro et al. 2004: rediagnosis, key to species; Garth and Kim 1983).

Quadrella Dana, 1851 Diagnosis. Carapace hexagonal; frontal margin with conspicuous V-shaped median notch between 2 pairs of teeth; anterolateral margin with 1 triangular epibranchial tooth at about midlength. Chelipeds dissimilar in size; merus anterior margin with teeth or tubercles. Pereopods 2–5 dactyli with acute tip, with facial transverse rows of thick setae. Maximum cl. 12 mm (Pl. 63f, g). Subtidal–slope (3–908 m; symbionts on antipatharian black corals, gorgonians, alcyonacean soft corals or azooxanthellate ahermatypic scleractinian corals). Indo-West Pacific, Tropical Eastern Pacific. 6 species (Castro et al. 2004: rediagnosis, key to 6 species; Galil 1986: redescriptions of 8 species, some since synonymised).

Sphenomerides Rathbun, 1897 Diagnosis. Carapace transversely ovate; frontal margin biconvex; anterolateral margin with 2 small teeth, second at about midlength. Chelipeds dissimilar in size; merus anterior margin with teeth. Pereopods 2–5 dactyli with acute tip, without facial transverse rows of thick setae. Maximum cl. 7.7 mm. Shelf, slope (80–660 m; symbiont of sponges). Western and Central Indo-Pacific. 1 species (Castro 2013: photos, host; Castro et al. 2004: rediagnosis).

14 – Brachyura – crabs

Trapezia Latreille, 1828 Diagnosis. Carapace rounded-trapezoidal; frontal margin straight or with short, round or triangular lobes; anterolateral margin with or without 1 triangular epibranchial tooth at about midlength. Chelipeds slightly dissimilar in size; merus anterior margin with teeth or tubercles. Pereopods 2–5 dactyli with hollowed, hoof-like tip, with transverse rows of spiniform setae on distal flexor margin. Maximum cl. 20 mm (Pl. 63h, i). Intertidal–shelf (0–90 m; symbionts of shallow-water scleractinian corals, usually pocilloporids). Indo-West Pacific, Tropical Eastern Pacific. 22 species (Castro 1996: figures of 4 eastern Pacific species; Castro et al. 2004: rediagnosis, key to 23 species, some refigured). References Castro P (1988) Animal symbioses in coral reef communities: a review. Symbiosis 5, 161–184. Castro P (1996) Eastern Pacific species of Trapezia (Crustacea, Brachyura: Trapeziidae), sibling species symbiotic with reef corals. Bulletin of Marine Science 58, 531–554. Castro P (1997) Trapeziid crabs (Brachyura: Xanthoidea: Trapeziidae) of New Caledonia, eastern Australia, and the Coral Sea. In Les fonds meubles des lagons de Nouvelle-Calédonie (sédimentologie, benthos). (Ed. Richer de Forges B) pp. 59–107. ORSTOM Editions, Paris. Castro P (1998) Systematic status and geographic distribution of Trapezia formosa Smith, 1869 (Crustacea, Brachyura, Trapeziidae), a symbiont of reef corals. Zoosystema 20, 177–181. Castro P (1999) Trapeziid crabs (Crustacea, Brachyura, Trapeziidae) of the Indian Ocean and the Red Sea. Zoosystema 21, 93–120. Castro P (2013) Brachyuran crabs (Crustacea, Brachyura: Crossotonotidae, Ethusidae, Euryplacidae, Goneplacidae, Latreilliidae, Palicidae, Tetraliidae, Trapeziidae) of the MAINBAZA, MIRIKI, and ATIMO VATAE expeditions to the Mozambique Channel and Madagascar. In: Ahyong ST, Chan T-Y, Corbari L and Ng PKL (Eds) Tropical Deep-Sea Benthos vol. 27. Mémoires du Muséum National d’Histoire Naturelle, Paris 2041, 437–466. Castro P (2015) Symbiotic Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 543–981. Brill, Leiden. Castro P, Ng PKL, Ahyong ST (2004) Phylogeny and systematics of the Trapeziidae Miers, 1886 (Crustacea: Brachyura), with the description of a new family. Zootaxa 643, 1–70. doi:10.11646/zootaxa.643.1.1 Galil B (1986) Quadrella (Brachyura: Xanthoidea: Trapeziidae): review and revision. Journal of Crustacean Biology 6, 275–293. doi:10.2307/​ 1547987 Galil B (1997) A new Hexagonalia (Brachyura, Xanthoidea, Trapeziidae), obligate commensal on Stylasteridae. Zoosystema 19, 449–452. Galil B, Clark PF (1990) Crustacea Decapoda: notes on trapeziid crabs from New Caledonia including descriptions of two new species. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 6. Mémoires du Muséum National d’Histoire Naturelle, Paris 145, 369–388. Garth JS (1946) Littoral brachyuran fauna of the Galapagos Archipelago. Allan Hancock Pacific Expeditions 5, 341–522, pls 49–87.

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Garth JS, Kim HS (1983) Crabs of the family Xanthidae (Crustacea: Brachyura) from the Philippine Islands and adjacent waters based largely on collections of the U.S. Fish Commission steamer Albatross in 1908–1909. Journal of Natural History 17, 663–729. doi:10.1080/​ 00222938300770561 Huber ME, Coles SL (1986) Resource utilization and competition among the five Hawaiian species of Trapezia (Crustacea, Brachyura). Marine Ecology Progress Series 30, 21–31. doi:10.3354/meps030021 Komai T, Higashiji T, Castro P (2010) A new genus and new species of deepwater trapeziid crab (Crustacea: Decapoda: Brachyura: Trapezioidea) from the Ryukyu Islands, Japan. Zootaxa 2555, 62–68. doi:10.11646/​ zootaxa.2555.1.4 Lai JCY, Ahyong ST, Jeng M-S, Ng PKL (2009) Are coral-dwelling crabs monophyletic? A phylogeny of the Trapezioidea (Crustacea:​ Decapoda:​Brachyura). Invertebrate Systematics 23, 402–408. doi:10.1071/​IS09012 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Takeda M, Galil BS (1980) A new species of Calocarcinus (Crustacea, Brachyura) from the Submarine Bank off the Izu Islands, Central Japan. Bulletin of the National Science Museum, Ser. Zoology 6, 191–194.

Trichopeltarioidea Tavares & Cleva, 2010 Trichopeltaroids are a group of more or less elongate, ornate crabs usually inhabiting deep waters from the lower shelf to mid-slope depths. Males have one cheliped considerably larger than the other. All have a projection of the lower orbital floor between the antennal peduncle and the suborbital tooth (termed an ‘suborbital intercalated piece’, Fig. 14.140g–h). This combination of characters differentiates trichopeltarioids from similar-looking corystid and atelecyclid crabs, with which they have been previously classified. There is only one family, of five genera, reviewed in detail by Tavares and Cleva (2010) on whose work this key and these diagnoses rely. Diagnosis. Carapace pentagonal, subcircular, or pyriform; front trilobed; orbits present; suborbital intercalated piece present. Antennule folding longitudinally or almost so. Buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate. Male chelipeds strongly unequal. Male pleonites 1–6 and telson free; male telson semicircular–subtriangular.

Trichopeltariidae Tavares & Cleva, 2010 Figure 14.140, Plate 63j, k

732

Marine Decapod Crustacea

Key to genera of Trichopeltariidae 1. – 2.



3. – 4. –

Carapace pyriform (Fig. 14.140b, f). Orbital cavity shallow (Fig. 14.140h, i). Antennal peduncles and flagella upper and lower faces with rows of opposing long setae, forming median antennal channel; flagella elongate or short������������������������������������������������������������������������������������������������������������������������������������������������������������2 Carapace subcircular or pentagonal (Fig. 14.140a, d–f). Orbital cavity deeply excavate (Fig. 14.140g, j, k). Antennae with or without incipient median setal channel; flagella much less than half carapace length�������������������3 Carapace dorsal surface densely covered with strong, acute or round tubercles. Second supraorbital tooth strong, triangular, acute. Hepatic tooth prominent. First epibranchial tooth strong or reduced, second epibranchial tooth long. Antennal peduncle articles 2+3 abutting but not fused with suborbital tooth tooth. Antennal peduncle article 5 shorter than 2+3 and 4; flagellum elongate (about half carapace length) (Fig. 14.140b, h)����������������������������������������������������������������������������������������������������������������������������������������� Podocatactes Carapace dorsal surface finely granulate. Second supraorbital tooth bump-like. Hepatic tooth bump-like. First epibranchial tooth absent; second epibranchial tooth a strong, laterally directed spine. Antennal peduncle articles 2, 3 separated by gap from suborbital tooth tooth. Antennal peduncle article 5 distinctly longer than 4; flagellum short (Fig. 14.140c, i)����������������������������������������������������������������������������� Pteropeltarion Antennal peduncle articles 2+3 separated by deep, narrow notch from suborbital tooth; peduncles and flagella upper and lower faces fringed by row of long setae (Fig. 14.140g). Male pleonites 3–5 fused��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������Peltarion Antennal peduncle articles 2+3 separated by deep, wide gap from suborbital tooth; peduncles and flagella surrounded by setae, with opposing setae of flagella forming incipient antennal channel (Fig. 14.140j, k). Male pleonites and telson free���������������������������������������������������������������������������������������������������������������������������������������������������4 Carapace pentagonal or subcircular. Hepatic, first and second epibranchial teeth prominent, usually armed with barbs. Pereopods 2–5 elongate, meri and propodi more than 3 times as long as wide (Fig. 14.140e, f)��������������������������������������������������������������������������������������������������������������������������������������������������� Trichopeltarion Carapace subcircular. Hepatic tooth a gently swollen tubercle; first and second epibranchial teeth minute, round, smooth similar tubercles. Pereopods 2–5 short, meri and propodi much less than 3 times as long as wide (Fig. 14.140d)�����������������������������������������������������������������������������������������������������������������������������Sphaeropeltarion

Fig. 14.140.  Trichopeltariidae. Carapace, habitus: a, Peltarion spinulosum (White, 1843); b, Podocatactes foresti (Guinot, 1989); c, Pteropeltarion novaezelandiae Dell, 1972; d, Sphaeropeltarion edentatum Tavares & Cleva, 2010; e, Trichopeltarion dextrum (Rathbun, 1898). f, Trichopeltarion alcocki Doflein in Chun, 1903. Antennules, antennae, ventral orbit (suborbital intercalated piece shaded): g, Peltarion; h, Podocatactes; i, Pteropeltarion; j, Sphaeropeltarion; k, Trichopeltarion.

14 – Brachyura – crabs

Peltarion Hombron & Jacquinot, 1846 Diagnosis. Carapace subcircular; dorsal surface finely granulate. Orbital cavity deeply excavate; second supraorbital tooth triangular. Hepatic tooth lobe-like; first and second epibranchial teeth lobe-like, armed with barbs. Antennal peduncle articles 2+3 separated by deep, deep, narrow notch from suborbital tooth. Antennal peduncles and flagella upper and lower faces fringed by row of long setae; antennal peduncle article 5 shorter than 2+3 and 4; flagella much less than half carapace length. Male pleonites 3–5 fused. Maximum cl. 57 mm. Intertidal–slope (0–305 m). Temperate South America. 1 species (Rathbun 1930: figure; Tavares and Cleva 2010: rediagnosis).

Podocatactes Ortmann, 1893 Diagnosis. Carapace pear-shaped; dorsal surface densely covered with strong, acute or round tubercles. Orbital cavity shallow; second supraorbital tooth triangular. Hepatic tooth prominent; first epibranchial tooth strong or reduced, second epibranchial tooth long. Antennal peduncle articles 2+3 abutting but not fused to suborbital tooth tooth. Antennal peduncles and flagella upper and lower face with rows of opposing long setae, forming median antennal channel; antennal peduncle article 5 shorter than 2+3 and 4; flagella elongate (about half carapace length). Male pleonites and telson free. Maximum cl. 29 mm (Pl. 63j). Shelf, slope (50–550 m). Temperate Northern W Pacific, Central Indo-Pacific. 3 species (Tavares and Cleva 2010: rediagnosis, key to species).

Pteropeltarion Dell, 1972 Diagnosis. Carapace pear-shaped; dorsal surface finely granulate. Orbital cavity shallow; second supraorbital tooth bump-like. Hepatic tooth bump-like; first epibranchial tooth absent; second epibranchial tooth strong, spine-like. Antennal peduncle articles 2+3 separated by gap from suborbital tooth tooth. Antennal peduncles and flagella upper and lower face with rows of opposing long setae, forming median antennal channel; antennal peduncle article 5 distinctly longer than 4; flagella short. Male pleonites and telson free. Maximum cl. 17 mm. Slope (499–903 m). Temperate Australasia (New Zealand only). 1 species (Dell 1972; Tavares and Cleva 2010: rediagnosis).

Sphaeropeltarion Tavares & Cleva, 2010 Diagnosis. Carapace subcircular; dorsal surface covered with thin coat of velvet and sparse short setae. Orbital cavity deeply excavate; second supraorbital tooth triangular. Hepatic tooth a gently swollen tubercle; first and second epibranchial teeth minute, round, smooth similar tubercles. Antennal peduncle articles 2+3 separated by deep, wide gap from suborbital tooth. Antennal peduncles and flagella surrounded by setae, with opposing setae of flagella forming incipient antennal channel; antennal peduncle article 5 shorter than 2+3 and 4; flagella much less than half carapace length. Pereopods 2–5 much less than twice as long as wide. Male pleonites and telson free. Maximum cl. 44 mm.

733

Slope (600–608 m). Central Indo-Pacific (Wallis and Futuna Is). 1 species (Tavares and Cleva 2010).

Trichopeltarion A. Milne-Edwards, 1880 Diagnosis. Carapace pentagonal, or subcircular; dorsal surface with granules or tubercles. Orbital cavity deeply excavate; second supraorbital tooth triangular. Hepatic tooth prominent; first and second epibranchial teeth prominent, usually armed with barbs. Antennal peduncle articles 2+3 separated by deep, wide gap from suborbital tooth. Antennal peduncles and flagella surrounded by setae, with opposing setae of flagella forming incipient antennal channel; antennal peduncle article 5 shorter than 2+3 and 4; flagella much less than half carapace length. Pereopods 2–5 elongate, meri and propodi more than 3 times as long as wide. Male pleonites and telson free. Maximum cl. 94 mm (Pl. 63k). Subtidal–slope (15–1700 m). Tropical Atlantic, Temperate Northern W Pacific, Western and Central Indo-Pacific, Tropical Eastern Pacific, Temperate South America, Temperate Southern Africa, Temperate Australasia. 23 species (Tavares and Cleva 2010: rediagnosis, 14 species illustrated). References Dell RK (1972) A new genus and species of atelecyclid crab from New Zealand. Journal of the Royal Society of New Zealand 2, 55–59. doi:10.1 080/03036758.1972.10423304 Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Tavares M, Cleva R (2010) Trichopeltariidae (Crustacea, Decapoda, Brachyura), a new family and superfamily of eubrachyuran crabs with description of one new genus and five new species. Papéis Avulsos de Zoologia 50, 97–157. doi:10.1590/S0031-10492010000900001

Xanthoidea MacLeay, 1838 Figure 14.141 For many decades, the xanthids, and later, the xanthoids, comprised a heterogeneous group of heterotrematous crabs revolving around a common typical body plan – stocky, transversely rounded-subhexagonal carapace with several anterolateral teeth, eyes set well inwards of the lateral extremities, and converging posterior margins; simple, stout walking legs and robust chelipeds with dark finger tips. The group has always been difficult to taxonomically diagnose and likewise, has always been complicated taxonomically, especially in the Indo-West Pacific. Progressive and ongoing revision of the xanthoids, substantially commenced by Guinot (1976), has clarified familial relationships, resolved many taxonomic ambiguities and removed multiple groups formerly considered as xanthids to other superfamilies, such as Aethroidea, Carpilioidea, Dairoidea, Eriphiioidea and Pilumnoidea (e.g. Serène 1984 (e.g. Davie et al. 2015; Karasawa and Schweitzer 2006; Lai

734

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et al. 2011; Ng et al. 2008; Serène 1984; Števčić 2005; Thoma et  al. 2014). Despite the removal of many xanthid-like groups from the orbit of Xanthoidea, the superfamily and its constituent families remain difficult to characterise. At present the xanthoids are united by similar, ‘uncomplicated’ organisation of the male gonopores (compared to that seen in goneplacoids), typical fusion of male pleonites 3–5, long gonopods 1 and short, straight gonopods 2. The xanthoid classification used here substantially follows that of Ng et  al. (2008) and Davie et  al. (2015). Currently, Xanthoidea are a highly diverse group of some 130 genera and 650 species of crabs, currently arrayed in four families distinguished primarily by the proportions of the thoracic sternites 3–4, exposure of sternite 8 in males and features of gonopod 1. Revision of the xanthoids is active and ongoing and the monophyly of all currently accepted families is questionable (Lai et al. 2011; Lai et al. 2014; Thoma et al. 2014);

further division of existing families is likely. Separation of genera and species, especially in Xanthidae, relies in part on the pattern of the elevated regions on the dorsal carapace. The alphanumeric labelling scheme devised by Dana (1852) is widely accepted and used here (Fig. 14.141a). Separation of families partly relies on the proportions of the anterior part of the male thoracic sternum (Fig. 14.141b). Diagnosis. Carapace hexagonal to oval, sometimes subcircular or subquadrate, usually wider than long, regions indicated. Antennule folding transversely or obliquely forwards. Endostomial ridges absent or rudimentary, rarely well developed. Buccal cavity approximately quadrate. Maxilliped 3 merus rounded to irregularly subquadrate or rectangular. Male pleonites 3–4, 3–5 or 4–5 fused, immovable, even if some or all sutures visible. Gonopod 1 stout to slender. Gonopod 2 usually shorter than one-third, occasionally half, gonopod 1 length.

Key to families of Xanthoidea 1. – 2. – 3. –

Male thoracic sternite 3–4 less than half as long as wide (width measured across posteriormost points of episternites 4) (Figs 14.142b, 14.147g)���������������������������������������������������������������������������������������������������������������������������������������2 Male thoracic sternite 3–4 half or more times as long as wide (Fig. 14.158h–n, 14.167e)�����������������������������������������������������3 Pereopods 2, 3 meri with spine on lower distal margin (Fig. 14.142d)����������������������������������������������������� Linnaeoxanthidae Pereopods 2, 3 meri without spine on lower distal margin (Fig. 14.145a, b)������������������������������������������ Pseudorhombilidae Males with part of lateral extremity of sternite 8 exposed, visible between lateral end of pleonite 2 and pereopod 5 coxa (sometimes very small) (Fig. 14.144j–l). Gonopod 1 tip with complex folds (Fig. 14.144a–g)��������������������������������������������������������������������������������������������������������������������������������������������������������� Panopeidae Males with sternite 8 usually completely covered by pleonite 2, rarely with lateral extremity of sternite 8 exposed. Gonopod 1 tip simple, with or without setae (Figs 14.153w, x, 14.158v–y, 14.167n–u)���������������� Xanthidae

References Dana JD (1852) Crustacea. Part I. United States Exploring Expedition during the years 1838, 1839, 1840, 1841, 1842 under the command of Charles Wilkes, U.S.N. 13, 1–685, with a folio atlas of 96 plates (published 1855). Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from

the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. Guinot D (1976) Constitution de quelques groupes naturels chez les Crustacés Décapodes Brachyoures: I. La superfamille des Bellioidea et trois sous-familles de Xanthidae (Polydectinae Dana, Trichiinae de Haan, Actaeinae Alcock). Mémoires du Muséum National d’Histoire Naturelle. Nouvelle Série. Série A. Zoologie 97, 1–308.

Fig. 14.141.  Xanthoidea. a, Dana’s (1852) scheme for labelling dorsal regions of the xanthoid carapace. b, thoracic sternites 1–3 (fused), conventional length and width dimensions fused sternites 3 and 4 indicated; e, episternite 4.

14 – Brachyura – crabs

Karasawa H, Schweitzer CE (2006) A new classification of the Xanthoidea sensu lato (Crustacea: Decapoda: Brachyura) based on phylogenetic analysis and traditional systematics and evaluation of all fossil Xanthoidea sensu lato. Contributions to Zoology 75, 23–73. doi:10.1163/18759866-0750102002 Lai JC, Mendoza JCE, Guinot D, Clark PF, Ng PKL (2011) Xanthidae MacLeay, 1838 (Decapoda: Brachyura: Xanthoidea) systematics: A multi-gene approach with support from adult and zoeal morphology. Zoologischer Anzeiger 250, 407–448. doi:10.1016/j.jcz.2011.07.002 Lai JCY, Thoma BP, Clark PF, Felder DL, Ng PKL (2014) Phylogeny of eriphioid crabs (Brachyura, Eriphioidea) inferred from molecular and morphological studies. Zoologica Scripta 43, 52–64. doi:10.1111/zsc.12030 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Serène R (1984) Crustacés Décapodes Brachyoures de l’Océan Indien Occidental et de la Mer Rouge, Xanthoidea: Xanthidae et Trapeziidae. Avec un addendum par Crosnier, A.: Carpiliidae et Menippidae. Faune Tropicale 24, 1–400. Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Thoma BP, Guinot D, Felder DL (2014) Evolutionary relationships among American mud crabs (Crustacea: Decapoda: Brachyura: Xanthoidea) inferred from nuclear and mitochondrial markers, with comments on adult morphology. Zoological Journal of the Linnean Society 170, 86–109. doi:10.1111/zoj.12093

Linnaeoxanthidae Števčić, 2005 Figure 14.142 The two species of Linnaeoxanthidae, one in the Central Indo-Pacific and the other in the tropical Western Atlantic, share numerous features, notably four sharp triangular anterolateral teeth; a broad, bilobed, deflexed front; large eyes; narrow, widely separate maxillipeds 3; long, unequal chelipeds, with spinose meri and carpi, flat chelae and short fingers; and spinose pereopods with a spine on the lower distal margin of at least pereopods 2 and 3 (Mendoza et al. 2012). These authors argued for their treatment as a family distinct from Xanthidae s.s. Diagnosis. Pereopods 2, 3 meri with spine on lower distal margin (Fig. 14.142d). Male thoracic sternite 3–4 less than half as long as wide (Fig. 14.142b). Gonopod 1 stout.

735

Linnaeoxantho Števčić, 2005 Diagnosis. Carapace 1.3 times as wide as long; front comprising 2 almost transverse lobes, with shallow lateral depression. Pereopods 2 and 3 with spine on distal lower margin. Maximum cl. 10 mm. Subtidal, shelf (6–55 m; amongst living or dead corals on reef slopes). Central Indo-Pacific. 1 species (Mendoza et al. 2012: rediagnosis, description of only species).

Melybia Stimpson, 1871 Diagnosis. Carapace 1.5 times as wide as long. Front comprising 2 oblique lobes, prominent submedially. Pereopods 2–4 with spine on distal lower margin. Maximum cl. 7.3 mm. Intertidal–slope (0–368 m; under Sargassum, coral and broken shell). Temperate Northern and Tropical W Atlantic. 1 species (Rathbun 1930: rediagnosis; Williams 1984: figure of only species). References Mendoza JCE, Clark PF, Ng PKL (2012) The identity of Pilumnoplax acanthomerus Rathbun, 1911 (Crustacea: Decapoda: Brachyura: Xanthidae), with new records from the central and western Pacific. Zootaxa 3367, 211–221. doi:10.11646/zootaxa.3367.1.20 Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Williams AB (1984) Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C.

Panopeidae Ortmann, 1893 Figures 14.143, 14.144, Plate 64a–c The panopeid crabs are largely confined to the Americas (with a few representatives off West Africa), inhabiting the shore to upper shelf from mostly muddy habitats. Accordingly, panopeids are commonly called mud crabs (not to be confused with the portunid genus, Scylla, which takes the same common name in the Indo-Pacific). Among the xanthoids, panopeids can be recognised by the combination of the proportionally narrow thoracic sternum, partially

Fig. 14.142.  Linnaeoxanthidae. Linnaeoxantho acanthomerus (Rathbun, 1911): a, habitus; b, thoracic sternum; c, maxilliped 3; d, pereopod 3; e, pereopod 5 merus. f, carapace, Melybia thalamita Stimpson 1871.

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Marine Decapod Crustacea

exposed thoracic sternite 8 in males, and elaborately ornamented and folded tip of gonopod 1. Panopeids have usually been treated as a subfamily of Xanthidae, but following review by Guinot (1978), are now well established as a separate family. Williams (1983) demonstrated a high level of unrealised diversity in the common genus Panopeus. Martin and Abele (1986) documented significant genus-level diversity in gonopod 1 morphology within the panopeids, but which nevertheless substantially conform to a common, trilobate ground-pattern. More recently, a close relationship between panopeids and pseudorhombilids was inferred by preliminary phylogenetic analyses, whose results also cast doubt on the monophyly of several panopeid genera, as well as the monophyly of all of the xanthoid families (Lai et al. 2011; Thoma et al. 2009). The close connections between panopeids and pseudorhombilids appear to corroborate, at least in part, Guinot’s (1969, 1978) observations of lineages having ‘fácies xanthien’ and ‘fácies gonéplacien’ among the panopeid-pseudorhombilid genera. Although Panopeidae and Pseudorhombilidae may be ultimately and substantially redefined or even merged, we treat the two families as separate, distinguished chiefly by the proportions of the thoracic

sternum. As such, several genera with a wide thoracic sternum placed in Panopeidae by Ng et al. (2008) are herein included in Pseudorhombilidae (Chasmophora, Cycloplax, Malacoplax, Prionoplax and Tetraplax). Homoioplax Rathbun, 1914, by default long retained in Panopeidae (e.g. Davie 2002; Ng et al. 2008), was shown to be a junior synonym of Carcinoplax in Goneplacidae (Ng et al. 2022). Diagnosis. Males with part of lateral extremity of sternite 8 exposed, visible between lateral end of pleonite 2 and pereopod 5 coxa (often barely so). Pleon half or more times as long as wide. Gonopod 1 tip with complex folds. Implicit generic characters. Carapace wider than long; front unimarginate; anterolateral margin with 4 teeth, first indistinguishable from exorbital, second-fourth not conspicuously curved forwards, triangular, pointed outward or slightly forward; posterior and posterolateral margins demarcated by angular or obtusely curved junctions, not forming evenly continuous curve. Endostomial ridges absent or not reaching to anterior margin of buccal frame. Pereopods 2–5 carpi upper margins convex, not bilobed. Male pleonite 3 reaching pereopod 5 coxae.

Key to genera of Panopeidae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8.

Carapace posterior and posterolateral margins forming continuous even, semicircular curve when viewed dorsally with carapace horizontal. Endostomial ridges complete, reaching to anterior margin of buccal frame (Fig. 14.143c)������������������������������������������������������������������������������������������������������������������������������������������������������Eurytium Carapace posterior and posterolateral margins demarcated by angular or obtusely curved junctions, not forming evenly continuous curve. Endostomial ridges absent or not reaching to anterior margin of buccal frame���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Pereopod 2–4 (sometimes also 5) carpi upper margins sinuous, concave medially, weakly to prominently bilobed AND/OR fronto-orbital border half carapace width in adults������������������������������������������������������������������������������3 Pereopod 2–5 carpi upper margins convex, not bilobed AND fronto-orbital border more than half carapace width in adults������������������������������������������������������������������������������������������������������������������������������������������������������������4 Fronto-orbital width usually half carapace width. Pereopods 2–5 meri upper margins smooth to weakly irregular (Fig. 14.144m)��������������������������������������������������������������������������������������������������� Lophopanopeus Fronto-orbital width 0.7 times carapace width (Fig. 14.143o. Pereopods 2–5 meri upper margins serrated, lined with angular or acute granules (Fig. 14.144n)���������������������������������������������������������������������Milnepanopeus Carapace as wide as long (Fig. 14.143n)��������������������������������������������������������������������������������������������������������������Metopocarcinus Carapace wider than long���������������������������������������������������������������������������������������������������������������������������������������������������������������5 Male pleonite 3 not reaching pereopod 5 coxae (Fig. 14.144j)���������������������������������������������������������������������������������������������������6 Male pleonite 3 reaching pereopod 5 coxae (Fig. 14.144k)��������������������������������������������������������������������������������������������������������9 Carapace front bilobed, each lobe concave (Fig. 14.143p). Male telson triangular���������������������������������������������Odontoplax Carapace front bilobed, each lobe straight or convex. Male telson triangular or rounded (Fig. 14.144j)��������������������������7 Carapace front bimarginate, lower margin visible in anterior and dorsal view (Fig. 14.143h, i). Male telson slightly longer than wide, margins in proximal half subparallel���������������������������������������Rhithropanopeus Carapace front unimarginate. Male telson wider than long, margins convergent����������������������������������������������������������������8 Carapace front gently convex (Fig. 14.143k). Maxilliped 3 merus distolateral angle square, blunt (Fig. 14.144h)��������������������������������������������������������������������������������������������������������������������������������������������������������������Eucratopsis

14 – Brachyura – crabs

– 9. – 10. – 11.

737

Carapace front approximately transverse (Fig. 14.143l). Maxilliped 3 merus distolateral angle produced, bluntly acute (Fig. 14.144i)��������������������������������������������������������������������������������������������������������������������������������������� Glyptoplax Carapace with posterior 3 anterolateral teeth swept forwards, apices usually acute, lateral margins curved forwards (Fig. 14.143a, b, f, g)��������������������������������������������������������������������������������������������������������������������������������������������������10 Carapace with posterior 3 anterolateral teeth triangular, pointed outward or slightly forward, lateral margins not conspicuously curved forwards (Fig. 14.143d, j, m)��������������������������������������������������������������������������������������13 Gonopod 1 without distal accessory process and lateral tooth (Fig. 14.144b)�������������������������������������������������� Dyspanopeus Gonopod 1 distal accessory process well developed, sharply tapering, distinctly longer than median process; lateral tooth present���������������������������������������������������������������������������������������������������������������������������������������������������11 Gonopod 1 accessory process lanceolate, largely straight, directed distally in line with gonopod axis; lateral tooth bilobed, directed laterally (Fig. 14.144g)��������������������������������������������������������������������������������������������������������Panopeus

Fig. 14.143.  Panopeidae. a, Dyspanopeus sayi (Say, 1869); b, Eurypanopeus abbreviatus (Stimpson, 1860); c, Eurytium limosum (Say, 1819); d, Hexapanopeus angustifrons (Benedict & Rathbun, 1891); e, Lophopanopeus bellus (Stimpson, 1860); f, Neopanope packardi (Kingsley, 1879); g, Panopeus lacustris Desbonne, 1867; h, Rhithropanopeus harrisii (Gould, 1841). i, front, orbits (oblique frontal view), Rhithropanopeus. Carapace: j, Acantholobulus; k, Eucratopsis; l, Glyptoplax; m, Lithopanopeus; n, Metopocarcinus; o, Milnepanopeus; p, Odontoplax.

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Marine Decapod Crustacea

Fig. 14.144.  Panopeidae. Gonopod 1 apex (sternomesial margin on right; ap = accessory process; mp = mesial process; lt = lateral tooth): a, Acantholobulus; b, Dyspanopeus; c Eurypanopeus; d, Hexapanopeus; e, Lithopanopeus; f, Neopanope; g, Panopeus. Maxilliped 3: h, Eucratopsis; i, Glyptoplax. Male sternites 7, 8, pleonites 1–6, telson, pereopodal coxa 5 (numbered): j, Glyptoplax; k, Lithopanopeus; l, Milnepanopeus; Pereopod 3: m, Lophopanopeus; n, Milnepanopeus.



Gonopod 1 accessory process straight or sharply bent over top of median process; lateral tooth minute, singular, slightly hooked (Fig. 14.144c, f)����������������������������������������������������������������������������������������������������������������������������� 12 12. Carapace transversely ovate, 1.4–1.6 times as wide as long; usually crossed by broken, transverse, granular lines (Fig. 14.143b). Maxilliped 3 ischium with large red patch on inner surface������������������������������������� Eurypanopeus – Carapace subhexagonal-subpentagonal, 1.3 times as wide as long; surface smooth, without broken, transverse, granular lines (Fig. 14.143f). Maxilliped 3 ischium without red patch on inner surface���������� Neopanope 13. Gonopod 1 sternomesial margin with row of broad, stout subacute teeth, row ending well short of lateral tooth (Fig. 14.144e)����������������������������������������������������������������������������������������������������������������������������������������������Lithopanopeus – Gonopod 1 sternomesial margin unarmed or at most with row of minute spinules (Fig. 14.144a, d)�����������������������������14 14. Gonopod 1 accessory process an acute spine, shorter than to overreaching median process (Fig. 14.144a)������������������������������������������������������������������������������������������������������������������������������������������������������ Acantholobulus – Gonopod 1 accessory process absent (Fig. 14.144d)��������������������������������������������������������������������������������������������Hexapanopeus Acantholobulus Felder & Martin, 2003

Dyspanopeus Martin & Abele, 1986

Diagnosis. Carapace front bilobed, usually bimarginate, lower margin visible in anterior and dorsal view; posterior 3 anterolateral teeth triangular, pointed outward or slightly forward, lateral margins not conspicuously curved forwards. Gonopod 1 accessory process an acute spine, shorter than to overreaching median process; sternomesial margin unarmed or at most with row of minute spinules. Maximum cl. 12 mm. Intertidal, subtidal (0–15 m). Tropical W Atlantic, Temperate South America (Atlantic coast). 7 species (Felder and Martin 2003). Acantholobulus pacificus (Edmondson, 1931) has been recorded from Hawaii, New Zealand and Japan as an extralimital species (Komai and Furota 2013). Eurypanopeus hyperconvexus Garth, 1986, and Hexapanopeus beebei Garth, 1961 are herein transferred to Acantholobulus based on carapace shape and gonopod features.

Diagnosis. Carapace front bilobed, gently convex; posterior 3 anterolateral teeth swept forwards, apices usually acute, lateral margins curved forwards. Gonopod 1 without distal accessory process and lateral tooth. Maximum cl. 21 mm. Estuarine, intertidal–shelf (0–79 m). Temperate Northern and Tropical W Atlantic. 2 species (Abele 1972: key to 2 species as Neopanope, redescriptions). One species, D. sayi (Smith, 1869) has been introduced to the north-eastern Atlantic, including to the Mediterranean Sea and the Black Sea (Micu et  al. 2010).

Eucratopsis Smith, 1869 Diagnosis. Carapace front bilobed, gently convex, front unimarginate; posterior 3 anterolateral teeth triangular, pointed

14 – Brachyura – crabs

outward or slightly forward, lateral margins not conspicuously curved forwards. Maxilliped 3 merus distolateral angle square, blunt. Male pleonite 3 not reaching pereopod 5 coxae. Male telson rounded, wider than long. Maximum cl. 21 mm. Shelf, slope (23–350 m). Tropical W Atlantic. 1 species (Rathbun 1918: description, figure).

Eurypanopeus A. Milne-Edwards, 1880 Diagnosis. Carapace transversely ovate, 1.4–1.6 times as wide as long; usually crossed by broken, transverse, granular lines; front bilobed, approximately transverse; posterior 3 anterolateral teeth swept forwards, apices usually acute, lateral margins curved forwards. Gonopod 1 with accessory process straight or sharply bent over top of median process; lateral tooth minute, singular, slightly hooked. Maximum cl. 24 mm. Estuarine, intertidal, subtidal. Temperate Northern W Atlantic, Tropical Atlantic, Tropical Eastern Pacific, Temperate South America (Atlantic coast). 13 species (Rathbun 1930). Eurypanopeus depressus (Smith, 1869) has been introduced to the Black Sea (Petrescu et al. 2016), and E. dissimilis (Benedict & Rathbun, 1891) has been introduced from the Tropical Western Atlantic to the Tropical Eastern Pacific via the Panama Canal (Abele and Kim 1989). Eurypanopeus hyperconvexus Garth, 1986, with a comparatively narrow carapace and blunt anterolateral teeth, is herein transferred to Acantholobulus.

Eurytium Stimpson, 1859 Diagnosis. Carapace front bilobed, approximately transverse; posterior 3 anterolateral teeth swept forwards, apices usually acute, lateral margins curved forwards; posterior and posterolateral margins forming continuous, even, semicircular curve when viewed dorsally with carapace horizontal. Endostomial ridges complete, extending anteriorly to epistome. Maximum cl. 34 mm (Pl. 64a). Estuarine, intertidal–shelf (0–71 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific. 4 species (Rathbun 1930: rediagnosis, key to 3 species).

739

Gonopod 1 accessory process absent; sternomesial margin unarmed or at most with row of minute spinules. Maximum cl. 20 mm. Intertidal–shelf (0–139 m). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific, Temperate South America. 9 species (Rathbun 1930: key to 6 species together with H. schmitti Rathbun, 1930 and H. caribbaeus (Stimpson, 1871), currently placed in Acantholobulus; Thoma et  al. 2009: phylogeny). Hexapanopeus ­beebei Garth, 1961 is transferred to Acantholobulus.

Lithopanopeus Felder & Thoma, 2020 Diagnosis. Carapace front bilobed, approximately transverse; fronto-orbital width exceeding half carapace width. Gonopod 1 sternomesial margin with row of broad, stout subacute teeth, row ending well short of lateral tooth. Maximum cl. 7 mm. Intertidal, subtidal (0–1 m). Tropical Atlantic (Gulf of Mexico). 1 species (Felder and Thoma 2020).

Lophopanopeus Rathbun, 1898 Diagnosis. Carapace front bilobed, approximately transverse; front-exorbital width 0.5 times carapace width. Pereopods 2–4 (sometimes also 5) carpi upper margins sinuous, concave medially, weakly to prominently bilobed. Pereopods 2–5 5 meri upper margins smooth to weakly irregular. Maximum cl. 23 mm. Intertidal–shelf (0–102 m). Temperate Northern and Tropical Eastern Pacific. 5 species (Menzies 1948: key to species).

Metopocarcinus Stimpson, 1860 Diagnosis. Carapace as wide as long; front bilobed, concave, fronto-orbital border more than half carapace width in adults; anterolateral margin with obsolete first lobe, 3 lobes on lateral margin. Pereopod 2–5 carpi upper margins convex, not bilobed. Maximum cl. 5.6 mm. Intertidal. Tropical Eastern Pacific, Temperate South America (Mexico to Chile). 2 species (Crane 1947: species comparison; Rathbun 1930).

Glyptoplax Smith, 1870

Milnepanopeus Thoma & Felder, 2012

Diagnosis. Front bilobed, approximately transverse; posterior 2 or 3 anterolateral teeth triangular, pointed outward or slightly forward, lateral margins not conspicuously curved forwards. Maxilliped 3 merus distolateral angle produced, bluntly acute. Male pleonite 3 not reaching pereopod 5 coxae. Male telson rounded to subtriangular, wider than long. Maximum cl. 21 mm. Shelf (20–110 m). Tropical W Atlantic, Tropical Eastern Pacific. 3 species (Hendrickx 1989: species comparison; Rathbun 1918).

Diagnosis. Carapace front bilobed, approximately transverse; front-exorbital width 0.7 times carapace width. Maxilliped 3 merus distolateral angle square, blunt. Pereopods 2–5 carpi upper margins sinuous, concave medially, weakly to prominently bilobed. Pereopods 2–5 meri upper margins serrated, lined with angular or acute granules. Maximum cl. 13 mm. Intertidal–shelf (0–84 m). Tropical W Atlantic, Tropical Eastern Pacific. 1 species (Thoma and Felder 2012).

Hexapanopeus Rathbun, 1898 Diagnosis. Carapace front bilobed, approximately transverse; posterior 3 anterolateral teeth triangular, pointed outward or slightly forward, lateral margins not conspicuously curved forwards.

Neopanope A. Milne-Edwards, 1880 Diagnosis. Carapace subhexagonal-subpentagonal, 1.3 times as wide as long; smooth; front bilobed, gently convex; posterior 3 teeth swept forwards, apices usually acute, lateral margins curved forwards. Gonopod 1 with accessory process sharply bent over top

740

Marine Decapod Crustacea

of median process; lateral tooth minute, singular, slightly hooked. Maximum cl. 15 mm. Subtidal, shelf (1–74 m). Tropical W Atlantic. 1 species (Abele 1972: description of only species, N. packardii (Kingsley, 1879)).

Odontoplax Garth, 1986 Diagnosis. Carapace front bilobed, each lobe concave; anterolateral margin with 3 conical submarginal teeth. Male pleonite 3 not reaching pereopod 5 coxae. Maximum cl. 11 mm. Subtidal, shelf (4–50 m). Tropical Eastern Pacific. 1 species (Garth 1986).

Panopeus H. Milne Edwards, 1834 Diagnosis. Carapace front bilobed, approximately transverse; posterior 3 anterolateral teeth swept forwards, apices usually acute, lateral margins curved forwards. Gonopod 1 with accessory process lanceolate, largely straight, directed distally in line with gonopod axis; lateral tooth bilobed, directed laterally. Maximum cl. 43 mm (Pl. 64b). Estuarine, intertidal–shelf (0–59 m). Temperate Northern W Atlantic, Tropical Atlantic, Tropical Eastern Pacific, Temperate South America (Atlantic, Pacific coasts). 16 species (Rathbun 1930: key to 14 species including P. bermudensis Benedict & Rathbun, 1891 (currently in Acantholobulus) and P. turgidus Rathbun, 1930 (currently in Eurypanopeus); Williams 1983: review of Panopeus herbstii complex and key to 6 species). Panopeus lacustris Desbonne, 1867 has been introduced to Hawaii (Brockerhoff and McLay 2011).

Rhithropanopeus Rathbun, 1898 Diagnosis. Carapace front bilobed, bimarginate, lower margin visible in anterior view. Male pleonite 3 not reaching pereopod 5 coxae. Male telson rounded, slightly longer than wide, margins in proximal half subparallel. Maximum cl. 16 mm (Pl. 64c). Freshwater, estuarine, intertidal–shelf (0–37 m). Temperate Northern and Tropical W Atlantic. 1 species (Squires 1990: figures; Williams 1984). The only species, R. harrisii (Gould, 1841) is invasive in the tropical eastern Pacific and north-eastern Atlantic, including the Mediterranean Sea, Black Sea and Caspian Sea (Brockerhoff and McLay 2011; Roche and Torchin 2007). References Abele LG (1972) A reevaluation of the Neopanope texana-sayi complex with notes on N. packardii (Crustacea: Decapoda: Xanthidae) in the western Atlantic. Chesapeake Science 13, 263–271. doi:10.2307/1351110 Abele LG, Kim W (1989) The decapod crustaceans of the Panama Canal. Smithsonian Contributions to Zoology 482, 1–50. doi:10.5479/si.​ 00810282.482 Brockerhoff A, McLay C (2011) Human-mediated spread of alien crabs. In In the Wrong Place – Alien Marine Crustaceans: Distribution, Biology and Impacts. (Eds Galil BS, Clark PF, Carlton JT) pp. 27–106. Springer, Dordrecht. Crane J (1947) Eastern Pacific Expeditions of the New York Zoological Society. XXXVIII. Intertidal brachygnathous crabs from the west

coast of tropical America with special reference to ecology. Zoologica 32, 69–95. Davie PJF (2002) Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells A, Houston WWK, Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing, Melbourne. Felder DL, Martin JW (2003) Establishment of a new genus for Panopeus bermudensis Benedict and Rathbun, 1891 and several other xanthoid crabs from the Atlantic and Pacific oceans (Crustacea: Decapoda: Xanthoidea). Proceedings of the Biological Society of Washington 116, 438–452. Felder DL, Thoma BP (2020) A new genus and species of mud crab (Crustacea, Brachyura, Panopeidae) from shoreline waters of the western Gulf of Mexico. Zootaxa 4845, 425–435. doi:10.11646/zootaxa.4845.3.7 Garth JS (1986) A new genus and species of goneplacid crab (Decapoda: Brachyura) from the west coast of South and Central America. Journal of Crustacean Biology 6, 543–546. doi:10.2307/1548193 Guinot D (1969) Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures. VII. Les Goneplacidae (suite et fin). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 41, 688–724, pls 1–5. Guinot D (1978) Principes d’une classification évolutive des Crustacés Décapodes Brachyoures. Bulletin Biologique de la France et de la Belgique 112, 211–292. Hendrickx ME (1989) Glyptoplax consagae new species (Crustacea, Decapoda, Brachyura, Panopeidae) from the Gulf of California, Mexico, with some notes on the distribution of G. pugnax Smith, 1870. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 11, 649–657. Komai T, Furota T (2013) A new introduced crab in the western North Pacific: Acantholobulus pacificus (Crustacea: Decapoda: Brachyura: Panopeidae), collected from Tokyo Bay, Japan. Marine Biodiversity Records 6, e96. doi:10.1017/S1755267213000699 Lai JC, Mendoza JCE, Guinot D, Clark PF, Ng PKL (2011) Xanthidae MacLeay, 1838 (Decapoda: Brachyura: Xanthoidea) systematics: A multi-gene approach with support from adult and zoeal morphology. Zoologischer Anzeiger 250, 407–448. doi:10.1016/j.jcz.2011.07.002 Martin JW, Abele LG (1986) Notes on male pleopod morphology in the brachyuran crab family Panopeidae Ortmann, 1893, sensu Guinot (1978) (Decapoda). Crustaceana 50, 182–198. doi:10.1163/​156854086X00205 Menzies RJ (1948) A revision of the brachyuran genus Lophopanopeus. Occasional Papers of the Allan Hancock Foundation 4, 1–45. Micu D, Niţă V, Todorova V (2010) First record of Say’s mud crab Dyspanopeus sayi (Brachyura: Xanthoidea: Panopeidae) from the Black Sea. Marine Biodiversity Records 3, e36. doi:10.1017/S1755267210000308 Ng PKL, Clark PF, Ahyong ST (2022) The identity of Homoioplax haswelli (Miers, 1884) (Crustacea: Decapoda: Brachyura). Zoological Studies 61, 6. doi:10.6620/ZS.2022.61-06 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Petrescu I, Petrescu A-M, Popescu-Mirceni R (2016) First record of Eurypanopeus depressus (Smith, 1869) (Brachyura, Panopeidae) from the Black Sea. Crustaceana 89, 141–149. doi:10.1163/15685403-00003519 Rathbun MJ (1918) The grapsoid crabs of America. Bulletin of the United States National Museum 97, 1–461, pls 1–161. Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Roche DG, Torchin ME (2007) Established population of the North American Harris mud crab, Rhithropanopeus harrisii (Gould 1841) (Crustacea: Brachyura: Xanthidae) in the Panamá Canal. Aquatic Invasions 2, 155–161. doi:10.3391/ai.2007.2.3.1 Squires HJ (1990) Decapod Crustacea of the Atlantic coast of Canada. Canadian Bulletin of Fisheries and Aquatic Sciences 221, 1–532.

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Thoma BP, Felder DL (2012) Redescription of Hexapanopeus lobipes and Its reassignment to Milnepanopeus n. gen. (Decapoda: Brachyura: Panopeidae). Journal of Crustacean Biology 32, 141–152. doi:10.1163/​ 193724011X615415 Thoma BP, Schubart CD, Felder DL (2009) Molecular phylogeny of Western Atlantic representatives of the genus Hexapanopeus (Decapoda: Brachyura: Panopeidae). In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 551–565. CRC Press, Boca Raton. Williams AB (1983) The mud crab, Panopeus herbstii, s.l. partition into six species (Decapoda: Xanthidae). Fishery Bulletin 81, 863–882. Williams AB (1984) Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C.

Pseudorhombilidae Alcock, 1900 Figures 14.145–14.148, Plate 64d, e Pseudorhombilids are xanthoid crabs mostly from the Americas in which the male thoracic sternite 3–4 is less than half as long as wide and the merus of pereopods 2 and 3 lacks a spine on the lower distal margin. The family name was introduced as a subfamily of Goneplacidae and its members have long been recognised has having affinities with this family (Guinot 1969a, 1969b; Hendrickx 1998), but Ng et al. (2008) argued for a closer relationship to the xanthoids, particularly panopeids. Hendrickx (1998) rediagnosed the family but included only six genera. This number was doubled in a phylogenetic analysis based on molecular data (Thoma et al. 2014). These authors suggested that the short broad thoracic sternum diagnosed the family but ambiguities remain in relation to whether pseudorhomibilds, panopeids and xanthids are each monophyletic with respect to

741

each other; further division and generic realignments among families are likely in the future (Lai et  al. 2011; Thoma et al. 2014). Here, we emphasise the wide thoracic sternum in diagnosing Pseudorhombilidae. The family was divided into five families or subfamilies and ten tribes by Števčić (2005), some now synonymised, moved to other families or no longer recognised (Ng et  al. 2008; Thoma et al. 2014). Ng et al. (2008) listed Chasmophora, Cycloplax, Malacoplax, Prionoplax and Tetraplax in Panopeidae, and Eucratodes, Garthiope and Gonopanope in Xanthidae. All have the male thoracic sternite 3–4 less than half as long as wide and we include these genera in Pseudorhombilidae. Guinope was tentatively placed in Linnaeoxanthidae (Thoma and Felder 2020) but it too is included in Pseudorhombilidae. The anterolateral margin of the carapace of pseudorhombilids usually comprises four teeth or lobes in addition to the exorbital angle. The exorbital tooth or angle is often more or less coalesced with the first tooth, often just an obtuse angle. The second and third teeth are the largest; the fourth is usually minute, sometimes absent. In Pseudopanopeus and Scopolius the first tooth is interpreted as being fused to the exorbital; Prionoplax and Trapezioplax have only two teeth besides the exorbital tooth. Diagnosis. Pereopods 2, 3 meri without spine on lower distal margin (Fig.  14.145a, b). Male thoracic sternite 3–4 less than half as long as wide (Fig. 14.147g). Gonopod 1 with distal convoluted, foliaceous apex, sometimes with strong, long setae and spines (Fig. 14.148). Implicit generic characters. Maxilliped 3 merus distolateral angle rounded. Pereopods 2–5 stout, longest little longer than carapace width.

Key to genera of Pseudorhombilidae 1. – 2. – 3. – 4. –

Carapace anterolateral margin with blunt exorbital tooth plus 3 teeth: first obtuse, second and third spine-like, set one behind other, well separated (Fig. 14.145b)�����������������������������������������������������������������Perunorhombila Carapace anterolateral margin usually with 4 teeth, rarely 2 plus exorbital, largest triangular, set more or less obliquely��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Carapace anterolateral margin with exorbital tooth and anterolateral teeth well separated, spinulose; front with lateral spinulose lobe (Fig. 14.145a)���������������������������������������������������������������������������������������������������������������������������������3 Carapace anterolateral margin with exorbital tooth and anterolateral teeth not spinulose; front without spinulose lateral lobe������������������������������������������������������������������������������������������������������������������������������������������������������������������4 Carapace exorbital tooth and first anterolateral tooth fused into spinulose margin (Fig. 14.145i). Gonopod 1 with spatulate spinulose apex, with short subapical spines (Fig. 14.148g)������������������������������������������������������Garthiope Carapace exorbital tooth and first anterolateral tooth separate (Fig. 14.145a). Gonopod 1 twisted, apex with subdistal lateral triangular flange, without spines below flange (Fig. 14.148f)������������������������������������������������������������������������������������������������������������������������������������������������������ Euphrosynoplax Carapace ovate; anterolateral margin without teeth (Fig. 14.145f). Gonopod 1 stout, with twisted tip with 2 apices (Fig. 14.148d)���������������������������������������������������������������������������������������������������������������������������������������� Cycloplax Carapace subovate or subquadrate; anterolateral margin with teeth. Gonopod 1 otherwise����������������������������������������������5

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5. – 6. – 7. – 8. – 9. – 10. – 11.

Marine Decapod Crustacea

Exorbital tooth prominent or acute, followed by 2 triangular anterolateral teeth set one behind other. Front-exorbital margin as wide or almost as wide as carapace (Fig. 14.146i, s)����������������������������������������������������������������6 Exorbital tooth usually obsolete, often fused with first anterolateral tooth; with 3 or 4 anterolateral teeth. Front-exorbital margin rarely almost as wide as carapace���������������������������������������������������������������������������������������������������7 Carapace exorbital tooth directed anteriorly (Fig. 14.146s). Gonopod 1 almost straight, with tubular apex, with subapical short spines (Fig. 14.148v)������������������������������������������������������������������������������������������������Trapezioplax Carapace exorbital tooth directed laterally (Fig. 14.146i). Gonopod 1 with 1 lobular and 1 acute apical projection; 1 short subdistal projection; short subapical spines (Fig. 14.148o)����������������������������������������������� Prionoplax Pereopods 2–5 flat, carpi with bilobed carina on extensor margin (Fig. 14.146c)������������������������������������������Lophoxanthus Pereopods 2–5 linear, carpi without carina on extensor margin����������������������������������������������������������������������������������������������8 Carapace anterolateral margin convex, all teeth obsolete: exorbital tooth coalesced with obtuse first anterolateral; second and third broad, short, elevated; fourth minute (Fig. 14.146d). Gonopod 1 slender, with bilobed apex and subapical projection; with apical setae (Fig. 14.148k)����������������������������������Malacoplax Carapace anterolateral margin with 1 or more teeth well defined. Gonopod 1 otherwise��������������������������������������������������9 Front-exorbital margin almost as wide as carapace; exorbital and 3 anterolateral teeth similar, well defined (Figs 14.145e, 14.146p)��������������������������������������������������������������������������������������������������������������������������������������������������������������10 Front-exorbital margin 0.8 or less carapace width; exorbital and anterolateral teeth dissimilar, some ill-defined�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������11 Carapace 1.6 times as wide as long, convex (Fig. 14.145e); anterior 3 anterolateral teeth of similar size, last tooth minute. Gonopod 1 with 2 subapical lobes, short spines on triangular apex (Fig. 14.148c)��������������������������������������������������������������������������������������������������������������������������������������������������������� Chasmophora Carapace 1.2 times as wide as long, flat (Fig. 14.146p); 4 similar anterolateral teeth. Gonopod 1 with 1 lobular and 1 acute apical projection; 1 short subdistal projection; short subapical spines (Fig. 14.148t)������������������������������������������������������������������������������������������������������������������������������������������������������������������Tetraplax Carapace ovate, ~1.7 times as wide as long (Fig. 14.146l). Gonopod 1 slender, with broad proximolateral flange, apex with terminal recurved flap (Fig. 14.148r)�������������������������������������������������������������������������������������� Robertsella

Fig. 14.145.  Pseudorhombilidae. a, Euphrosynoplax campechiensis Vázquez-Bader & Gracia, 1991; b, Perunorhombila nitida Chace, 1940. Carapace: c, Bathyrhombila; d, Chacellus; e, Chasmophora; f, Cycloplax; g, Cyrtoplax; h, Eucratodes; i, Garthiope.

14 – Brachyura – crabs

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Fig. 14.146.  Pseudorhombilidae. Carapace: a, Gonopanope; b, Guinope; c, Lophoxanthus (with pereopods); d, Malacoplax: e, Micropanope; f, Nanoplax; g, Oediplax; h, Panoplax; i, Prionoplax; j, Pseudopanopeus; k, Pseudorhombila; l, Robertsella; m, Scopolius; n, o, Speocarcinus (dorsal, lateral); p, Tetraplax; q, Tetraxanthus; r, Thalassoplax; s, Trapezioplax (with pereopods).

– Carapace ovate or subquadrate, no more than 1.5 times as wide as long. Gonopod 1 otherwise������������������������������������ 12 12. Carapace exorbital and first anterolateral tooth indistinguishable; front half carapace width (Fig. 14.146j, m)������������������������������������������������������������������������������������������������������������������������������������������������������������������������ 13 – Carapace exorbital and first anterolateral tooth coalesced but distinguishable; front less than half carapace width�������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������14 13. Carapace anterolateral margin with second anterolateral (first separate) tooth triangular (Fig. 14.146m). Male with major chela twice diameter of minor chela. Gonopod 1 ‘distally bulbous, with recurved apical projection’ (not figured)���������������������������������������������������������������������������������������������������������������������������������������������� Scopolius – Carapace anterolateral margin with second anterolateral (first separate) tooth truncate (Fig. 14.146j). Male with major chela slightly larger than minor chela. Gonopod 1 with subapical large, proximally directed spine (sometimes 2), with apical weakly curved finger (Fig. 14.148p)�������������������������������������Pseudopanopeus 14. Carapace margin between exorbital tooth and first anterolateral up to twice orbit width. Front-exorbital margin half carapace width (Figs 14.145c, d, 14.146g, q)����������������������������������������������������������������������������������������������������15 – Carapace margin between exorbital tooth and first anterolateral as long or less than orbit width. Frontexorbital margin distinctly greater than carapace width���������������������������������������������������������������������������������������������������18 15. Carapace third anterolateral tooth most prominent; fourth and fifth teeth absent (Fig. 14.146q)��������������� Tetraxanthus – Carapace second and third anterolateral teeth similar, neither prominent; at least fourth present���������������������������������16 16. Carapace second anterolateral tooth lobe-like (Fig. 14.145c). Gonopod 1 twisted, with 2 subapical series of long spines, with subterminal process (Fig. 14.148a)������������������������������������������������������������������������������������ Bathyrhombila – Carapace second anterolateral tooth acute. Gonopod 1 otherwise����������������������������������������������������������������������������������������17

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Marine Decapod Crustacea

Fig. 14.147.  Pseudorhombilidae. Maxilliped 3: a, Chacellus; b, Cyrtoplax; c, Guinope; d, Nanoplax; e, Pseudorhombilia. f, g, male pleon, thoracic sternites, Nanoplax.

Fig. 14.148.  Pseudorhombilidae. Gonopod 1 and/or detail of apex: a, Bathyrhombila; b, Chacellus; c, Chasmophora; d, Cycloplax; e, Cyrtoplax; f, Euphrosynoplax; g, Garthiope; h, Gonopanope ; i, j, Guinope (with gonopod 2); k, Malacoplax; l, Micropanope; m, Nanoplax; n, Panoplax; o, Prionoplax; p, Pseudopanopeus; q, Pseudorhombila; r, Robertsella; s, Speocarcinus; t, Tetraplax; u, Thalassoplax; v, Trapezioplax.

14 – Brachyura – crabs

17. – 18. – 19. – 20. – 21. – 22. – 23. – 24. – 25. – 26. –

745

Carapace front straight, with median notch; fifth anterolateral tooth absent (Fig. 14.145d). Gonopod 1 slender, tapering, with twisted apical lappet (Fig. 14.148b)�����������������������������������������������������������������������������������Chacellus Carapace front bilobed, with broad median excavation; fifth anterolateral tooth sharp (Fig. 14.146g). Gonopod 1 unknown���������������������������������������������������������������������������������������������������������������������������������������������������Oediplax Maxilliped 3 merus distolateral angle produced laterally (Fig. 14.147d, e)���������������������������������������������������������������������������19 Maxilliped 3 merus distolateral angle square or rounded, not produced laterally (Fig. 14.147b, c)���������������������������������21 Carapace anterolateral margin with second and third teeth prominent, acute (Fig. 14.146r). Gonopod 1 stout, apex twisted, with subdistal row of short spines (Fig. 14.148u)���������������������������������������������������������� Thalassoplax Carapace anterolateral margin with second and third teeth triangular, blunt. Gonopod 1 with reflexed apex��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 20 Gonopod 1 slender, apex with subterminal or lateral upturned flange, with patch of strong spines below flange (Fig. 14.148q). Carapace subquadrate-trapezoidal, posterior margin wider than front-orbital width (Fig. 14.146k)������������������������������������������������������������������������������������������������������������������������������������������ Pseudorhombila Gonopod 1 twisted, flat, apex with flap folded proximally, with long subterminal spines (Fig. 14.148m). Carapace oval-trapezoidal, posterior margin about as wide as front-orbital width (Fig. 14.146f)���������������� Nanoplax Carapace exorbital tooth and first anterolateral tooth separated by short concavity�������������������������������������������������������� 22 Carapace exorbital tooth and first anterolateral tooth almost indistinguishable��������������������������������������������������������������� 25 Carapace subquadrate to suboval or hexagonal, 1.7 times as wide as long (Fig. 14.146h). Gonopod 1 slender, curved, with apical curved lappet (Fig. 14.148n)����������������������������������������������������������������������������������������������������� Panoplax Carapace subquadrate to suboval, 1.4–1.5 times as wide as long. Gonopod 1 otherwise�������������������������������������������������� 23 Carapace subovate or rounded-subquadrate; anterolateral teeth slightly upturned; front 0.3 carapace width (Fig. 14.146n, o). Gonopod 1 slender, with reflexed flattened apex, with short subapical spines (Fig. 14.148s)������������������������������������������������������������������������������������������������������������������������������������������������������������Speocarcinus Carapace subquadrate or hexagonal; anterolateral teeth not upturned; front 0.4 carapace width. Gonopod 1 otherwise����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 24 Carapace subhexagonal, with second and third anterolateral teeth triangular (Fig. 14.146e). Gonopod 1 stout, with short apical and subapical flaps (Fig. 14.148l)��������������������������������������������������������������������������������Micropanope Carapace subquadrate, with second and third anterolateral teeth subquadrate, with acute tip (Fig. 14.145g). Gonopod 1 slender, curved, with 1 subapical acute lobe and 1 rounded lappet (Fig. 14.148e)����������������������� Cyrtoplax Carapace exorbital tooth and first anterolateral tooth coalesced as denticulate margin; second and third anterolateral teeth acutely triangular; front bilobed. Eyestalk 1.5 times as long as wide. (Fig. 14.146b). Gonopod 1 slender, sinuous, with subdistal constriction and spatulate apex (Fig. 14.148i)������������������������������Guinope Carapace exorbital tooth and first anterolateral tooth coalesced as smooth margin; second and third anterolateral teeth obsolete or triangular. Eyestalk twice as long as wide (Figs 14.145h, 14.146a). Gonopod 1 otherwise�������������������������������������������������������������������������������������������������������������������������������������������������������������� 26 Distance between exorbital and first anterolateral tooth equal to orbit width; third and fourth teeth blunt (Fig. 14.145h). Gonopod 1 unknown���������������������������������������������������������������������������������������������������������������������� Eucratodes Distance between exorbital and first anterolateral tooth one-half orbit width; third and fourth teeth acute (Fig. 14.146a). Gonopod 1 stout, with twisted apex carrying 2 groups of strong setae (Fig. 14.148h)����������������������������������������������������������������������������������������������������������������������������������������������������������� Gonopanope Bathyrhombila Hendrickx, 1998

Diagnosis. Carapace subquadrate, 1.4 times as wide as long; front 0.3 carapace width, almost straight; front-exorbital width 0.5 times carapace width; anterolateral margin with exorbital tooth and first anterolateral tooth obsolete; second–fourth triangular, lying almost one behind other. Gonopod 1 twisted, with 2 subapical series of long spines, with subterminal process. Maximum cl. 18 mm.

Shelf, slope (55–644 m). Tropical Eastern Pacific. 1 species (Hendrickx 1998). Chacellus Guinot, 1969 Diagnosis. Carapace trapezoidal, 1.4 times as wide as long; front 0.3 carapace width, almost straight, with median notch; frontexorbital width 0.45 times carapace width; anterolateral margin with exorbital tooth small; first anterolateral blunt, separated

746

Marine Decapod Crustacea

from exorbital by shallow depression, distance between two greater than orbit width; second–fourth triangular, acute, diminishing, more prominent than second. Maxilliped 3 merus distolateral angle produced. Pereopods 2–5 elongate, longest ~1.5 times carapace width. Gonopod 1 slender, tapering, with twisted apical lappet. Maximum cl. 43 mm. Shelf–slope (61–240 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Guinot 1969b: diagnosis; Hendrickx 1989: second species).

Chasmophora Rathbun, 1914 Diagnosis. Carapace subquadrate, 1.6 times as wide as long, convex; front 0.4 carapace width, almost straight, with median notch; front–exorbital width 0.9 times carapace width; anterolateral margin with prominent triangular exorbital tooth; 2 similar triangular teeth; third tooth minute; oblique-longitudinal. Maxilliped 3 merus distolateral angle produced. Gonopod 1 stout, with 2 subapical lobes, short spines on triangular apex. Maximum cl. 3 mm. Shelf (92 m). Tropical Eastern Pacific (Panama). 1 species (Guinot 1969b: figures; Rathbun 1918: description, figures).

Cycloplax Guinot, 1969 Diagnosis. Carapace ovate, 1.2 times as wide as long; front 0.3 carapace width, slightly projecting, with small median notch; front-exorbital width 0.6 times carapace width; anterolateral margin with obsolete exorbital tooth, convex, without teeth. Maxilliped 3 merus distolateral angle produced. Gonopod 1 stout, with twisted tip with 2 apices. Maximum cl. 7 mm. Subtidal (15–20 m). Tropical Atlantic (French Guiana). 1 species (Guinot 1969a).

Cyrtoplax Rathbun, 1914 Diagnosis. Carapace subquadrate, 1.5 times as wide as long; front 0.4 carapace width, with 2 weak lobes; front-exorbital width 0.7 times carapace width; anterolateral margin with exorbital tooth small; first anterolateral lobiform, separated from exorbital by shallow depression; second and third subquadrate, with acute tip; fourth small, acute. Gonopod 1 slender, curved, with 1 subapical acute lobe, 1 rounded lappet. Maximum cl. 15 mm. Intertidal–shelf (0–126 m). Tropical W Atlantic, Tropical Eastern Pacific. 4 species (Garth 1940: species description; Guinot 1969a: remarks).

Eucratodes A. Milne-Edwards, 1880 Diagnosis. Carapace ovate, 1.4 times as wide as long; front 0.3 carapace width, almost straight; front-exorbital width 0.6 times carapace width; anterolateral margin with obtuse exorbital tooth; first and second anterolateral teeth obtuse, third and fourth blunt, triangular; oblique. Maximum cl. 6.2 mm. Slope (250–313 m). Tropical Atlantic (Mexico). 1 species (Rathbun 1930: rediagnosis).

Euphrosynoplax Guinot, 1969 Diagnosis. Carapace subovate, 1.3 times as wide as long; front 0.3 carapace width, with U-shaped median notch and lateral tooth; front-exorbital width 0.5 times carapace width; anterolateral margin with exorbital tooth separated from sharp first anterolateral by concavity; second–fourth triangular, anteriorly-directed, all denticulate. Maxilliped 3 merus distolateral angle produced. Gonopod 1 stout, strong, twisted, apex with subdistal lateral triangular flange, without spines below flange. Maximum cl. 24 mm. Shelf (47–171 m). Tropical W Atlantic. 2 species (Guinot 1969b: diagnosis; Vázquez-Bader and Gracia 1991: species compared).

Garthiope Guinot, 1990 Diagnosis. Carapace ovate, 1.5 times as wide as long, spinulose; front 0.3 carapace width, with U-shaped median notch and lateral tooth; front-exorbital width 0.7 times carapace width; anterolateral margin with exorbital tooth and first anterolateral tooth together spinulose; second–fourth teeth well separated, spinulose. Gonopod 1 stout, curved, with spatulate spinulose apex, with short subapical spines. Maximum cl. 10.4 mm. Intertidal–shelf, marine anchialine caves (0–82 m). Tropical W Atlantic, Tropical Eastern Pacific (Galapagos). 4 species (Guinot 1990: diagnosis, key to 3 species; Guinot and Iliffe 1991: fourth species). In a molecular analysis, Thoma et  al. (2009) recovered Garthiope barbadensis (Rathbun, 1921) as closer to pseudorhombilids than to panopeids.

Gonopanope Guinot, 1967 Diagnosis. Carapace subquadrate, 1.5 times as wide as long; front 0.4 carapace width, with 2 weak lobes; front-exorbital width 0.8 times carapace width; anterolateral margin with exorbital and first anterolateral tooth together sinuous; second and third teeth acute; fifth minute. Gonopod 1 stout, with twisted apex carrying 2 groups of strong setae. Maximum cl. 7 mm. Subtidal (13–20 m). Tropical Eastern Pacific. 3 species (Guinot 1967).

Guinope Thoma & Felder, 2020 Diagnosis. Carapace trapezoidal, 1.5 times as wide as long; front 0.4 carapace width, with 2 lobes separated by shallow angle; frontexorbital width 0.8 times carapace width; anterolateral margin with exorbital tooth coalesced with obsolete first anterolateral tooth as denticulate margin; second anteriorly-directed, triangular, sharp; third triangular, smaller; fourth obsolete. Gonopod 1 slender, sinuous, with subdistal constriction and spatulate apex. Maximum cl. 4.5 mm. Shelf (58–143, possibly to 300 m). Tropical W Atlantic. 1 species (Thoma and Felder 2020).

Lophoxanthus A. Milne-Edwards, 1879 Diagnosis. Carapace trapezoidal, 1.4 times as wide as long; front 0.3 carapace width, almost straight; front-exorbital width 0.5

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times carapace width; anterolateral margin with exorbital tooth coalesced with obsolete first and second anterolateral teeth; third rounded-triangular; fourth acute, smaller, oblique. Maxilliped 3 merus distolateral angle produced. Pereopods 2–5 stout, longest about carapace width, almost laminiform, carpi with cristate upper margins. Maximum cl. 8.4 mm. Intertidal, subtidal. Tropical Eastern Pacific. 1 species (Rathbun 1930: rediagnosis, description).

Malacoplax Guinot, 1969 Diagnosis. Carapace subquadrate, 1.5 times as wide as long; front 0.25 carapace width, almost straight, with median notch; frontexorbital width 0.8 times carapace width; anterolateral margin with exorbital tooth coalesced with obtuse first anterolateral; second and third broad, short, elevated; fourth minute; convex. Gonopod 1 slender, with bilobed apex and subapical projection; with apical setae. Maximum cl. 16 mm. Intertidal. Tropical Eastern Pacific (California). 1 species (Guinot 1969a; Rathbun 1918: figure as Speocarcinus californiensis (Lockington, 1877)).

Micropanope Stimpson, 1871 Diagnosis. Carapace subhexagonal, 1.4–1.5 times as wide as long; front 0.4 carapace width, almost straight, with median notch; front-exorbital width 0.6 times carapace width; anterolateral margin with exorbital tooth separated from first anterolateral tooth by concave margin; second prominent, triangular, blunt; third acute, elevated laterally; fourth small. Gonopod 1 stout, with short apical and subapical flaps. Maximum cl. 7 mm. Subtidal–slope (10–355 m). Tropical W Atlantic, Tropical Eastern Pacific. 7 species (Guinot 1967: revision; Rathbun 1930: key to species, some now in other genera; Williams 1965: figures of type species as Micropanope sculptipes Stimpson, 1871).

Nanoplax Guinot, 1967 Diagnosis. Carapace suhexagonal, 1.4 times as wide as long; front 0.4 carapace width, with oblique lobes separated by median notch; front-exorbital width 0.7 times carapace width; anterolateral margin with exorbital coalesced with first anterolateral lobe; second and third triangular; fourth minute. Maxilliped 3 merus distolateral angle produced. Pereopods 2–5 elongate, longest ~1.5 times carapace width. Gonopod 1 twisted, flat, apex with flap folded proximally, with long subterminal spines. Maximum cl. 11 mm. Shelf, slope (9–333 m). Tropical W Atlantic. 2 species (Felder 2020: rediagnosis, second species; Guinot 1967: diagnosis).

Oediplax Rathbun, 1894 Diagnosis. Carapace subovate, 1.5 times as wide as long; front 0.25 carapace width, with 2 lobes separated by shallow angle; frontexorbital width 0.4 times carapace width; anterolateral margin with exorbital tooth separated from first anterolateral angle by straight transverse margin; second obtusely triangular; third and fourth similar, narrow, triangular. Maximum cl. 33 mm.

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Shelf (60 m). Tropical Eastern Pacific. 1 species (Guinot 1969b: discussion; Rathbun 1918: species description).

Panoplax Stimpson, 1871 Diagnosis. Carapace trapezoidal, 1.7 times as wide as long; front 0.4 carapace width, with 2 weak lobes; front-exorbital width 0.6 times carapace width; anterolateral margin with exorbital tooth separated from obtuse first anterolateral tooth by concavity; second quadrate with anterior angle; third acute; fourth minute. Gonopod 1 slender, curved, with apical curved lappet. Maximum cl. 9 mm (Pl. 64d). Subtidal, shelf (5–101 m). Tropical W Atlantic, Tropical Eastern Pacific (doubtful). 2 species (Guinot 1969a; Rathbun 1918: both discussed as Eucratoplax elata A. Milne-Edwards, 1890). Panoplax mundata Glassell, 1935 is possibly a member of Cyrtoplax.

Perunorhombila Števčić, 2005 Diagnosis. Carapace subquadrate, 1.2 times as wide as long; front 0.4 carapace width, with 2 lobes separated by shallow angle; frontexorbital width 0.65 times carapace width; anterolateral margin with small exorbital tooth, separated from first anterolateral tooth by straight margin; second and third spine-like, set one behind other, well separated. Maxilliped 3 merus distolateral angle produced. Pereopods 2–5 elongate, longest ~1.5 times carapace width. Gonopod 1 with subdistal long setae. Maximum cl. 10 mm. Slope (212–440 m). Tropical W Atlantic. 1 species (Chace 1940: species description; Števčić 2005: diagnosis).

Prionoplax H. Milne Edwards, 1852 Diagnosis. Carapace trapezoidal, 1.6 times as wide as long; front 0.3 carapace width, with 2 weak lobes; front-exorbital width 1 times carapace width; anterolateral margin with sharp exorbital angle; first tooth close-by, acute; third tooth shorter, acute. Maxilliped 3 merus distolateral angle produced. Pereopods 2–5 elongate, longest ~1.5 times carapace width. Gonopod 1 stout, with 1 lobular and 1 acute apical projection; 1 short subdistal projection; short subapical spines. Maximum cl. 16 mm. Subtidal, shelf (18–42 m). Tropical W Atlantic, Central IndoPacific, Tropical Eastern Pacific. 3 species (Guinot 1969a: rediagnosis; Rathbun 1918: figure of P. atlantica Kendall, 1891).

Pseudopanopeus Felder, 2020 Diagnosis. Carapace trapezoidal, 1.7 times as wide as long; front 0.5 carapace width, with 2 lobes separated by shallow angle; frontexorbital width 0.8 times carapace width; anterolateral margin with exorbital tooth indistinguishable from first anterolateral tooth; second triangular, separated from first by shallow depression; third triangular; fourth minute or obsolete. Gonopod 1 with subapical large, proximally directed spine (sometimes 2), with apical weakly curved finger. Maximum cl. 4 mm. Subtidal–slope (7–311 m). Tropical W Atlantic. 1 species (Felder 2020).

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Marine Decapod Crustacea

Pseudorhombila H. Milne Edwards, 1837

Tetraplax Rathbun, 1901

Diagnosis. Carapace subquadrate-trapezoidal, 1.5 times as wide as long; front 0.3 carapace width, almost straight, with median notch; front-exorbital width 0.7 times carapace width; anterolateral margin with exorbital tooth separated from obsolete first anterolateral tooth by shallowly concave margin; second blunt; third triangular, oblique; fourth minute if present. Maxilliped 3 merus distolateral angle produced. Gonopod 1 slender, apex with subterminal or lateral upturned flange, with patch of strong spines below flange. Maximum cl. 15 mm. Shelf (20–112 m). Tropical W Atlantic, Tropical Eastern Pacific. 5 species (Garth 1940: species description; Vazquez-Bader and Gracia 1995: key to 4 species).

Diagnosis. Carapace oval-trapezoidal, 1.2–1.6 times as wide as long; front 0.2–0.3 carapace width, with 2 weak lobes; front-exorbital width 0.9 times carapace width; anterolateral margin with prominent triangular exorbital tooth; 3 similar triangular teeth; longitudinal. Pereopods 2–5 elongate, longest ~1.5 times carapace width. Gonopod 1 stout, with 1 lobular and 1 acute apical projection; 1 short subdistal projection; short subapical spines. Maximum cl. 11 mm. Subtidal, shelf (4–46 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Guinot 1969a: rediagnosis; Rathbun 1918: figures; Türkay 1967: figures).

Robertsella Guinot, 1969 Diagnosis. Carapace ovate, 1.7 times as wide as long; front 0.3 carapace width, almost straight, with median notch; front-­exorbital width 0.5 times carapace width; anterolateral margin with exorbital tooth separated from first anterolateral tooth by concave margin; second prominent, triangular, blunt; third acute, elevated laterally; fourth small. Maxilliped 3 merus distolateral angle produced. Gonopod 1 slender, with broad proximolateral flange, apex with terminal recurved flap. Maximum cl. 22 mm. Shelf, slope (20–353 m). Temperate Northern and Tropical W Atlantic. 2 species (Gouvêa 2013: discussion; Guinot 1969b: diagnosis; Tavares and Gouvêa 2013: species description).

Scopolius Števčić, 2011 Diagnosis. Carapace subhexagonal, 1.3 times as wide as long; front 0.5 carapace width, almost straight, with median notch; frontexorbital width 0.8 times carapace width; anterolateral margin with exorbital tooth indistinguishable from first anterolateral tooth; second–fourth triangular, diminishing, one behind other; fifth absent. Maxilliped 3 merus distolateral angle produced. Gonopod 1 ‘distally bulbous, with recurved apical projection’. Maximum cl. 7 mm. Shelf, slope (150–480 m). Tropical W Atlantic. 1 species (Rathbun 1930: species description as Micropanope nuttingi (Rathbun, 1898); Števčić 2011: diagnosis).

Speocarcinus Stimpson, 1859 Diagnosis. Carapace ovate-subquadrate, dorsally elevated, 1.3–1.4 times as wide as long; front 0.3 carapace width, almost straight; front-exorbital width 0.7 times carapace width; anterolateral margin with exorbital tooth coalesced with first anterolateral tooth, together shorter than orbit width; second broad, blunt; third triangular; fourth small; all slightly upturned; convex. Gonopod 1 slender, with reflexed flattened apex, with short subapical spines. Maximum cl. 18 mm (Pl. 64e). Estuarine, subtidal, shelf (6–130 m). Tropical W and temperate SW Atlantic, Tropical Eastern Pacific. 8 species (Brandão et  al. 2012: remarks, key to species; Felder and Rabalais 1986: species descriptions; Guinot 1969b: 4 species).

Tetraxanthus Rathbun, 1898 Diagnosis. Carapace trapezoidal, 1.4 times as wide as long; front 0.3 carapace width, almost straight, with median notch; frontexorbital width 0.6 times carapace width; anterolateral margin with exorbital tooth coalesced with first anterolateral tooth, separated by straight margin; second and third obtusely triangular, blunt, well spaced; fourth and fifth absent. Maxilliped 3 merus distolateral angle produced. Maximum cl. 18 mm. Shelf, slope (20–622 m). Tropical W and temperate SW Atlantic. 2 species (Rathbun 1930: rediagnosis, key to species).

Thalassoplax Guinot, 1969 Diagnosis. Carapace subquadrate, 1.2 times as wide as long; front 0.4 carapace width, almost straight, with median notch; frontexorbital width 0.7 times carapace width; anterolateral margin with exorbital tooth coalesced with and indistinguishable from obtuse first anterolateral; second triangular, acute; third narrowly triangular, elevated, longer; fourth minute. Maxilliped 3 merus distolateral angle produced. Pereopods 2–5 elongate, longest ~1.5 times carapace width. Gonopod 1 stout, apex twisted, with subdistal row of short spines. Maximum cl. 10 mm. Slope (203–260 m). Tropical W Atlantic. 1 species (Guinot 1969b: diagnosis).

Trapezioplax Guinot, 1969 Diagnosis. Carapace trapezoidal, 1.6 times as wide as long; front 0.3 carapace width, almost straight; front-exorbital width 0.9 times carapace width; anterolateral margin with prominent acute exorbital tooth; first and second anterolateral teeth triangular, set one behind other. Maxilliped 3 merus distolateral angle produced. Pereopods 2–5 elongate, longest ~1.5 times carapace width. Gonopod 1 stout, almost straight, with tubular apex, with subapical short spines. Maximum cl. 18 mm. Subtidal, shelf (13–33 m). Tropical W Atlantic. 1 species (Guinot 1969b: figure of gonopod; figure of habitus on pl. 5 Fig. 3; Ng et  al. 2008: figure). Thoma et  al. (2014) placed Trapezioplax in Pseudorhombilidae. References Brandão MC, Coelho-Filho PA, Tavares M (2012) A review of the genus Speocarcinus Stimpson, 1859 (Crustacea: Brachyura: Xanthidae), with

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a key to its species and the description of one new species. Zootaxa 3327, 1–19. doi:10.11646/zootaxa.3327.1.1 Chace FA (1940) Reports on the scientific results of the Atlantis expeditions to the West Indies, under the joint auspices of the University of Havana and Harvard University, The brachyuran crabs. Torreia 1–67. Felder DL (2020) A new crab of the genus Nanoplax from the Gulf of Mexico, and assignment of Micropanope pusilla to a new genus (Crustacea, Brachyura, Pseudorhombilidae). Zootaxa 4810, 531–545. doi:10.11646/zootaxa.4810.3.9 Felder DL, Rabalais NN (1986) The genera Chasmocarcinus Rathbun and Speocarcinus Stimpson on the continental shelf of the Gulf of Mexico, with descriptions of two new species (Decapoda: Brachyura: Goneplacidae). Journal of Crustacean Biology 6, 547–575. doi:10.2307/1548194 Garth JS (1940) Some new species of brachyuran crabs from Mexico and the Central and South American mainland. Allan Hancock Pacific Expeditions 5, 53–95, pls 11–26. Gouvêa ACd (2013) Diferenciação morfológica entre os gêneros Euphrosynoplax Guinot, 1969; Robertsella Guinot, 1969; Thalassoplax Guinot, 1969; Panoplax Stimpson, 1871 e Eucratopsis Smitth, 1869, com a descrição de novas espécies. MS thesis. Universidade de São Paulo, Instituto de Biociências, São Paulo. Guinot D (1967) Recherches préliminaires sur les groupements naturels chez les Crustacés décapodes brachyoures. II. Les anciens genres Micropanope Stimpson et Medaeus Dana. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 39, 345–374. Guinot D (1969a) Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures. VII. Les Goneplacidae. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 41, 241–265, pl. 1. Guinot D (1969b) Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures. VII. Les Goneplacidae (suite et fin). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 41, 688–724, pls 1–5. Guinot D (1990) Éstablissement du genre Garthiope gen. nov., ses relations avec le genre Coralliope Guinot, 1967, et leurs affinités avec les Trapeziidae sensu lato (Crustacea Decapoda Brachyura). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 12, 469–487, pls I, II. Guinot D, Iliffe TM (1991) Garthiope anchialina sp. nov., espèce anchialine des Galapagos, île Isabella, Cueva de la Cadena, avec remarques sur la faune carcinologique anchialine (Crustacea Decapoda Brachyura). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 12, 607–62, pl. 1. Hendrickx ME (1989) Chacellus pacificus, new species (Crustacea, Decapoda, Brachyura, Goneplacidae), from the continental shelf of the Gulf of California, México. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 11, 193–202. Hendrickx ME (1998) A new genus and species of “goneplacid-like” brachyuran crab (Crustacea: Decapoda) from the Gulf of California, Mexico, and a proposal for the use of the family Pseudorhombilidae Alcock, 1900. Proceedings of the Biological Society of Washington 111, 634–644. Lai JC, Mendoza JCE, Guinot D, Clark PF, Ng PKL (2011) Xanthidae MacLeay, 1838 (Decapoda: Brachyura: Xanthoidea) systematics: A multi-gene approach with support from adult and zoeal morphology. Zoologischer Anzeiger 250, 407–448. doi:10.1016/j.jcz.2011.07.002 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Rathbun MJ (1918) The grapsoid crabs of America. United States National Museum Bulletin 97, 1–461, pls 1–161. Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230.

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Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159. Števčić Z (2011) Addition to the reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Part I. New taxa. Natura Croatica 20, 125–139. Tavares M, Gouvêa A (2013) A new species of Robertsella Guinot, 1969 (Crustacea: Brachyura: Panopeidae) from the southwestern Atlantic. Zootaxa 3734, 72–80. doi:10.11646/zootaxa.3734.1.8 Thoma BP, Felder DL (2020) A new genus and species of xanthoid crab (Decapoda: Brachyura) from offshore hard bank habitats in the Gulf of Mexico. Zootaxa 4731, 403–413. doi:10.11646/zootaxa.4731.3.8 Thoma BP, Schubart CD, Felder DL (2009) Molecular phylogeny of Western Atlantic representatives of the genus Hexapanopeus (Decapoda: Brachyura: Panopeidae). In Crustacean Issues Vol. 18: Decapod Crustacean Phylogenetics. (Eds Martin JW, Crandall KA, Felder DL) pp. 551–565. CRC Press, Boca Raton. Thoma BP, Guinot D, Felder DL (2014) Evolutionary relationships among American mud crabs (Crustacea: Decapoda: Brachyura: Xanthoidea) inferred from nuclear and mitochondrial markers, with comments on adult morphology. Zoological Journal of the Linnean Society 170, 86–109. doi:10.1111/zoj.12093 Türkay M (1967) Neue Brachyuren von der Westküste Südamerikas (Crustacea, Decapoda). Senckenbergiana Biologica 48, 361–364. Vázquez-Bader AR, Gracia A (1991) Euphrosynoplax campechiensis, new species (Crustacea, Decapoda, Brachyura, Goneplacidae), from the continental shelf of Southwestern Gulf of Mexico. Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 13, 433–438. Vazquez-Bader AR, Gracia A (1995) A new crab species of the genus Pseudorhombila H. Milne-Edwards, 1837 (Crustacea: Decapoda: Goneplacidae). Proceedings of the Biological Society of Washington 108, 254–265. Williams AB (1965) Marine decapod crustaceans of the Carolinas. Fishery Bulletin 65, 1–298.

Xanthidae MacLeay, 1838 rock crabs Figures 14.149–14.168, Plates 64f–r, 65–67 The xanthids are widely considered to be among the most difficult crabs to identify because of their high speciesrichness, intergrading morphology and diversity of carapace and pereopodal ornamentation. Many of the diagnostic characters are subtle or qualitative, especially with regards to carapace ornamentation, making identification often challenging for the novice and expert alike. Xanthids are currently united by the comparatively elongated thoracic sternites 3–4 (being as long as or longer than half the width), and relatively simple gonopod 1 that lacks convoluted apical folds and complex protrusions. Xanthids can be among the most common crabs of tropical shallow water rocky and coral reefs worldwide, and many extend into temperate waters, as well as outer shelf or slope habitats. They are generally slow moving, usually have dark finger tips on the chelipeds, and are often found under boulders and within tight crevices. Most of the more than 130 genera and 630 species are from the Indo-West Pacific but

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many have become established in the Mediterranean Sea as Lessepsian migrants from the Red Sea or as introductions via shipping from the western Atlantic (Galil et al. 2002; Mavidis et al. 2008). Several crabs are poisonous to humans if consumed, the majority being xanthids (Llewellyn et  al. 2002; Tsai et al. 2006; Yasumoto et al. 1983). The classification of Xanthidae has changed considerably in recent decades, and several groups formerly included are now considered separate families (many even now in other superfamilies). Guinot (1967a, 1967b, 1967c, 1968, 1969, 1976, 1978, 1979) commenced substantial revision of the xanthids that continues to this day but Serène’s (1984) major synopsis of the family, with keys to the subfamilies, most genera and most species from the Indo-West Pacific, remains

the single most comprehensive taxonomic work on the family. Xanthidae currently includes 14 subfamilies (Davie et al. 2015; Ng et al. 2008), but recent studies based on molecular data and larval morphology question the monophyly of most of these, and possibly even the family itself (Lai et al. 2011; Thoma et al. 2014). The xanthid system remains a work in progress. Here, we substantially follow the classification of Ng et al. (2008) and De Grave et al. (2009) with minor modifications as outlined under each subfamily entry. Diagnosis. Thoracic sternites 3–4 half or more times as long as wide. Male thoracic sternite 8 usually fully concealed by pleonite 2. Gonopod 1 slender, apex relatively simple, apex without convoluted folds or complex protrusions. Male gonopore coxal.

Key to subfamilies of Xanthidae 1. – 2. – 3.

– 4. – 5. – 6.

– 7.



Pereopods 2–5 short, laterally compressed, especially distal articles; dactyli blade-like (Fig. 14.161a, c, g, h)�������������������������������������������������������������������������������������������������������������������������������������������������Kraussiinae Pereopods 2–5 not broadly flattened, cylindrical or carinate at most�������������������������������������������������������������������������������������2 Chelipeds equal, similar, slender; fingers long, thin, cutting edges armed with slender upright teeth for holding sea anemones (Fig. 14.163e–g)���������������������������������������������������������������������������������������������������������������Polydectinae Chelae with fingers variously armed, not especially slender or lined with pointed teeth����������������������������������������������������3 Cheliped finger tips rounded, expanded, hollowed on occlusal surface, spoon-shaped, meeting when closed (Fig. 14.155d) AND pereopods 2–5 with dactylo-propodal ‘lock’ (Fig. 14.153s–u) formed by rounded prolongation of propodal lateral margin sliding against and beneath projecting button situated proximally on lateral margin of dactylus (if not clearly differentiated then dactylus apically bifid)����������������������������4 Cheliped finger tips acute or expanded, spoon-shaped. Pereopods 2–5 usually without dactylo-propodal ‘lock’; if dactylo-propodal ‘lock’ present, cheliped fingers tips not expanded, spoon-shaped���������������������������������������5 Front wider than one-third maximum carapace width, typically with 2 large submedian lobes and 2 small lateral lobules, or nearly straight, without lateral sinuses; anterolateral margin typically with 4 teeth behind exorbital angle (sometimes 2 or 3) (Fig. 14.153). Antennules folding transversely�������������������Chlorodiellinae Front narrower than one-third maximum carapace width, 2- or 4-lobed, with median fissure, projecting between lateral sinuses demarcating supraorbital angles; anterolateral margin with 4–8 teeth behind exorbital angles (Fig. 14.155). Antennules folding obliquely forwards������������������������������������������������������������������ Etisinae Antennules folding obliquely forwards�����������������������������������������������������������������������������������������������������������������������������������������6 Antennules folding transversely or almost so������������������������������������������������������������������������������������������������������������������������������7 Carapace subcircular or longitudinally subovate, slightly longer to scarcely wider than long, flat, regions usually indistinct, lateral margins weakly dentate, sparsely to densely setose. Chelipeds strongly unequal, dissimilar in both sexes; fingers curved conical, tips rounded, hollowed, meeting. Symbionts of scleractinian corals (Fig. 14.154)������������������������������������������������������������������������������������������������������������������������������ Cymoinae Carapace subcircular to transversely subovate, wider than long, convex, regions distinct, lateral margins weakly to strongly dentate, densely covered with short or long setae. Chelipeds subequal, similar in both sexes; fingers laterally compressed, thin, tips pointed, usually crossing. Free living (Fig. 14.152)������������ Banareiinae Carapace posterolateral margin cristate, interrupted by posterior end of sub-branchial channel curving onto posterior branchial region; cardiac region swollen, sometimes overhanging deep cardio-intestinal groove (Fig. 14.151a–c). Cheliped fingers pigmented along cutting edge and on no more than distal half (Fig. 14.151g, h)������������������������������������������������������������������������������������������������������������������������������������������������ Antrocarcininae Carapace posterolateral margin not cristate, not interrupted by posterior end of sub-branchial channel; cardiac region not swollen. Cheliped fingers pigmented over most of length������������������������������������������������������������������8

14 – Brachyura – crabs

751

8.

Carapace anterolateral margin descending anteriorly, not meeting exorbital angle but continuing below orbit towards anterolateral corner of buccal frame���������������������������������������������������������������������������������������������������������������9 – Carapace anterolateral margin horizontal, meeting exorbital angle�������������������������������������������������������������������������������������10 9. Carapace front distally downcurved, appearing 4-lobed in anterior view; subhepatic surface with, more-or less vertical grooves descending from gaps between anterolateral teeth (Fig. 14.160)����������� Glyptoxanthinae – Carapace front horizontal, appearing subtly 2-lobed, almost straight in anterior view; subhepatic surface without vertical grooves descending from gaps between anterolateral teeth (Figs 14.156–14.158)�����������������������������������������������������������������������������������������������������������������������������������������������Euxanthinae 10. With all or combination of the following: 1) cheliped with thin, inwardly inclined crest on upper margin of palm, fitting closely against carapace when folded; 2) pereopods 2–5 meri, carpi and propodi upper margins with thin crest; 3) carapace anterolateral margin with thin crested crimped rim or 4 broad, flat rimmed lobes or teeth behind exorbital angle, or margins rounded, only subtly interrupted (Fig. 14.168)�����������������������������������������������������������������������������������������������������������������������������������������������������������������Zosiminae – Cheliped upper inner margin without thin crest. Pereopods 2–5 usually ovate in section, upper margin smooth, spinous or granular, rarely crested. Carapace anterolateral margin not crested, typically with 4 or 5 teeth or broad lobes, sometimes scarcely indicated or subdivided into as many as 10 teeth�����������������������������11 11. Male thoracic sternites 3–4 length equal to or slightly less than half maximum width (measured across lateral extremities of episternites) (Fig. 14.159b)����������������������������������������������������������������������������������������������������������������� 12 – Male thoracic sternites 3–4 length greater than half maximum width���������������������������������������������������������������������������������13 12. Carapace up to twice as wide as long; anterolateral teeth absent or as rounded convex lobes. Cheliped finger tips hollowed on occlusal margin, not crossing. Pereopods 2–5 merus upper margins smooth or granular. Gonopods and female gonopores always bilaterally symmetrical (Fig. 14.162)����������������������� Liomerinae – Carapace up to 1.7 times as wide as long; anterolateral teeth absent or as sharp, curved spines. Cheliped finger tips acute, not hollowed, crossing. Pereopods 2–5 meri upper margins smooth or spinose. Gonopods and female gonopores bilaterally symmetrical (Garthiella) or with male gonopods longer on right side and female gonopore larger on left side (Mabui) (Fig. 14.159)������������������������������������������������Garthiellinae 13. Carapace regions weakly or well defined, lobulate, surface occasionally smooth but usually densely granular, tuberculate or spinose, with short bristly setae and longer clavate setae; when present, surface granulation or spination distributed over whole surface. Cheliped and pereopods 2–5 densely granular or spinose, not smooth (Figs 14.149, 14.150)����������������������������������������������������������������������������������������������������������� Actaeinae – Carapace regions not or well defined, usually smooth or minimally granular, glabrous or with scattered setae, occasionally densely granulate, finely, evenly setose; when present, surface granulation usually confined largely to anterior half, rarely over entire surface. Cheliped and pereopods 2–5 usually smooth or weakly granular, sometimes densely tuberculate or spinous (Figs 14.164–14.167)�������������������������������������Xanthinae

Subfamily Actaeinae Alcock, 1898 Actaeine crabs tend to be more highly ornamented, and more setose than xanthines, which share a more elongate thoracic sternum than other xanthids. They are usually brightly coloured, often patterned with red. They are generally slow moving and most have compact walking legs. Actaeines tend to inhabit intertidal and shallow reef environments in rubble, under boulders and in deep crevices. Serène (1984) provided a key to 14 genera from the Indo-West Pacific. Psaumis and Pseudactaea, previously considered to be actaeines, were shown to belong to Epixanthinae and Banareinae, respectively (Lai et  al. 2011). Here we recognise 18 actaeine genera, most of which occur in the Indo-West Pacific; one is endemic to the Atlantic.

Diagnosis. Carapace regions weakly or well defined, lobulate, surface occasionally smooth but usually densely granular, tuberculate or spinose, with short bristly setae and longer clavate setae; when present, surface granulation or spination distributed over whole surface. Male thoracic sternites 3–4 length greater than half maximum width. Antennules folding transversely. Cheliped and pereopods 2–5 densely granular or spinose; chelipeds with finger tips acute or expanded, spoon-shaped; pereopods 2–5 usually without dactylo-propodal ‘lock’. Implicit generic characters. Carapace posterolateral margin nearly straight. Cheliped palms not noticeably swollen; fingers relatively elongate, straight to weakly curved, without paired brushes of rigid setae.

752

Marine Decapod Crustacea

Key to genera of Actaeinae 1.

Cheliped palms very swollen, fingers short, dactylus strongly curved; each finger with prominent brush of rigid setae on either side of central tooth of cutting margin (Fig. 14.150e, h) (brushes sometimes reduced in P. speciosa, but uniquely with distally directed brush of setae completely encircling tip of pereopod 2 dactylus) (Fig. 14.150i, j)�������������������������������������������������������������������������������������������������������������������������������������Pseudoliomera – Cheliped palms not noticeably swollen; fingers relatively elongate, straight to weakly curved, without paired brushes of rigid setae������������������������������������������������������������������������������������������������������������������������������������������������������2 2. Carapace posterolateral margin nearly straight; if concave pereopods 2–5 spinose but without carinae�������������������������3 – Carapace posterolateral margin forming deep depression to accommodate posterior pereopods when folded, granular or carinate, but never spinose or nodular (Fig. 14.149c, f)��������������������������������������������������������������������17 3. Pereopods 2–5 laterally compressed, carinate, upper margin cristate (Fig. 14.150g)�����������������������������������������������Serenius – Pereopods 2–5 smooth, granular, nodular or spinose, not carinate or cristate����������������������������������������������������������������������4 4. Pereopods 2–5 slender, pereopod 5 merus ~3 times as long as wide or more�������������������������������������������������������������������������5 – Pereopods 2–5 stout, pereopod 5 merus less than 2.5 times as long as wide��������������������������������������������������������������������������7 5. Carapace regions covered in small, well spaced, rounded granules (Fig. 14.150f)��������������������������������������������������������� Rata – Carapace regions covered in crowded, bulging lobules or granules, or irregular, rugose, eroded ‘slabs’��������������������������6 6. Carapace surface glabrous; regions covered in bulging lobules or granules, or irregular, rugose, eroded ‘slabs’ (Fig. 14.149k). Male thoracic sternites 3, 4 partially demarcated by short groove laterally (Fig. 14.150n)��������������������������������������������������������������������������������������������������������������������������������������������������������������� Meractaea – Carapace surface sparsely setose; regions covered in crowded, bulging granules (Fig. 14.150a). Male thoracic sternites 3, 4 demarcated by wide central transverse groove (Fig. 14.150p)�������������������������������������������Odhnea 7. Male thoracic sternites 3, 4 demarcated only at lateral margins of sternal plastron (Fig. 14.150l, o)���������������������������������8 – Male thoracic sternites 3, 4 demarcated by complete transverse groove (Fig. 14.150m, q)������������������������������������������������13 8. Carapace, chelipeds and pereopods 2–5 covered with evenly spaced granules, covered with short velvet pile or surrounded by short, rigid black or brown setae. Carapace regions weakly projecting, separated by shallow, broad furrows���������������������������������������������������������������������������������������������������������������������������������������������������������������9 – Carapace, chelipeds and usually pereopods 2–5 densely covered with spines, crowded nodules or compound granules, glabrous or with symmetrically placed tufts of long setae. Carapace regions projecting, separated by distinct furrows; furrows wide or indicated only by fissure���������������������������������������������������11 9. Male thoracic sternite 4 with median groove concealed by telson, not evident on exposed surface anterior to telson (Fig. 14.150o). Pereopod 5 merus less than twice as long as wide (Fig. 14.149l)������������������������������� Novactaea – Male thoracic sternite 4 with median groove in front of telson, fully visible when pleon closed (Fig. 14.150l). Pereopod 5 merus 2–2.5 times as long as wide���������������������������������������������������������������������������������������������������������������������10 10. Carapace length: width 1.5–1.6; anterolateral divisions absent or scarcely indicated, weakly convex (Fig. 14.150d)������������������������������������������������������������������������������������������������������������������������������������������������������������� Platyactaea – Carapace length: width 1.3–1.5 (usually 1.3); anterolateral divisions distinct, bluntly rounded to angular or dentiform (Fig. 14.149g)�������������������������������������������������������������������������������������������������������������������������������������������� Forestiana 11. Carapace glabrous, regions separated by narrow grooves, with mosaic-like, squamiform or nodular tubercles (Fig. 14.149a, b)�������������������������������������������������������������������������������������������������������������������������������������������������Actaea – Carapace setose, regions usually separated by wide furrows, with rounded, simple or compound tubercles��������������� 12 12. Carapace regions marked by large, smooth, lobular glabrous tubercles, setose between tubercles. Pereopods 2–5 meri, carpi and propodi upper margin spinose (Fig. 14.149d)���������������������������������������������������������������������� Allactaea – Carapace regions marked by compound tubercles and crowded granules, variously setose. Pereopods 2–5 meri, carpi and propodi upper margin nodulose (Fig. 14.149e)�������������������������������������������������������������������������� Epiactaea 13. Carapace anterolateral teeth sharp curved spines of differing lengths. Pereopod 2–5 meri and propodi upper margins lined with spines (Fig. 14.149i)��������������������������������������������������������������������������������������������������� Heteractaea – Carapace anterolateral teeth, if indicated, blunt angular or nodular lobes. Pereopod 2–5 meri and propodi upper margins unarmed or tubercular but not spinose������������������������������������������������������������������������������������������������������14

14 – Brachyura – crabs

753

Fig. 14.149.  Xanthidae, Actaeinae. a, Actaea calculosa (H. Milne-Edwards, 1834); b, Actaea peronii (H. Milne-Edwards, 1834); c, Actaeodes hirsutissimus (Rüppell, 1830); d, Allactaea lithostrota Williams, 1974; e, Epiactaea margaritifera (Odhner, 1925); f, Epiactaeodes tessellatus (Pocock, 1890); g, Forestiana lucius Ng, 2014; h, Gaillardiellus orientalis (Odhner, 1925); i, Heteractaea ceratopus (Stimpson, 1860); j, Lobiactaea lobipes (Odhner, 1925); k, Meractaea takunan Komatsu & Takeda, 2011; l, Novactaea bella Guinot, 1976.

14. –

Carapace regions distinct but not grossly swollen, surfaces granulate, granules numerous but spaced as to not be appressed, usually with short and long stiff setae; regions separated by shallow, largely open furrows; region 3M undivided������������������������������������������������������������������������������������������������������������������������������������������������15 Carapace regions prominently swollen, bulging, densely covered with crowded granules or large rounded nodules, usually with fine downy setae; regions separated by more or less broad but deep furrows, with

754

15. – 16. – 17. –

Marine Decapod Crustacea

or without fine tomentum; region 3M divided into three parts (slender anterior portion and two adjacent posterior portions)����������������������������������������������������������������������������������������������������������������������������������������������������16 Pereopods 2–5 carpi and propodi with prominent wart-like and pointed tubercles; meri upper margins with subdistal tooth (Fig. 14.149j)�������������������������������������������������������������������������������������������������������������������������� Lobiactaea Pereopods 2–5 granulation uniform, similar to that of carapace, never nodular; meri upper margins without subdistal tooth (Fig. 14.149h)���������������������������������������������������������������������������������������������������������������Gaillardiellus Cheliped finger tips blunt, slightly hollowed, particularly that of fixed finger, tips not crossing when closed (Fig. 14.150b)���������������������������������������������������������������������������������������������������������������������������������������������������������������� Paractaea Cheliped finger tips pointed, not hollowed, crossing when closed (Fig. 14.150c)��������������������������������������������Paractaeopsis Cheliped fingers with cutting edges dentate along full length, tips rounded, slightly spooned or pointed. Carapace regions 3M and 4M subdivisions close together, not forming fleur-de-lys pattern (Fig. 14.149c)����������������������������������������������������������������������������������������������������������������������������������������������������������������� Actaeodes Cheliped fingers with cutting edges thin, with 2 or 3 low submedian (fixed finger) or proximal (dactylus) teeth but otherwise blade-like, tips pointed. Carapace regions 3M and 4M subdivisions forming fleur-de-lys pattern (Fig. 14.149f)������������������������������������������������������������������������������������������������������������������������Epiactaeodes

Fig. 14.150.  Xanthidae, Actaeinae. a, Odhnea echinus (Alcock, 1898); b, Paractaea rufopunctata (H. Milne Edwards, 1834); c, Paractaeopsis quadriareolata (Takeda & Miyake, 1968); d, Platyactaea setigera (H. Milne Edwards, 1834); e, Pseudoliomera violacea (A. Milne-Edwards, 1873); f, Rata chalcal Davie, 1997; g, Serenius kuekenthali (De Man, 1902). h, chela, Pseudoliomera. i, j, pereopod 2, Pseudoliomera. Male thoracic sternites 1–4, pleonite 6, telson: k, Actaea; l, Forestiana; m, Gaillardiellus; n, Meractaea; o, Novactaea; p, Odhnea; q, Paractaea.

14 – Brachyura – crabs

Actaea De Haan, 1833 Diagnosis. Carapace, chelipeds and pereopods 2–5 densely covered with spines, crowded nodules or compound granules; carapace glabrous, regions prominent, separated by narrow grooves, with mosaic-like tubercles, squamiform or nodular tubercles. Male thoracic sternites 3, 4 demarcated only at lateral margins of sternal plastron. Maximum cl. 23 mm (Pl. 64f). Intertidal–slope (0–300 m). Tropical W Atlantic, Indo-West Pacific, Temperate Southern Africa, Temperate Australia. 31 species (Guinot 1969: rediagnosis; Rathbun 1930: key to 8 American species; Serène 1984: key to 18 Indo-West Pacific species).

Actaeodes Dana, 1851 Diagnosis. Carapace posterolateral margin forming deep depression to accommodate posterior pereopods when folded, granular or carinate, but never spinose or nodular. Male thoracic sternite 3–4 demarcated by complete transverse groove. Cheliped fingers with cutting edges dentate along full length, tips rounded, slightly spooned or pointed. Maximum cl. 20 mm. Intertidal, subtidal (0–30 m). Indo-West Pacific, Temperate Australia. 6 species (Guinot 1976: rediagnosis; Serène 1984: key to species).

Allactaea Williams, 1974 Diagnosis. Carapace regions marked by large, smooth, lobular glabrous, tubercles, regions separated by wide furrows, setose between tubercles. Male thoracic sternites 3, 4 demarcated only at lateral margins of sternal plastron. Pereopods 2–5 stout, pereopod 5 merus less than 2.5 times as long as wide; meri, carpi and propodi upper margins spinose. Maximum cl. 18 mm. Shelf, slope (50–640 m). Temperate Northern and Tropical W Atlantic. 1 species (Soto 1980; Williams 1974).

Epiactaea Serène, 1984 Diagnosis. Carapace regions marked by compound tubercles and crowded granules, regions separated by wide furrows, variously setose. Male thoracic sternites 3, 4 demarcated only at lateral margins of sternal plastron. Pereopods 2–5 stout, pereopod 5 merus less than 2.5 times as long as wide; meri, carpi and propodi upper margins nodulose. Maximum cl. 18 mm (Pl. 64g). Intertidal–shelf (0–96 m). Indo-West Pacific. 3 species (Serène 1984: key to 2 species).

Epiactaeodes Serène, 1984 Diagnosis. Carapace regions 3M and 4M subdivided to form fleur-de-lys pattern; posterolateral margin forming deep depression to accommodate posterior pereopods when folded, granular or carinate, but never spinose or nodular. Male thoracic sternite 3–4 demarcated by complete transverse groove. Cheliped fingers with cutting edges thin, with 2 or 3 low submedian (fixed finger)

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or proximal (dactylus) teeth but otherwise blade-like, tips pointed. Maximum cl. 14 mm. Intertidal, subtidal (0–14 m). Western and Central IndoPacific. 2 species (Serène 1984).

Forestiana Guinot & Low, 2010 Diagnosis. Carapace length: width 1.3–1.5 (usually 1.3); carapace, chelipeds and pereopods 2–5 covered with evenly spaced granules, surrounded by rigid black or brown setae; regions weakly projecting, separated by shallow, broad furrows; anterolateral margin with distinct, bluntly rounded to angular or dentiform divisions. Male thoracic sternite 4 posterior part with median groove in front of telson, fully visible when pleon closed. Pereopods 2–5 stout, pereopod 5 merus 2–2.5 times as long as wide. Maximum cl. 33 mm. Intertidal–shelf (0 to at least 200 m). Western and Central Indo-Pacific. 5 species (Ng 2014: species compared; Serène 1984: key to 3 species as Forestia, preoccupied name).

Gaillardiellus Guinot, 1976 Diagnosis. Carapace regions distinct but not grossly swollen, surfaces granulate, granules numerous but spaced as to not be appressed, usually with short and long stiff setae; regions separated by shallow, largely open furrows; region 3M undivided. Male thoracic sternite 3–4 demarcated by complete transverse groove. Pereopods 2–5 granulation uniform, similar to that of carapace, never nodular; meri upper margins without subdistal tooth. Maximum cl. 24 mm (Pl. 64h). Intertidal–slope (0–312 m). Indo-West Pacific, Temperate Australasia (Kermadec Islands), Temperate Southern Africa. 6 species (Davie 1997: species description; Serène 1984: key to 3 species; Takeda and Komatsu 2010: species description).

Heteractaea Lockington, 1877 Diagnosis. Carapace regions distinct, coarsely granular, densely setose; anterolateral margin with sharp curved spines of differing lengths. Male thoracic sternites 3–4 demarcated by complete transverse groove. Pereopods 2–5 meri and propodi upper margins lined with spines. Maximum cl. 21 mm (Pl. 64i). Intertidal–shelf (0–130 m). Tropical W Atlantic, Tropical Eastern Pacific. 3 species (Garth 1940: third species; Rathbun 1930: key to 2 species).

Lobiactaea Sakai, 1983 Diagnosis. Carapace regions distinct but not grossly swollen, surfaces granulate, granules numerous but spaced as to not be appressed, with short and long stiff setae; regions separated by shallow, largely open furrows; region 3M undivided. Male thoracic sternites 3–4 demarcated by complete transverse groove. Pereopods 2–5 carpi and propodi with prominent wart-like and pointed tubercles; meri upper margins with subdistal tooth. Maximum cl. 10 mm.

756

Marine Decapod Crustacea

Shelf (40–87 m). Temperate Northern Pacific (South China Sea, Japan). 1 species (Odhner 1925: figure of type species as Actaea lobipes Odhner1925; Sakai 1983).

Meractaea Serène, 1984 Diagnosis. Carapace surface glabrous; regions covered in bulging lobules or granules, or irregular, rugose, eroded ‘slabs’. Male thoracic sternite 3–4 partially demarcated by short groove laterally. Pereopods 2–5 slender, pereopod 5 merus ~3 times as long as wide or more. Maximum cl. 21 mm. Intertidal–slope (0–285 m). Temperate Northern W Pacific, Indo-West Pacific. 4 species (Komatsu and Takeda 2011: key to species).

Paractaeopsis Serène, 1984 Diagnosis. Carapace regions prominently swollen, bulging, covered with large rounded nodules and scattered long setae; regions separated by more or less broad but deep, glabrous furrows; region 3M divided into three parts (slender anterior portion and two adjacent posterior portions). Male thoracic sternites 3–4 demarcated by complete transverse groove. Cheliped fingers with pointed tips, not hollowed, crossing when closed. Pereopods 2–5 stout, pereopod 5 merus less than 2.5 times as long as wide. Maximum cl. 7.4 mm. Intertidal, subtidal (0–5 m). Western and Central Indo-Pacific. 1 species (Serène 1984; Takeda and Miyake 1968: description of only species as Actaea quadriareolata Takeda & Miyake, 1968).

Novactaea Guinot, 1976

Platyactaea Guinot, 1967

Diagnosis. Carapace, chelipeds and pereopods 2–5 covered with evenly spaced granules, surrounded by short, rigid black setae; carapace regions weakly projecting, separated by shallow, broad furrows. Male thoracic sternite 4 posterior part with median groove concealed by telson, not evident on exposed surface anterior to telson. Male thoracic sternites 3–4 partially demarcated by short groove laterally. Pereopods 2–5 stout, pereopod 5 merus less than twice as long as wide. Maximum cl. 18 mm. Intertidal–shelf (0–157 m). Indo-West Pacific, Temperate Australasia (Kermadec Islands). 4 species (Guinot 1976: diagnosis; Serène 1984: key to species).

Diagnosis. Carapace length: width 1.5–1.6; carapace, chelipeds and pereopods 2–5 covered with evenly spaced granules, surrounded by short, rigid black setae; regions weakly projecting, separated by shallow, broad furrows; anterolateral divisions absent or scarcely indicated, weakly convex. Male thoracic sternites 3, 4 demarcated only at lateral margins of sternal plastron; thoracic sternite 4 posterior part with median groove in front of telson, fully visible when pleon closed. Pereopods 2–5 stout, pereopod 5 merus 2–2.5 times as long as wide. Maximum cl. 17 mm (Pl. 64l). Intertidal–shelf (0–65 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Guinot 1976; Rathbun 1930: figures as Actaea setigera A Milne Edwards, 1865).

Odhnea Ng & Low, 2010 Diagnosis. Carapace surface sparsely setose; regions covered in crowded, bulging granules. Male thoracic sternites 3–4 demarcated by wide transverse groove. Pereopods 2–5 slender, pereopod 5 merus ~3 times as long as wide or more. Maximum cl. 23 mm (Pl. 64j). Subtidal–shelf (15–52 m). Temperate Northern W Pacific, Western Indo-Pacific. 2 species (Mendoza et  al. 2012: figure). Replacement name for Odhneria Sakai, 1983.

Paractaea Guinot, 1969 Diagnosis. Carapace regions prominently swollen, bulging, densely covered with crowded granules and usually with fine downy setae; regions separated by more or less broad but deep furrows lined with fine tomentum; region 3M divided into three parts (slender anterior portion and two adjacent posterior portions). Male thoracic sternites 3–4 demarcated by complete transverse groove. Cheliped fingers with blunt tips, slightly hollowed, particularly that of fixed finger, tips not crossing when closed. Maximum cl. 24 mm (Pl. 64k). Intertidal–shelf (0–200 m). Temperate Northern and Tropical Atlantic, including Mediterranean, Indo-West Pacific. 16 species (Deb 1989: rediagnosis, key to 8 species; Guinot 1969: key to 18 species and forms; Serène 1984: key to 13 species and subspecies).

Pseudoliomera Odhner, 1925 Diagnosis. Cheliped palms very swollen, fingers short, dactylus strongly curved; each finger with prominent brush of rigid setae on either side of central tooth of cutting margin (brushes sometimes reduced in P. speciosa, but uniquely with distally directed brush of setae completely encircling tip of pereopod 2 dactylus. Maximum cl. 23 mm (Pl. 64m). Intertidal–shelf (0–31 m). Western and Central Indo-Pacific. 10 species (Serène 1984: key to 8 species).

Rata Davie, 1993 Diagnosis. Carapace regions covered in small, well spaced, rounded granules; posterolateral margin nearly straight. Pereopods 2–5 slender, dorsally unarmed, non-cristate; pereopod 5 merus ~3 times as long as wide or more. Maximum cl. 14.5 mm. Shelf–slope (170–270 m). Central and Eastern Indo-Pacific. 2 species (Davie 1993, 1997).

Serenius Guinot, 1976 Diagnosis. Carapace dorsal surface covered with flat, rounded lobules or flat compound tubercles, usually margined with stiff setae; posterolateral margin nearly straight. Pereopods 2–5

14 – Brachyura – crabs

laterally compressed, carinate, upper margin cristate. Maximum cl. 11 mm. Intertidal, subtidal (0–10 m). Western and Central IndoPacific. 6 species (Guinot 1976).

Subfamily Antrocarcininae Ng & Chia, 1994 Antrocarcinines are highly sculptured crabs from rocky environments at shelf and upper slope depths. They are recognised by the cristate posterolateral margin that is interrupted by the posterior end of sub-branchial channel curving on to posterior branchial region. The fingers of the chelipeds are pigmented only at tip and along cutting edge

757

(Fig. 141.151g, h) unlike most other xanthoids in which the fingers are pigmented to their bases. The subfamily was revised in detail by Ng and Chia (1994) who diagnosed all genera and species. Naruse et al. (2020) refined the subfamily diagnosis. Diagnosis. Carapace posterolateral margin cristate, interrupted by posterior end of sub-branchial channel curving onto posterior branchial region; cardiac region swollen, sometimes overhanging deep cardio-intestinal groove. Antennules folding transversely. Chelipeds with fingers pigmented along cutting edge and on no more than distal half; pereopods 2–5 without dactylo-propodal ‘lock’.

Key to genera of Antrocarcininae 1.



Carapace front with broad shallow notch, frontal lobes usually inclined obliquely outwards; anterolateral margin not lamelliform (Fig. 14.151a). Cheliped palm upper margin with inward-folding crest, median part with spines or teeth (Fig. 14.151g). Pleonites 2–4 with 3 transverse, granulate swellings, pleonites 5 and 6 with large median granulate swelling (Fig. 14.151e)������������������������������������������������������������������������������Antrocarcinus Carapace front with narrow fissure, frontal lobes directed anteriorly; anterolateral margin lamelliform (Fig. 14.151b–d). Cheliped palm upper margin not cristate, median part without spines or teeth (Fig. 14.151i, j). Pleonites 2–5 without swellings��������������������������������������������������������������������������������������������������������������������2

Fig. 14.151.  Xanthidae, Antrocarcininae. a, Antrocarcinus petrosus Ng & Chia, 1994; b, Cyrtocarcinus truncatus (Rathbun, 1906); c, d, Glyptocarcinus lophopus Takeda, 1973. e, male pleon, Antrocarcinus. f, thoracic sternites 1–4, Cyrtocarcinus. Chela, upper, lateral views: g, h, Antrocarcinus; i, j, Cyrtocarcinus.

758

2.



Marine Decapod Crustacea

Carapace anterolateral teeth low, broadly triangular, lobiform (Fig. 14.151b). Cheliped palm inner surface swollen, surfaces rounded; carpus with 1 broad, blunt tooth at inner distal angle (Fig. 14.151i). Thoracic sternite 4 posterior part sloping steeply, forming strong transverse ridge (Fig. 14.151f)����������������������������������������������������������������������������������������������������������������������������������������������������������Cyrtocarcinus Carapace anterolateral teeth well developed, acutely triangular (Fig. 14.151c). Cheliped palm inner surface with longitudinal ridge; carpus with 1 large sharp tooth at inner distal angle, 1 sub-basal tooth (Fig. 14.151j). Thoracic sternite 4 posterior part sloping gradually, forming low transverse ridge��������� Glyptocarcinus Antrocarcinus Ng & Chia, 1994

Diagnosis. Carapace front with broad shallow notch, frontal lobes usually inclined obliquely outwards. Cardio-intestinal region shallow, cardiac fold not over-reaching groove; anterolateral margin not lamelliform. Thoracic sternite 4 posterior part sloping gradually forming low transverse ridge. Pleonites 2–4 with 3 transverse, granulate swellings, pleonites 5 and 6 with large median granulate swelling. Cheliped palm upper margin with inward-folding crest, median part with spines or teeth. Maximum cl. 9 mm. Slope (235–244 m; near volcanic vent). Central Indo-Pacific (New Caledonia). 1 species (Ng and Chia 1994).

Cyrtocarcinus Ng & Chia, 1994 Diagnosis. Front with narrow fissure, frontal lobes directed anteriorly. Cardio-intestinal region deep, cardiac fold over-reaching groove, posterior part truncate; anterolateral margin lamelliform, with low, broadly triangular, lobiform teeth. Thoracic sternite 4 posterior part sloping steeply forming strong transverse ridge. Pleonites 2–6 smooth. Cheliped palm upper margin rounded, inner surface strongly swollen. Maximum cl. 20 mm (Pl. 64n). Subtidal–slope (3–424 m). Central and Eastern Indo-Pacific. 1 species (Naruse et al. 2020: redescription; Ng and Chia 1994).

Glyptocarcinus Takeda, 1973 Diagnosis. Front with narrow fissure, frontal lobes directed anteriorly. Cardio-intestinal region deep, cardiac fold over-reaching

groove, posterior part broadly triangular. Anterolateral margin lamelliform, with well developed, acutely triangular teeth. Thoracic sternite 4 posterior part sloping gradually forming low transverse ridge. Pleonites 2–6 smooth. Chela upper margin slightly cristate; inner surface gently convex. Maximum cl. 11.4 mm. Shelf, slope (150–275 m; rocky substrates). Central IndoPacific. 2 species (Ng and Chia 1994: key to species).

Subfamily Banareiinae Števčić, 2005 Banareiines are large, usually hairy crabs, often associated with soft corals. Grouping the four enigmatic genera together has been much discussed (Guinot 1967c; Serène 1984; Števčić 2005) but they remain weakly diagnosed. Much early discussion used the subfamily names Zalasiinae Serène, 1968 or Trichiinae Serène, 1984. Lai et  al. (2011) showed that Pseudactaea, formerly considered to be an actaeine, is a banareiine. Diagnosis. Carapace subcircular to transversely subovate, wider than long, convex, regions usually distinct, lateral margins weakly to strongly dentate, densely covered with short or long setae. Antennules folding obliquely forwards. Chelipeds subequal, similar in both sexes; fingers laterally compressed, thin, tips pointed, usually crossing. Pereopods 2–5 without dactylo-­ propodal ‘lock’.

Key to genera of Banareiinae 1. – 2. – 3. –

Buccal cavity long, oval, narrow. Maxilliped 3 elongate; merus longer than wide, distolateral angle obsolete; meri in contact (Fig. 14.152g)������������������������������������������������������������������������������������������������������������������������� Trichia Buccal cavity quadrate, broad. Maxilliped 3 rectangular; merus usually wider than long, distolateral angle square; meri separated by space occupied by palps (Fig. 14.152e, f)�������������������������������������������������������������������������2 Carapace anterolateral margins rimmed with narrow crest (Fig. 14.152b). Cheliped carpus and pereopods 2–5 carpi and propodi upper margins crested (Fig. 14.152h)���������������������������������������������������������������������������Pseudactaea Carapace anterolateral margins without narrow crest. Cheliped carpus and pereopods 2–5 carpus and propodus upper margins without crests����������������������������������������������������������������������������������������������������������������������������������3 Carapace dorsal surface weakly convex, with regions distinct or not, granular, with felt of long setae; anterolateral margin with 4 lobes, more or less developed, granular (Fig. 14.152a)�������������������������������������������Banareia Carapace dorsal surface strongly convex, with regions indistinct, minutely granular or smooth, with felt of short setae; anterolateral margin convex, entire (Fig. 14.152c)������������������������������������������������������ Calvactaea

14 – Brachyura – crabs

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Fig. 14.152.  Xanthidae, Banareiinae. a, Banareia fatuhiva Davie, 1993 (setae cleared from half carapace); b, Pseudactaea corallina (Alcock, 1898). Carapace, setae not shown: c, Calvactaea; d, Trichia. Front, antennae, antennules, maxillipeds 3: e, Banareia; f, Calvactaea; g, Trichia. h, pereopods 3, 4, Pseudactaea.

Banareia A. Milne-Edwards, 1869 Diagnosis. Carapace weakly convex, with regions distinct or not, granular, with felt of long setae; anterolateral margin with 4 lobes, more or less developed, granular. Buccal cavity quadrate, broad. Maxilliped 3 rectangular; merus usually wider than long, distolateral angle square; meri separated by space occupied by palps. Maximum cl. 35 mm (Pl. 64o). Intertidal–slope (0–250 m). Tropical W Atlantic, Temperate Northern W Pacific, Indo-West Pacific, Temperate Australia. 16 species (Guinot 1976: species descriptions; Serène 1984: key to 15 Indo-West Pacific species).

Calvactaea Ward, 1933 Diagnosis. Carapace strongly convex, with regions indistinct, minutely granular or smooth, with felt of short setae; anterolateral margin evenly convex, entire. Buccal cavity quadrate, broad. Maxilliped 3 rectangular; merus usually wider than long, distolateral angle square; meri separated by space occupied by palps. Maximum cl. 20 mm (Pl. 64p). Subtidal, shelf (10–106 m; inside cavities in the walls of, and climbing between, alcyonarian soft corals). Temperate Northern W Pacific, Central Indo-Pacific, Temperate SE Australia. 1 species (Poore 2004: figure; Sakai 1976: figure; Ward 1933).

Pseudactaea Serène, 1962 Diagnosis. Carapace weakly convex, with regions distinct or not, granular, with felt of short setae; anterolateral margin evenly

convex or with 3 broadly obtuse angular lobes. Buccal cavity quadrate, broad. Maxilliped 3 rectangular; merus wider than long, distolateral angle square; meri separated by space occupied by palps. Cheliped carpus and pereopods 2–5 carpus and propodus upper margins crested. Maximum cl. 11 mm. Subtidal, shelf (15–150 m). Western and Central Indo-Pacific. 3 species (Takeda and Marumura 2002: figures, comparison of species).

Trichia De Haan, 1839 Diagnosis. Carapace ovate to hexagonal, strongly convex, granular, densely covered in long setae; anterolateral margin angular, produced. Buccal cavity long, oval, narrow. Maxilliped 3 elongate; merus longer than wide, distolateral angle obsolete; meri in contact. Maximum cl. 57 mm (Pls 64q, 65a). Intertidal, subtidal (0–115 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate. Australia. 6 species (Guinot 1976: species descriptions). Zalasius Rathbun, 1897 is a synonym.

Subfamily Chlorodiellinae Ng & Holthuis, 2007 Chlorodielline xanthids have spoon-shaped cheliped finger tips and pereopods 2–5 with a dactylo-propodal ‘lock’ formed by rounded prolongation of the propodal lateral margin. The front is especially wide, typically with large submedian lobes and 2 small lateral lobules. Most occur on

760

Marine Decapod Crustacea

Indo-West Pacific shallow coral or rocky reefs, some extending into warm temperate regions. Only one species occurs in the tropical Western Atlantic. Molecular evidence has suggested that the subfamily is not monophyletic (­Lasley et al. 2015). These authors rediagnosed Chlorodiellinae but included only five genera. They discussed the morphological differences and similarities of the excluded genera but could not decide on subfamily placement. Our key retains the ‘traditional’ arrangement for pragmatic reasons. Diagnosis. Front wider than one-third maximum carapace width, typically with 2 large submedian lobes and 2

small lateral lobules, or nearly straight, without lateral sinuses; anterolateral margin typically with 4 teeth behind exorbital angle (sometimes 2 or 3). Antennules folding transversely. Chelipeds with finger tips rounded, expanded, hollowed on occlusal surface, spoon-shaped, meeting when closed; pereopods 2–5 with dactylo-propodal ‘lock’ formed by rounded prolongation of propodal lateral margin sliding against and beneath projecting button situated proximally on lateral margin of dactylus (if not clearly differentiated then dactylus apically bifid.

Key to genera of Chlorodiellinae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. – 9. –

Antennal basal article with elongated distolateral extension completely blocking or extending halfway or more, partially or completely closing hiatus, excluding free articles of antennal peduncle from orbit (Fig. 14.153m–o)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Antennal basal article with distolateral extension absent or minimally blocking orbital hiatus, enabling free articles of antennal peduncle to freely enter orbit (Fig. 14.153k, l)������������������������������������������������������������������������������6 Carapace surface smooth, unadorned, with only slightly raised, indistinctly defined regions (Fig. 14.153h)��������� Ratha Carapace surface with raised, distinct regions, adorned with setae and granules����������������������������������������������������������������3 Cheliped merus with longitudinal groove on upper surface (Fig. 14.153f). Pereopods 2–5 dactyli with distal corneous unguis distinctly smaller than adjacent flexor spine (Fig. 14.153u)����������������������������������������������������Sulcodius Cheliped merus without longitudinal groove on upper surface. Pereopods 2–5 dactyli with distal corneous unguis larger than adjacent flexor spine (Fig. 14.153t)����������������������������������������������������������������������������������������������������������4 Antennal basal article with elongated distolateral extension extending almost halfway into orbital hiatus (Fig. 14.153o)���������������������������������������������������������������������������������������������������������������������������������������������������������������������Soliella Antennal basal article with elongated distolateral extension extending beyond halfway or completely blocking orbital hiatus (Fig. 14.153m, n)���������������������������������������������������������������������������������������������������������������������������������5 Carapace anterolateral teeth simple (Fig. 14.153c)��������������������������������������������������������������������������������������������������������� Luniella Carapace anterolateral teeth with secondary spines or acute granules (Fig. 14.153d)��������������������������������������������� Pilodius Carapace regions prominent and projecting (2M and 3M always well defined, usually also L), surface rugose, distinctly granular (Fig. 14.153b)��������������������������������������������������������������������������������������������������������������������������������7 Carapace regions undefined or only poorly defined, surface smooth or finely granular (Fig. 14.153a)����������������������������9 Chelipeds merus extending little beyond lateral margin of the carapace; fixed finger truncate, occlusal margin almost perpendicular to axis of palm (Fig. 14.153p). Pereopods 2–5 short, robust; meri hardly longer than maximum width������������������������������������������������������������������������������������������������������������������������������������ Tweedieia Chelipeds elongate, merus extending laterally well beyond the lateral margin of the carapace; fixed finger elongated, occlusal margin almost in line with axis of palm. Pereopods 2–5 typical; meri about twice as long as wide (Fig. 14.153b)���������������������������������������������������������������������������������������������������������������������������������������������������������8 Cheliped fingers with tufts of setae near cutting margins (Fig. 14.153q). Pereopods 2–5 dactyli with corneous unguis subdistal, pectinate, distinctly smaller than adjacent distal flexor spine (Fig. 14.153v)����� Vellodius Cheliped fingers without tufts of setae near cutting margins. Pereopods 2–5 dactyli with corneous unguis distal, equal to or larger than adjacent flexor spine (Fig. 14.153r)������������������������������������������������������������������������Cyclodius Distance between inner orbital angles equal to half carapace width (Fig. 14.153a). Gonopod 1 with subapical setae long, directed proximally or short, perpendicular to gonopod axis (Fig. 14.153w)���������� Chlorodiella Distance between inner orbital angles 0.4 times carapace width (Fig. 14.153g). Gonopod 1 with subapical setae long, directed distally (Fig. 14.153x)��������������������������������������������������������������������������������������������������������� Liocarpilodes

14 – Brachyura – crabs

761

Fig. 14.153.  Xanthidae, Chlorodiellinae. a, Chlorodiella cytherea (Dana, 1852); b, Cyclodius obscurus (Hombron & Jacquinot, 1846); c, Luniella pugil (Dana, 1852); d, Pilodius areolatus (H. Milne Edwards, 1834); e, Soliella flava (Rathbun, 1894); f, Sulcodius deflexus (Dana, 1852). Carapace: g, Liocarpilodes; h, Ratha; i, Tweedieia; j, Vellodius. Left antenna (basal article shaded), orbit hiatus: k, Chlorodiella; l, Cyclodius; m, Luniella; n, Pilodius; o, Soliella. Chela, outer face: p, Tweedieia; q, Vellodius. Pereopod 5 dactylus: r, Cyclodius; s, Chloridiella; t, Pilodius; u, Sulcodius; v, Vellodius. Gonopod 1 apex: w, Chlorodiella; x, Liocarpilodes.

Chlorodiella Rathbun, 1897 Diagnosis. Carapace regions undefined or only poorly defined, surface smooth or finely granular; front-exorbital width 0.5 times carapace width. Antennal basal article with distolateral extension absent or minimally blocking orbital hiatus, enabling free articles of antennal peduncle to enter orbit. Pereopods 2–5 dactyli with distal

corneous unguis distinctly smaller than adjacent flexor spine. Gonopod 1 with subapical setae long, directed proximally or short, perpendicular to gonopod axis. Maximum cl. 14 mm (Pl. 65c). Intertidal–shelf (0–124 m). Indo-West Pacific, Temperate ­Australasia. 11 species (Lasley et al. 2015: rediagnosis; Serène 1984: key to 10 species).

762

Marine Decapod Crustacea

Cyclodius Dana, 1851 Diagnosis. Carapace regions prominent and projecting (2M and 3M always well defined, usually also L), surface rugose, distinctly granular. Antennal basal article with distolateral extension slightly blocking orbital hiatus, enabling free articles of antennal peduncle to enter orbit. Chelipeds elongate, merus extending laterally well beyond the lateral margin of the carapace; fixed finger elongated, occlusal margin almost in line with axis of palm; fingers without tufts of setae near cutting margins. Pereopods 2–5 dactyli with distal corneous unguis larger than adjacent flexor spine. Maximum cl. 21 mm (Pl. 65d). Intertidal–shelf (0–97 m). Indo-West Pacific, Temperate Southern Africa. 8 species (Lasley et al. 2015: rediagnosis; Serène 1984: key to 7 species as Phymodius A. Milne-Edwards, 1863, now a junior synonym).

Liocarpilodes Klunzinger, 1913 Diagnosis. Carapace regions undefined or only poorly defined, surface smooth or finely granular; front-exorbital width 0.4 times carapace width. Antennal basal article with distolateral extension absent or minimally blocking orbital hiatus, enabling free articles of antennal peduncle to enter orbit. Gonopod 1 with subapical setae long, directed distally. Maximum cl. 8.4 mm (Pl. 65e). Intertidal–shelf (0–30 m). Indo-West Pacific. 6 species (Serène 1984: key to 5 species and subspecies).

Subtidal, shelf (5–154 m). Tropical W Atlantic. 1 species (Lasley et al. 2013).

Soliella Lasley, Klaus & Ng, 2015 Diagnosis. Carapace regions well defined; anterolateral margin with 4 tipped with anteriorly directed spine and smaller accessory spines. Antennal basal article with elongated distolateral extension extending almost halfway into orbital hiatus, partially closing hiatus. Pereopods 2–5 dactyli with distal corneous unguis larger than adjacent flexor spine. Maximum cl. 11 mm. Intertidal, subtidal. Indo-West Pacific. 2 species (Lasley et al. 2015).

Sulcodius Clark & Ng, 1999 Diagnosis. Carapace regions well defined. Antennal basal article with elongated distolateral extension completely blocking hiatus. Cheliped merus with longitudinal groove on upper surface. Pereopods 2–5 dactyli with distal corneous unguis distinctly smaller than adjacent flexor spine. Maximum cl. 15 mm. Subtidal, shelf (1–29 m). Western and Central Indo-Pacific. 1 species (Clark and Ng 1999: comparison of Sulcodius, Pilodius, Phymodius and Chlorodiella).

Tweedieia Ward, 1934

Diagnosis. Carapace regions well defined; anterolateral margin with simple teeth. Antennal basal article with elongated distolateral extension extending well into orbital hiatus, completely closing hiatus. Pereopods 2–5 dactyli with distal corneous unguis larger than adjacent flexor spine. Maximum cl. 11 mm. Intertidal–shelf (0–30 m). Indo-West Pacific. 4 species (Lasley et al. 2015; Serène 1984: key to species as Pilodius).

Diagnosis. Carapace regions well defined. Antennal basal article with distolateral extension slightly blocking orbital hiatus, enabling free articles of antennal peduncle to enter orbit. Chelipeds merus extending little beyond lateral margin of the carapace; fixed finger truncate, occlusal margin almost perpendicular to axis of palm. Pereopods 2–5 short, robust; meri hardly longer than maximum width. Maximum cl. 7 mm (Pl. 65b). Intertidal, subtidal (0–20 m). Indo-West Pacific. 2 species (­Lasley et al. 2022).

Pilodius Dana, 1851

Vellodius Ng & Yang, 1998

Diagnosis. Carapace regions well defined; anterolateral margin with teeth having secondary spines or acute granules. Antennal basal article with elongated distolateral extension extending beyond halfway into orbital hiatus, almost closing hiatus. Pereopods 2–5 dactyli with distal corneous unguis larger than adjacent flexor spine. Maximum cl. 42 mm (Pl. 65f). Intertidal–shelf (0–25 m). Indo-West Pacific, Temperate Southern Africa, Temperate Australasia. 10 species (Clark and Galil 1993: rediagnosis, key to 15 species, some now in other genera; Lasley et al. 2015: rediagnosis).

Diagnosis. Carapace regions well defined, 2F separate, 2F fused with 1M, 2M entire, 2M and 3L fused, 5L and 4L separate. Antennal basal article with distolateral extension absent, enabling free articles of antennal peduncle to enter orbit. Chelipeds elongate, merus extending laterally well beyond the lateral margin of the carapace; fixed finger elongated, occlusal margin almost in line with axis of palm; cheliped fingers with tufts of setae near cutting margins. Pereopods 2–5 dactyli with distal corneous unguis distinctly smaller than adjacent flexor spine. Maximum cl. 10 mm. Subtidal. Central Indo-Pacific (South China Sea, Japan). 1 species (Ng and Yang 1998; Takeda and Miyake 1968: description of only species as Pilodius etisoides Takeda & Miyake, 1968).

Luniella Lasley, Klaus & Ng, 2015

Ratha Lasley, Lai & Thoma, 2013 Diagnosis. Carapace surface smooth, unadorned, with only slightly raised, indistinctly defined regions. Antennal basal article with elongated distolateral extension extending halfway, partially closing hiatus. Pereopods 2–5 dactyli without flexor spine. Maximum cl. 13 mm (Pl. 65g).

Subfamily Cymoinae Alcock, 1898 The subfamily is represented only by Cymo, a genus recognised by its flat, circular or longitudinally ovate carapace.

14 – Brachyura – crabs

763

Diagnosis. Front narrower than one-third maximum carapace width, 2- or 4-lobed, with median fissure, projecting between lateral sinuses separating it from supraorbital angles; anterolateral margin with 4–8 teeth behind exorbital angles. Antennules folding obliquely forwards. Chelipeds with finger tips rounded, expanded, hollowed on occlusal surface, spoon-shaped, meeting when closed; pereopods 2–5 with dactylo-propodal ‘lock’ formed by rounded prolongation of propodal lateral margin sliding against and beneath projecting button situated proximally on lateral margin of dactylus (if not clearly differentiated then dactylus apically bifid). Etisus H. Milne Edwards, 1834

Fig. 14.154.  Xanthidae, Cymoinae. Cymo quadrilobatus Miers, 1884.

Species of Cymo are obligate associates of living acroporid and pocilloporid scleratinian corals. Diagnosis. Carapace subcircular or longitudinally subovate, slightly longer to scarcely wider than long, flat, regions usually indistinct, lateral margins weakly dentate, sparsely to densely setose. Antennules folding obliquely forwards. Chelipeds strongly unequal, dissimilar in both sexes; fingers curved conical, tips rounded, hollowed, meeting; pereopods 2–5 without dactylo-propodal ‘lock’. Cymo De Haan, 1833 Maximum cl. 15 mm (Pl. 65h, i). Intertidal, shelf (0–30 m). IndoWest Pacific, Temperate Southern Africa, Temperate Australia. 8 species (Ho and Ng 2005: key to species).

Diagnosis. Carapace transversely ovate to pentagonal; surface weakly convex to flattened transversely and longitudinally; regions distinct in adults, 3M well demarcated; front anterior margin variously shaped, not with even, semicircular concavity; anterolateral margin with 4–8 angular or sharp anteriorly or anterolaterally inclined teeth behind exorbital angle. Maximum cl. 80 mm. (Pl. 66a). Intertidal–shelf (0–95 m). Tropical W Atlantic, Indo-West Pacific, Temperate Southern Africa, Temperate Australia. 26 species (Felder and Thoma 2010: Gulf of Mexico species; Sakai 1976: key to 7 Japanese species; Serène 1984: key to 16 species).

Paraetisus Ward, 1933 Diagnosis. Carapace subhexagonal, 1.2 times as wide as long; surface strongly transversely and longitudinally convex; regions indistinct or largely absent in adults, 3M faintly indicated anteriorly only by slender median part; front anterior margin with even, semicircular concavity; anterolateral margin with 4 prominent blunt, rounded anterolateral lobes behind exorbital angle. Maximum cl. 34 mm. Subtidal (18–27 m). Central Indo-Pacific (NE Australia). 1 species (Serène 1984: photo of Australian specimen of only species, Paraetisus globulus Ward, 1933; Ward 1933). Serène’s (1984: pl. 32F) figured specimen from Madagascar appears to represent a juvenile of Etisus godeffroyi (A. Milne Edwards, 1873).

Subfamily Etisinae Ortmann, 1893 spooner crabs Etisine crabs are cryptic, residing under coral and boulders on coral and rocky reefs but are commonly reported in faunal surveys. All have spoon-shaped tips on the cheliped fingers. A button on the dactylus and a modified propodal margin of pereopods 2–5 result in a characteristic form of articulation between these two articles. Etisus is speciose and widespread but Paraetisus is monotypic and known from only a few specimens from Australia. Etisids differ from chlorodiellines, which share these features, in having a narrower, more projecting front.

Subfamily Euxanthinae Alcock, 1898 Euxanthine xanthids are notable for the way in which the anterior part of anterolateral margin of the carapace descends and ends anteriorly below the orbit near the upper outer corner of the buccal frame (Figs 14.156i, k, 14.157g) rather than joining the exorbital tooth. This feature, however, is not well expressed in all genera (Ng and Clark 2003; Ng et al. 2008). Euxanthines share this feature with glyptoxanthines which differ in having a more or less projecting front. Many genera have the chelipeds and pereopods, when folded, closely fitting to depressions on the sides of

764

Marine Decapod Crustacea

Fig. 14.155.  Xanthidae, Etisinae. a, Etisus sp. b, Paraetisus globulus Ward, 1933 (holotype); c, chela, Etisus. d, dactylo-propodal ‘lock’, pereopod 5, facial and upper views, Etisus.

the carapace (Mendoza and Ng 2008a). The subfamily is unlikely to be monophyletic (Lai et al. 2011). Jacforus, with anterolateral carapace margins that descend below the orbits (Ng and Clark 2003), is treated as a euryxanthine rather than xanthine. Similarly, Psaumis is a euxanthine rather than actaeine as demonstrated by Lai et al. (2011). Euxanthines inhabit rocky environments at shelf and slope depths, more rarely in the intertidal, but their ecology has attracted little attention. Many genera have a strong

basal tooth on the dactylus of the major chela that may be used to prey on gastropods (Ng and Tan 1985). Diagnosis. Front horizontal, appearing subtly 2-lobed, almost straight in anterior view; anterolateral margin descending anteriorly, not meeting exorbital angle but continuing below orbit towards anterolateral corner of buccal frame. Antennules folding transversely. Chelipeds with finger tips acute or expanded, spoon-shaped; pereopods 2–5 without dactylo-propodal ‘lock’.

Key to genera of Euxanthinae 1.

– 2. – 3. – 4. – 5.

Carapace with prominent triangular projection near midlength of posterolateral margin. Cheliped and pereopods 2–5, when folded against carapace, coapting to form deep tube-like hollow bounded by posterior margin of cheliped, upper margins of pereopods 2–5 meri and posterolateral margin of carapace (Fig. 14.157l)�������������������������������������������������������������������������������������������������������������������������������������������� Pleurocolpus Carapace posterolateral margin straight or concave, without triangular projection near midlength. Cheliped and pereopods 2–5, when folded against carapace, not forming deep tube-like hollow��������������������������������2 Carapace subhepatic cavities clearly defined�������������������������������������������������������������������������������������������������������������������������������3 Carapace subhepatic cavities absent����������������������������������������������������������������������������������������������������������������������������������������������5 Carapace with upper part of subhepatic cavities forming concave ‘cut-out’ on anterolateral margins (Fig. 14.156j, k)�������������������������������������������������������������������������������������������������������������������������������������������������������� Hepatoporus Carapace with openings of subhepatic cavities without concave ‘cut-out’ on anterolateral margins (Figs 14.156h, i, 14.158d)������������������������������������������������������������������������������������������������������������������������������������������������������������4 Carapace broadly trapezoidal–ovate, distinctly wider than long; anterolateral margin lobate or dentate; surface rugose, regions indicated (Fig. 14.156l)��������������������������������������������������������������������������������������������������� Hypocolpus Carapace narrowly ovate, little wider than long; anterolateral margin entire; surface finely granular near margins, otherwise smooth, surface without trace of regions (Fig. 14.156h, i)���������������������������������������������� Guinotellus Chelipeds with junction of carpus and propodus forming large circular hollow when folded against carapace (Fig. 14.158r)���������������������������������������������������������������������������������������������������������������������������������������������Carpoporus

14 – Brachyura – crabs

– 6. – 7.

765

Chelipeds not forming circular hollow at junction of carpus and propodus when folded���������������������������������������������������6 Pereopods 2–5 usually folded into concavity of posterolateral margin by coaptation����������������������������������������������������������7 Pereopods 2–5 not folded by coaptation against posterolateral margin of carapace which is nearly straight��������������� 12 Movable articles of antennal peduncle and flagellum not visible in anterior view, concealed within orbit (Fig. 14.158b, e)��������������������������������������������������������������������������������������������������������������������������������������������� Lipaesthesius

Fig. 14.156.  Xanthidae, Euxanthinae. a, Alainodaeus akiaki Davie, 1993; b, Batodaeus urinator (A. Milne-Edwards, 1880); c, Cranaothus deforgesi Ng, 1993; d, Danielea Ng, 2003; e, Epistocavea mururoa Davie, 1993; f, Edwardsium spinimanus (H. Milne Edwards, 1834); g, Euxanthus exsculptus; h, i, Guinotellus melvillensis Serène, 1971 (dorsal, anterior); j, k, Hepatoporus pumex Mendoza & Ng, 2008 (dorsal, anterior); l, Hypocolpus mararae Crosnier, 1991; m, Jacforus cavatus (Rathbun, 1907).

766

– 8. –

Marine Decapod Crustacea

Movable articles of antennal peduncle and flagellum visible in anterior view����������������������������������������������������������������������8 Carapace and chelipeds deeply pitted, often forming reticulated pattern (Fig. 14.157m, p). Male thoracic sternum with large, rounded, basin-like cavity anterior to telson (Fig. 14.158m, n)��������������������������������������������������������9 Carapace and chelipeds variously smooth or rugose, with or without tubercles and ridges, but not deeply pitted and reticulated. Male thoracic sternum smooth or eroded, without large, rounded, basin-like cavity anterior to telson (Fig. 14.158h, i)��������������������������������������������������������������������������������������������������������������������������������10

Fig. 14.157.  Xanthidae, Euxanthinae. a, Ladomedaeus fungillus Manuel-Santos & Ng, 2007; b, Medaeops granulosus (Haswell, 1882); c, Medaeus grandis Davie, 1993; d, Miersiella haswelli (Miers, 1886); e, Monodaeus guinotae Forest, 1976; f, g, Olenothus uogi Ng, 2002 (dorsal, anterior); h, Palatigum trichostoma Davie, 1997; i, Paramedaeus megagomphios Davie, 1997; j, Paraxanthodes obtusidens (Sakai, 1964); k, Pilomedaeus okutanii Takeda, 2011; l, Pleurocolpus boileaui Crosnier, 1995; m, Psaumis cavipes Dana, 1852; n, Rizalthus anconis Mendoza & Ng, 2008; o, Takedax gabrielae Mendoza & Ng, 2012; p, Visayax osteodictyon Mendoza & Ng, 2008.

14 – Brachyura – crabs

767

9.

Pereopods 2–5 carpi and propodi with prominent, strongly curved crest, hollowed on anterior face forming shallow ‘cup’, pereopod surfaces otherwise unridged (Pl. 66j)���������������������������������������������������������������� Psaumis – Pereopods 2–5 carpi and propodi smooth, with few low ridges and granules or deeply eroded and pitted (Fig. 14.158u)��������������������������������������������������������������������������������������������������������������������������������������������������������Visayax 10. Carapace anterolateral margin with prominent teeth (Fig. 14.156g)����������������������������������������������������������������������Euxanthus – Carapace anterolateral margin curved, irregular or evenly granular, but without prominent teeth�������������������������������11 11. Carapace regions well defined, densely covered with large granules (Fig. 14.157n). Male pleonite 6 longitudinally rectangular (Fig. 14.158p)����������������������������������������������������������������������������������������������������������������� Rizalthus – Carapace regions weakly defined, finely granulate (Fig. 14.157f, g) Male pleonite 6 slightly wider than long, appearing square�����������������������������������������������������������������������������������������������������������������������������������������������Olenothus 12. Maxilliped 3 entirely and densely covered with short setae (Fig. 14.158q)�������������������������������������������������������������Palatigum – Maxilliped 3 at most sparsely setose�������������������������������������������������������������������������������������������������������������������������������������������13 13. Carapace densely covered by short, dense tomentum obscuring regions and granulation (Fig. 14.157k). Fused male pleonites 3–5 with complete transverse grooves demarcating pleonites���������������������������������� Pilomedaeus – Carapace surface glabrous or with scattered setae not obscuring regions and granulation. Fused male pleonites 3–5 usually without transverse sutures demarcating pleonites, at most partially indicated at lateral margins or between 3 and 4����������������������������������������������������������������������������������������������������������������������������������������14 14. Carapace anterolateral margin longer than posterolateral margin����������������������������������������������������������������������������������������15 – Carapace anterolateral margin as long as or shorter than posterolateral margin����������������������������������������������������������������17

Fig. 14.158.  Xanthidae, Euxanthinae. Carapace: a, Crosnierius; b, Lipaesthesius; c, Lipkemedaeus. d, Hypocolpus, left subhepatic region. Left antennule, antenna, orbit: e, Lipaesthesius; f, Paraxanthodes (with epistome); g, Visayax. Male thoracic sternites 1–4, pleonite 6, telson: h, Batodaeus; i, Cranaothus; j, Danielea; k, Jacforus; l, Ladomedaeus; m, Psaumis; n, Visayax. o, left male pleonites 1, 2, thoracic sternites 8 (shaded), pereopodal coxa 5, Monodaeus. p, male pleonite 6, telson, Rizalthus. q, maxilliped 3, Palatigum. Chela: r, Carpoporus; s, Crosnierius. Pereopod 5: t, Crosnierius; u, Visayax. Gonopod 1: v, Alainodaeus; w, Medaeops; x, Miersiella; y, Paraxanthodes.

768

15. – 16. – 17. – 18. – 19. – 20. – 21. – 22. – 23. – 24. – 25. – 26. – 27. – 28. –

Marine Decapod Crustacea

Carapace frontal lobes produced well in advance of orbits, anterior margins sloping posteriorly (Fig. 14.156c). Male telson not reaching midlength of sternite 4 (Fig. 14.158i)���������������������������������������������� Cranaothus Carapace frontal lobes weakly produced in advance of orbits, anterior margins transverse. Male telson extending anteriorly to or beyond midlength of sternite 4���������������������������������������������������������������������������16 Carapace 1.2–1.5 times as wide as long. Cheliped palm upper inner surface bluntly irregular, without crest (Fig. 14.156m). Male telson half width of pleonite 3, reaching beyond midlength of thoracic sternite 4 (Fig. 14.158k)������������������������������������������������������������������������������������������������������������������������������������������������������������������� Jacforus Carapace 1.5–1.6 times as wide as long. Cheliped palm upper inner surface with thin crest (Fig. 14.156d). Male telson about one-third width of pleonite 3, reaching midlength of thoracic sternite 4 (Fig. 14.158j)�������������������������������������������������������������������������������������������������������������������������������������������������������������������Danielea Carapace anterolateral margin shorter than posterolateral margin (Fig. 14.156f)��������������������������������������������� Epistocavea Carapace anterolateral margin as long as posterolateral margin�������������������������������������������������������������������������������������������18 Carapace anterolateral margin with slender spines, often with secondary spinules basally (Fig. 14.158c)���������Lipkemedaeus Carapace anterolateral margin with blunt, nodular or triangular lobes, not spiniform (Fig. 14.158a)����������������������������19 Cheliped palm upper margin with thin, smooth, evenly curved inwardly inclined crest (Fig. 14.158s). Pereopods 2–5 meri upper margins with upright spines; carpi and propodi upper margins with thin rounded crests (Fig. 14.158t)�������������������������������������������������������������������������������������������������������������������������� Crosnierius Cheliped palm upper margin variously ornamented but without thin curved crest. Pereopods 2–5 meri upper margin usually unarmed or minutely dentate, but if spinous, carpi and propodi upper margins without crests���������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 20 Male telson reaching beyond midlength of sternite 4 (Fig. 14.158h)�������������������������������������������������������������������������������������21 Male telson not reaching midlength of sternite 4 (Fig. 14.158l)��������������������������������������������������������������������������������������������� 22 Carapace region 4M not indicated; regions 2R and 3R fused; region P1 not indicated, without transverse ridge (Fig. 14.156b)���������������������������������������������������������������������������������������������������������������������������������� Batodaeus Carapace region 4M distinct; regions 2R and 3R separated by groove; region P1 with transverse ridge (Fig. 14.157c)�������������������������������������������������������������������������������������������������������������������������������������������������������������������Medaeus Male pleonite 6 as long as or longer than wide, appearing longitudinally rectangular����������������������������������������������������� 23 Male pleonite 6 wider than long, usually appearing transversely rectangular�������������������������������������������������������������������� 25 Carapace region 4M not indicated (Fig. 14.157i). Fused male pleonites 3–5 without complete transverse grooves demarcating pleonites, at most indicated at lateral margins����������������������������������������Paramedaeus Carapace region 4M distinct (Fig. 14.156e). Fused male pleonites 3–5 with transverse grooves demarcating pleonites������������������������������������������������������������������������������������������������������������������������������������������������������������� 24 Male telson semicircular�������������������������������������������������������������������������������������������������������������������������������������������� Edwardsium Male telson triangular (Fig. 14.158l)����������������������������������������������������������������������������������������������������������������������� Ladomedaeus Male thoracic sternite 8 with small part visible between edge of pleonite 2 and condyle of pereopod 5 coxa (Fig. 14.158o)����������������������������������������������������������������������������������������������������������������������������������������������������������������� 26 Male thoracic sternite 8 fully concealed by pleonite 2������������������������������������������������������������������������������������������������������������� 27 Carapace last anterolateral tooth directed posterolaterally; surface just inward of posterolateral corner with large conical tubercle (Fig. 14.157o)�������������������������������������������������������������������������������������������������������������������Takedax Carapace last anterolateral tooth directed laterally or anterolaterally; posterolateral surface without large conical tubercle, at most with low flat swelling (Fig. 14.157e)����������������������������������������������������������������� Monodaeus Pereopod 5 merus 4 times as long as wide (Fig. 14.156a)�������������������������������������������������������������������������������������������������������� 28 Pereopod 5 merus less than 3.5 times as long as wide (Fig. 14.157b, j)��������������������������������������������������������������������������������� 29 Carapace region 2M undivided or at most partially divided in anterior one-fourth by shallow groove; region 4M indistinct (Fig. 14.156a). Gonopod 1 with convex flange over proximal two-thirds, distal third strongly tapering (Fig. 14.158v)�������������������������������������������������������������������������������������������������������������������Alainodaeus Carapace region 2M longitudinally divided for more than half length; region 4M distinct (Fig. 14.157d). Gonopod 1 thick, weakly tapering, margins subparallel for most of distal two-thirds (Fig. 14.158x)���������� Miersiella

14 – Brachyura – crabs

29. –

769

Endostomial ridges complete, extending anteriorly to epistome (Fig. 14.158f). Gonopod 1 slender, distally curved, with short subdistal spines (Fig. 14.158y)�������������������������������������������������������������������������� Paraxanthodes Endostomial ridges short, indicated proximally only, not extending anteriorly to epistome. Gonopod 1 slender, curved, with lateral keel and long, subdistal setae (Fig. 14.158w)������������������������������������ Medaeops

Alainodaeus Davie, 1993 Diagnosis. Carapace ovate, ~1.4 times as wide as long; region 2M undivided or at most partially divided in anterior onefourth by shallow groove; region 4M indistinct; anterolateral margin with 4 teeth. Pereopod 5 merus 4 times as long as wide. Male pleonite 6 wider than long, usually appearing transversely rectangular. Gonopod 1 with convex flange over proximal twothirds, distal third strongly tapering. Maximum cl. 24 mm (Pl. 66b). Shelf, slope (35–573 m). Central and Eastern Indo-Pacific. 5 species (Mendoza and Ng 2008b: remarks, key to species).

Batodaeus Vázquez-Bader & Gracia, 2004

Shelf (180–200 m). Central Indo-Pacific. 2 species (Ng et  al. 2005: second species; Serène and Vadon 1981).

Danielea Ng & Clark, 2003 Diagnosis. Carapace 1.5–1.6 times as wide as long; regions weakly defined; front lobes weakly produced in advance of orbits, anterior margins transverse; anterolateral margin longer than posterolateral margin. Cheliped palm upper inner surface with thin crest. Male telson about one-third width of pleonite 3, reaching midlength of thoracic sternite 4. Maximum cl. 10 mm (Pl. 66c). Shelf (25–30 m). Temperate Northern W Pacific, Indo-West Pacific. 1 species (Ng and Clark 2003).

Edwardsium Guinot, 1967

Diagnosis. Carapace subhexagonal, wider than long; region 4M not indicated; regions 2R and 3R fused; region P1 not indicated, without transverse ridge; anterolateral margin with prominent teeth. Male telson reaching beyond midlength of sternite 4. Maximum cl. 17 mm. Shelf, slope (146–458 m). Temperate Northern W and Tropical Atlantic. 1 species (Vázquez-Bader and Gracia 2004; Williams 1984: description, figure).

Diagnosis. Carapace ovate, ~1.4 times as wide as long; regions well defined; anterolateral margin with 4 teeth. Male pleonites 3–5 with complete transverse grooves demarcating pleonites. Male pleonite 6 longitudinally rectangular. Maximum cl. 20 mm (Pl. 66d). Subtidal–slope (10–275 m). Tropical W Atlantic, Tropical Eastern Pacific. 4 species (Guinot 1967a; Rathbun 1930: figure as Medaeus spinimanus A. Milne Edwards, 1834).

Carpoporus Stimpson, 1871

Epistocavea Davie, 1993

Diagnosis. Carapace as wide as long; anterolateral margin with 3 small teeth. Chelipeds with junction of carpus and propodus forming large circular hollow when folded against carapace. Maximum cl. 13 mm. Shelf (32–120 m). Temperate Northern W Atlantic. 1 species (Rathbun 1930: rediagnosis).

Diagnosis. Carapace ovate, ~1.4 times as wide as long; surface rugose, regions indicated; anterolateral margin shorter than posterolateral margin, dentate and granulate. Maximum cl. 29 mm. Shelf, slope (50–500 m). Eastern Indo-Pacific (French Polynesia). 1 species (Davie 1993).

Cranaothus Ng, 1993 Diagnosis. Carapace subquadrate, 1.5 times as wide as long; regions weakly defined; front lobes produced well in advance of orbits, anterior margins sloping posteriorly; anterolateral margin longer than posterolateral margin. Male telson not reaching midlength of sternite 4. Maximum cl. 6 mm. Shelf (50 m). Central Indo-Pacific. 1 species (Ng 1993b).

Crosnierius Serène & Vadon, 1981 Diagnosis. Carapace subhexagonal, wider than long; regions weakly defined; anterolateral margin with prominent teeth. Cheliped palm upper margin with thin, smooth, evenly curved inwardly inclined crest. Pereopods 2–5 meri upper margins with upright spines; carpi and propodi upper margins with thin rounded crests. Maximum cl. 10 mm.

Euxanthus Dana, 1851 Diagnosis. Carapace as wide or wider than long; regions well defined; anterolateral margin with prominent teeth. Pereopods 2–5 usually folded into concavity of posterolateral margin by coaptation. Maximum cl. 38 mm. Intertidal–shelf (0–80 m). Indo-West Pacific, Temperate SW Australia. 6 species (Serène 1984: key to 6 species).

Guinotellus Serène, 1971 Diagnosis. Carapace narrowly ovate, little wider than long, domed, strongly convex; finely granular near margins, otherwise smooth, surface without trace of regions; anterolateral margin entire. Subhepatic cavity opening without concave ‘cut-out’ on anterolateral margins. Maximum cl. 41 mm (Pl. 66f). Intertidal, subtidal (0–20 m). Central Indo-Pacific. 1 species (Mendoza et al. 2009b: redescription).

770

Marine Decapod Crustacea

Hepatoporus Serène, 1984

Medaeops Guinot, 1967

Diagnosis. Carapace as wide or wider than long; anterolateral margin lobed between hepatic and branchial cavities. Subhepatic cavities upper part forming concave ‘cut-out’ on anterolateral margins. Maximum cl. 9 mm. Subtidal, shelf (2–85 m). Western and Central Indo-Pacific. 5 species (Mendoza and Ng 2008a: remarks, species comparison).

Diagnosis. Carapace subhexagonal, wider than long; regions well defined; anterolateral margin with 4 or 5 prominent or broad teeth. Endostomial ridges short, indicated proximally only, not extending anteriorly to epistome. Pereopod 5 merus less than 3.5 times as long as wide. Male pleonite 6 wider than long, usually appearing transversely rectangular. Gonopod 1 slender, curved, with lateral keel and long, subdistal setae. Maximum cl. 20 mm (Pl. 66h). Intertidal–shelf (0–236 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia (Kermadec Islands), Temperate SE South Africa. 6 species (Mendoza et al. 2009a: comment; Ng and McLay 2007: key to 6 species).

Hypocolpus Rathbun, 1897 Diagnosis. Carapace trapezoidal–broadly ovate, distinctly wider than long; surface rugose, regions indicated; anterolateral margin lobate or dentate. Subhepatic cavities openings without concave ‘cut-out’ on anterolateral margins. Maximum cl. 53 mm (Pl. 66e). Intertidal–shelf (0–150 m). Western and Central Indo-Pacific. 12 species (Crosnier 1991: species comparison; Serène 1984: key to 8 species).

Jacforus Ng & Clark, 2003 Diagnosis. Carapace 1.2–1.5 times as wide as long; regions well defined; front lobes weakly produced in advance of orbits, anterior margins transverse; anterolateral margin longer than posterolateral margin. Cheliped palm upper inner surface bluntly irregular, without crest. Male telson half width of pleonite 3, reaching beyond midlength of thoracic sternite 4. Maximum cl. 7 mm. Intertidal, subtidal (0–5 m). Indo-West Pacific. 1 species (Ng and Clark 2003).

Ladomedaeus Števčić, 2005 Diagnosis. Carapace subhexagonal, wider than long; regions well defined; anterolateral margin with 4 teeth. Male pleonites 3–5 with complete transverse grooves demarcating pleonites. Male pleonite 6 longitudinally rectangular. Maximum cl. 16 mm (Pl. 66g). Shelf, slope (50–712 m). Central Indo-Pacific. 2 species (Manuel-Santos and Ng 2007: rediagnosis).

Lipaesthesius Rathbun, 1898 Diagnosis. Carapace subhexagonal, about as wide as long; regions weakly defined; anterolateral margin entire. Antennal peduncle movable articles and flagellum not visible in anterior view, concealed within orbit. Pereopods 2–5 usually folded into concavity of posterolateral margin by coaptation. Maximum cl. 8.4 mm. Subtidal, shelf (5–73 m). Tropical Eastern Pacific. 1 species (Rathbun 1930: rediagnosis).

Lipkemedaeus Števčić, 2011 Diagnosis. Carapace ovate, ~1.4 times as wide as long; regions weakly defined; anterolateral margin with slender spines, often with secondary spinules basally. Maximum cl. 9 mm. Intertidal–shelf (0–134 m). Tropical Eastern Pacific. 2 species (Garth 1946; Rathbun 1930: figures as Medaeus spinulifer (Rathbun, 1898)).

Medaeus Dana, 1851 Diagnosis. Carapace much wider than long; region 4M distinct; regions 2R and 3R separated by groove; region P1 with transverse ridge; anterolateral margin with 3 lobes. Male telson reaching beyond midlength of sternite 4. Maximum cl. 25 mm (Pl. 66i). Subtidal–slope (15–367 m). Central and Eastern Indo-Pacific. 5 species.

Miersiella Guinot, 1967 Diagnosis. Carapace subhexagonal, wider than long; region 2M longitudinally divided for more than half length; region 4M distinct; anterolateral margin with 3 small teeth. Pereopod 5 merus 4 times as long as wide. Male pleonite 6 wider than long, usually appearing transversely rectangular. Gonopod 1 thick, weakly tapering, margins subparallel for most of distal two-thirds. Maximum cl. 9 mm. Shelf, slope (80–527 m). Western and Central Indo-Pacific, Temperate Australasia. 2 species (Guinot 1967a; Takeda 1989).

Monodaeus Guinot, 1967 Diagnosis. Carapace subhexagonal, wider than long; regions well defined; anterolateral margin with 4 teeth; anterolateral margin with last tooth directed laterally or anterolaterally; posterolateral surface without large conical tubercle, at most with low flat swelling. Male thoracic sternite 8 with small part visible between edge of pleonite 2 and condyle of pereopod 5 coxa. Male pleonite 6 wider than long, usually appearing transversely rectangular. Maximum cl. 30 mm. Subtidal–slope (6–800 m). Temperate Northern and Tropical E Atlantic, Indo-West Pacific, Temperate Southern Africa. 9 species (Guinot and Macpherson 1988).

Olenothus Ng, 2002 Diagnosis. Carapace ovate, 1.7 times as wide as long; regions weakly defined; anterolateral margin curved, irregular or evenly granular. Pereopods 2–5 folding into concavity of posterolateral

14 – Brachyura – crabs

margin by coaptation. Male pleonite 6 slightly wider than long, appearing square. Maximum cl. 14 mm. Subtidal, shelf (1–23 m). Central Indo-Pacific (Guam). 1 species (Ng 2002a).

Palatigum Davie, 1997 Diagnosis. Carapace ovate, ~1.4 times as wide as long; regions well defined; anterolateral margin with prominent teeth. Maxilliped 3 entirely and densely covered with short setae. Maximum cl. 10 mm. Slope (234–242 m). Central Indo-Pacific (New Caledonia). 1 species (Davie 1997).

Paramedaeus Guinot, 1967 Diagnosis. Carapace ovate, ~1.4 times as wide as long; region 4M not indicated; anterolateral margin with prominent teeth, as long posterolateral margin. Male pleonite 6 longitudinally rectangular; telson not reaching midlength of sternite 4. Maximum cl. 9 mm. Intertidal–slope (0–288 m). Indo-West Pacific. 5 species (Davie 1997: key to 5 species).

Paraxanthodes Guinot, 1968 Diagnosis. Carapace subhexagonal, wider than long; regions well defined; anterolateral margin with 4 teeth. Endostomial ridges complete, extending anteriorly to epistome. Pereopod 5 merus 2.5 times as long as wide. Male pleonite 6 wider than long, usually appearing transversely rectangular. Gonopod 1 slender, distally curved, with short subdistal spines. Maximum cl. 30 mm. Intertidal–shelf (0–210 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 3 species (Davie 1993: rediagnosis, species comparison).

771

Psaumis Kossmann, 1877 Diagnosis. Carapace much wider than long; regions well demarcated, eroded; anterolateral margin with broad teeth. Male thoracic sternum with large, rounded, basin-like cavity anterior to telson. Pereopods 2–5 folding into concavity of posterolateral margin by coaptation; carpi and propodi with prominent, strongly curved crest, hollowed on anterior face forming shallow ‘cup’. Maximum cl. 8.4 mm (Pl. 66j). Intertidal–shelf (0–25 m). Indo-West Pacific. 2 species (Serène 1984).

Rizalthus Mendoza & Ng, 2008 Diagnosis. Carapace subhexagonal, about as wide as long; regions well defined, densely covered with large granules; anterolateral margin curved, irregular or evenly granular. Pereopods 2–5 usually folding into concavity of posterolateral margin by coaptation. Male pleonite 6 longitudinally rectangular. Maximum cl. 8 mm. Subtidal, shelf (3–37 m). Central Indo-Pacific (Philippines). 1 species (Mendoza and Ng 2008a).

Takedax Mendoza & Ng, 2012 Diagnosis. Carapace subhexagonal, wider than long; regions well defined; anterolateral margin with 4 teeth; anterolateral margin with last tooth directed posterolaterally; surface just inward of posterolateral corner with large conical tubercle. Male thoracic sternite 8 with small part visible between edge of pleonite 2 and condyle of pereopod 5 coxa. Male pleonite 6 wider than long, usually appearing transversely rectangular. Maximum cl. 11 mm. Shelf, slope (50–356 m). Western and Central Indo-Pacific (SW India, Philippines). 1 species (Mendoza and Ng 2012).

Visayax Mendoza & Ng, 2008 Pilomedaeus Takeda & Komatsu, 2011 Diagnosis. Carapace ovate, ~1.4 times as wide as long; densely covered by short, dense tomentum obscuring regions and granulation; anterolateral margin with prominent teeth. Male pleonites 3–5 with complete transverse grooves demarcating pleonites. Maximum cl. 11 mm. Shelf (115–211 m). Temperate Northern Pacific (Izu Is, Japan). 1 species (Takeda and Komatsu 2011).

Diagnosis. Carapace subhexagonal, about as wide as long; regions well demarcated, eroded; anterolateral margin curved, irregular or evenly granular. Male thoracic sternum with large, rounded, basin-like cavity anterior to telson. Pereopods 2–5 folding into concavity of posterolateral margin by coaptation; smooth, with few low ridges and granules or deeply eroded and pitted. Maximum cl. 5.3 mm. Subtidal, shelf (3–25 m). Central Indo-Pacific (Philippines). 2 species (Mendoza and Ng 2008a).

Pleurocolpus Crosnier, 1995 Diagnosis. Carapace 1.45 times as wide as long; regions well demarcated, eroded; anterolateral margin with 3 lobes. Carapace posterolateral margin with prominent triangular projection near midlength. Chelipeds and pereopods 2–5, when folded against carapace, coapting to form deep tube-like hollow bounded by posterior margin of cheliped, upper margins of pereopods 2–5 meri and posterolateral margin of carapace. Maximum cl. 21 mm. Shelf (190 m). Eastern Indo-Pacific (Tuamotu). 1 species (Crosnier 1995).

Subfamily Garthiellinae Mendoza & ManuelSantos, 2012 Garthiellinae includes two genera, Mabui and Garthiella, whose species inhabit coarse coral and sandy sediments and rocky environments from the shore down to shallow shelf depths in the Pacific. The two genera differ strongly in external ornamentation but share the comparatively wide thoracic sternum and simple rather than hollowed cheliped

772

Marine Decapod Crustacea

Fig. 14.159.  Xanthidae, Garthiellinae. a, Garthiella sitatuna Mendoza & Manuel-Santos, 2012. b, Mabui calculus Naruse, Maenosono &.Ng, 2021. c, male thoracic sternites, 1–4, pleonite 6, telson, Garthiella. d, right and left gonopod 1, Mabui.

finger tips. Garthiella was found not to belong in Chlorodiellinae where it had been placed (Lai et al. 2011) and a new subfamily was erected for it (Mendoza and Manuel-Santos 2012). Phylogenetic analysis of Mabui also placed it together with Garthiella in Garthiellinae (Naruse et al. 2021). Diagnosis. Carapace up to 1.7 times as wide as long; anterolateral margin unarmed or with sharp, curved spines. Male thoracic sternites 3–4 length slightly less than half maximum width. Antennules folding transversely. Chelipeds with finger tips acute, not hollowed, crossing; pereopods 2–5 meri upper margins smooth or spinose; pereopods 2–5 without dactylo-propodal ‘lock’. Garthiella Titgen, 1986 Diagnosis. Carapace transversely hexagonal, up to 1.7 times as wide as long; anterolateral teeth as sharp, curved spines. Cheliped palms coarsely granular or spinose; dactylus of larger cheliped without prominent, basal cutting tooth. Pereopods 2–5 merus upper margin spinose. Gonopods and female gonopores bilaterally symmetrical. Maximum cl. 7 mm. Intertidal–shelf (0–48 m). Central and Eastern Indo-Pacific. 2 species (Mendoza and Manuel-Santos 2012: rediagnosis, species descriptions; Titgen 1986).

Mabui Naruse, Maenosono & Ng, 2021 Diagnosis. Carapace transversely ovate, 1.6–1.7 times as wide as long; anterolateral teeth absent or indicated at most by 2 or 3 inconspicuous notches. Cheliped palms smooth or finely granular; dactylus of larger cheliped with prominent, rounded, basal cutting tooth. Pereopods 2–5 merus upper margin smooth. Male gonopods longer on right side. Female gonopore larger on left side. Maximum cl. 5 mm (Pl. 66p). Intertidal, subtidal (0–3 m). Central Indo-Pacific (Okinawa). 1 species (Naruse et al. 2021).

Subfamily Glyptoxanthinae Mendoza & Guinot, 2011 Glyptoxanthus is the only genus. Its species inhabit rocky environments down to mid-shelf depths on both sides of the tropical Atlantic, and the Red Sea (Mendoza and Guinot 2011). The dorsal carapace is typically ornamented with prominent ridges formed by fused granules, arranged in regular reticulate patterns. The front has four lobes, allowing free movement of antennules. Diagnosis. Carapace front distally downcurved, appearing 4-lobed in anterior view; anterolateral margin descending anteriorly, not meeting exorbital angle but continuing below orbit towards anterolateral corner of buccal frame; subhepatic surface with, more-or less vertical grooves descending from gaps between anterolateral teeth (Fig. 14.160). Antennules folding transversely. Chelipeds with finger tips acute or expanded, spoon-shaped; pereopods 2–5 without dactylopropodal ‘lock’. Glyptoxanthus A. Milne-Edwards, 1879 Maximum cl. 36 mm. Intertidal–shelf (0–90 m). Tropical Atlantic, Western Indo-Pacific (Red Sea), Tropical Eastern Pacific. 9 species (Mendoza and Guinot 2011: diagnosis, key to species).

Subfamily Kraussiinae Ng, 1993 Kraussiines are round or oval domed crabs with flattened legs used for burrowing in sandy sediments around coral reefs (Serène 1972). All except one of the several species described as members of Kraussia (as in the key of Serène 1972) are now recognised as belonging to Palapedia (Ng 1993a).

14 – Brachyura – crabs

773

Fig. 14.160.  Xanthidae, Glyptoxanthinae, Glyptoxanthus hancocki Garth, 1939. a, dorsal; b, anterior; c, ventral views.

Diagnosis. Carapace transversely ovate. Antennules folding obliquely forwards, or longitudinally. Chelipeds with finger tips acute or expanded, spoon-shaped;

pereopods 2–5 short, laterally compressed, especially distal articles; dactyli blade-like; pereopods 2–5 without dactylo-propodal ‘lock’.

Key to genera of Kraussiinae 1. – 2. –

Carapace oval, ~1.3 times as wide as long (Fig. 14.161a). Pereopod 5 elongate, merus 1.8–2.7 times as long as wide (Fig. 14.161a,g)�����������������������������������������������������������������������������������������������������������������������������������������������Garthasia Carapace subcircular, about as wide as or little wider than long (Fig. 14.161b, c). Pereopod 5 stout, merus ~1.6 times as long as wide (Fig. 14.161c, h)������������������������������������������������������������������������������������������������������������������������������2 Carapace anterolateral margin dentiform or uneven (Fig. 14.161c). Cheliped fingers with slightly flattened or sharp tips����������������������������������������������������������������������������������������������������������������������������������������������������������������� Palapedia Carapace anterolateral margin with 4 sharp epibranchial teeth (Fig. 14.161b). Cheliped fingers with spoon-shaped tips (Fig. 14.161f)���������������������������������������������������������������������������������������������������������������������������������� Kraussia Garthasia Ng, 1993

Diagnosis. Carapace oval, ~1.3 times as wide as long; anterolateral margin shorter than posterolateral margin, smooth. Maxilliped 3

rectangular, ~2.4 times as long as wide, carpus not swollen. Cheliped fingers with sharp tips. Pereopod 5 elongate, merus ~1.8–2.7 times as long as wide. Maximum cl. 11 mm.

Fig. 14.161.  Xanthidae, Kraussiinae. a, Garthasia americana (Garth, 1939); b, carapace, Kraussia; c, carapace, pereopod 5, Palapedia. Maxilliped 3: d, Garthasia; e, Kraussia. f, chela, Kraussia. Pereopod 5: g, Garthasia; h, Kraussia.

774

Marine Decapod Crustacea

Subtidal, shelf (2–73). Central Indo-Pacific, Tropical Eastern Pacific. 2 species (Garth 1939: species description as Kraussia; Hsueh 2020: rediagnosis, second species; Ng 1993a: diagnosis).

Intertidal–shelf (0–128 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 17 species (Ng 1993a: diagnosis; Serène 1972: descriptions and key to 10 species as Kraussia).

Kraussia Dana, 1852 Diagnosis. Carapace circular, about as wide as long; anterolateral margin longer than posterolateral margin, with 4 sharp epibranchial teeth and spinules. Maxilliped 3 elongate, ~3 times as long as wide, carpus swollen. Cheliped fingers with spoon-shaped tips. Pereopod 5 stout, merus ~1.6 times as long as wide. Maximum cl. 17 mm. Intertidal, shallow subtidal. Temperate Northern W Pacific, Central Indo-Pacific, Temperate Southern Africa. 1 species (Ng 1993a: rediagnosis, description of only species).

Palapedia Ng, 1993 Diagnosis. Carapace subcircular, as wide as or little wider than long; anterolateral margin longer than posterolateral margin, dentiform or uneven. Maxilliped 3 elongate, ~3 times as long as wide, carpus swollen. Cheliped fingers with slightly flattened or sharp tips. Pereopod 5 stout, merus ~1.6 times as long as wide. Maximum cl. 20 mm (Pl. 66l).

Subfamily Liomerinae Sakai, 1976 Liomerine xanthids tend to have a less ornamented and more evenly ovate carapace than other xanthids, and are often broader. They have spoon-shaped fingers lacking significant teeth. They are often brightly coloured, including vivid red. Despite there being only six genera, the subfamily may be paraphyletic (Lai et al. 2011). Diagnosis. Carapace transversely ovate or hexagonal, up to twice as wide as long or more; anterolateral margin unarmed or with blunt lobes, not sharp spines. Male thoracic sternites 3–4 length equal to or slightly less than half maximum width. Antennules folding transversely. Chelipeds with finger tips hollowed on occlusal margin, not crossing; pereopods 2–5 with or without dactylo-propodal ‘lock’.

Fig. 14.162.  Xanthidae, Liomerinae. a, Actiomera erythra (Lanchester, 1902); b, Bruciana pediger (Alcock, 1898); c, Liomera monticulosa (A. Milne-Edwards, 1873); d, Lipkemera iejima Naruse & Fujita, 2015; e, Neoliomera insularis (Adams & White, 1849); f, Paraliomera longimana (A. Milne-Edwards, 1865). g, thoracic sternites 1–4, telson, Neoliomera. h, pereopod 5, propodus, dactylus, Neoliomera.

14 – Brachyura – crabs

775

Key to genera of Liomerinae 1. – 2. – 3. – 4. – 5. –

Carapace regions usually poorly defined; surface little flattened transversely (Fig. 14.162e, f). Pereopods 2–5 with dactylo-propodal ‘lock’ (Fig. 14.162h)���������������������������������������������������������������������������������������������������������������������2 Carapace regions well defined; surface strongly convex longitudinally and transversely. Pereopods 2–5 without dactylo-propodal ‘lock’�����������������������������������������������������������������������������������������������������������������������������������������������3 Carapace surface smooth. Chelipeds markedly unequal (Fig. 14.162f)��������������������������������������������������������������� Paraliomera Carapace surface rugose, granular. Chelipeds equal (Fig. 14.162e)����������������������������������������������������������������������� Neoliomera Carapace front projecting anteriorly beyond orbits and breaking general outline of anterior margin. Surface of carapace, chelipeds, thoracic sternum and pleon finely granular and conspicuously punctate, forming reticulated texture. Pereopods 2–5 flattened, upper and lower margins cristate (Fig. 14.162a)�����������������������Actiomera Carapace front not projecting, not breaking general curve of anterior margin. Carapace, chelipeds, sternal plastron and abdomen smooth, granular or finely punctate but never forming reticulated texture. Pereopods 2–5 without cristiform carinae on upper and lower margins���������������������������������������������������������������������������4 Pereopods 2–5 short, stout; pereopod 5 merus less than 3 times as long as wide (Fig. 14.162c)����������������������������� Liomera Pereopods 2–5 elongate, slender; pereopod 5 merus more than 3 times as long as wide�����������������������������������������������������5 Carapace regions distinct, well separated by narrow grooves (Fig. 14.162b). Antennal basal article inclined less than 30° from midline������������������������������������������������������������������������������������������������������������������������������������������ Bruciana Carapace regionalization incomplete, at most indicated on anterior half (Fig. 14.162d). Antennal basal article inclined ~45° from midline�������������������������������������������������������������������������������������������������������������������������� Lipkemera Actiomera Ng, Guinot & Davie, 2008

Liomera Dana, 1851

Diagnosis. Carapace well defined; surface strongly convex longitudinally and transversely; surface of carapace, chelipeds, thoracic sternum and pleon finely granular and conspicuously punctate, forming reticulated texture; front projecting anteriorly beyond orbits and breaking general outline of anterior margin. Chelipeds subequal. Pereopods 2–5 flattened, upper and lower margins cristate; without dactylo-propodal ‘lock’. Maximum cl. 8 mm. Shelf (20–100 m). Western and Central Indo-Pacific. 3 species (Serène 1984: key to 2 species as Liomera (Actites), figures). Actiomera is a replacement name for Actites Lanchester, 1902.

Diagnosis. Carapace well defined; surface strongly convex longitudinally and transversely; front not projecting. Chelipeds subequal. Pereopods 2–5 short, stout; without dactylo-propodal ‘lock’; pereopod 5 merus less than 3 times as long as wide. Maximum cl. 32 mm (Pl. 66n). Intertidal–shelf (0–144 m). Indo-West Pacific, Temperate SE South Africa, Temperate Australia. 30 species (Serène 1984: key to c. 20 species as Liomera (Liomera)).

Bruciana Serène, 1977 Diagnosis. Carapace regions distinct, well separated by narrow grooves; front not projecting. Antennal basal article inclined less than 30° from midline. Chelipeds subequal. Pereopods 2–5 short, stout, without dactylo-propodal ‘lock’; pereopod 5 merus less than 3 times as long as wide. Maximum cl. 6.5 mm (Pl. 66m). Subtidal, shelf (10–40 m). Western and Central Indo-Pacific. 1 species (Serène 1984: only species as Liomera (Bruciana)).

Neoliomera Odhner, 1925 Diagnosis. Carapace regions poorly defined, rugose, granular; front slightly projecting. Chelipeds equal. Pereopods 2–5 short, stout, with dactylo-propodal ‘lock’; pereopod 5 merus less than 3 times as long as wide. Maximum cl. 24 mm (Pl. 66n). Intertidal–shelf (0–50 m), including in submarine caves. IndoWest Pacific, Temperate SE South Africa. 19 species (Ng 2002b; Poupin and Starmer 2013; Serène 1984: key to 13 species).

Lipkemera Davie, 2010 Diagnosis. Carapace regionalization incomplete, at most indicated on anterior half; front not projecting. Antennal basal article inclined ~45° from midline. Chelipeds subequal. Pereopods 2–5 short, stout; without dactylo-propodal ‘lock’; pereopod 5 merus less than 3 times as long as wide. Maximum cl. 24 mm. Subtidal–slope (17–270 m, including in submarine caves). Central and Eastern Indo-Pacific (Philippines, Japan, French Polynesia). 5 species (Davie 1993: diagnosis as Meriola, superseded name; Naruse and Fujita 2015: key to species).

Paraliomera Rathbun, 1930 Diagnosis. Carapace regions poorly defined, smooth; front not projecting. Chelipeds strongly unequal. Pereopods 2–5 short, stout; with dactylo-propodal ‘lock’; pereopod 5 merus less than 3 times as long as wide. Maximum cl. 8 mm. Intertidal–shelf (0–154 m). Tropical W Atlantic. 2 species (Rathbun 1930).

776

Marine Decapod Crustacea

Subfamily Polydectinae Dana, 1851 boxer crabs Boxer crabs are inhabitants of coral reefs in the Indo-West Pacific notable for carrying one or two sea anemones, or sometimes nudibranchs in their small chelipeds. The

fingers of the chelae are armed with sharp teeth. The anemones probably catch food for the crab and are used in fighting between individuals (Duerden 1905; Karplus et  al. 1998). The relationships between two genera has been long understood; a third more recently added (Guinot 1976; Mendoza and Ng 2011).

Key to genera of Polydectinae 1. – 2. –

Carapace and pereopods densely tomentose. Carapace lateral margin strongly convex, smooth (Fig. 14.163c)���������������������������������������������������������������������������������������������������������������������������������������������������������������� Polydectus Carapace and pereopods weakly setose. Carapace lateral margin slightly convex, or anterolateral and posterolateral margins discontinuous (Fig. 14.163a, b d)�����������������������������������������������������������������������������������������������������2 Carapace subhexagonal, anterolateral and posterolateral margins discontinuous. Pereopod 2 more slender and shorter than pereopods 3, 4 (Fig. 14.163a, b)����������������������������������������������������������������������������������������������������������� Lybia Carapace subovate, anterolateral and posterolateral margins continuous, denticulate. Pereopod 2 similar to pereopods 3, 4 (Fig. 14.163d)���������������������������������������������������������������������������������������������������������������������������������������� Tunebia Lybia H. Milne Edwards, 1834

Tunebia Mendoza & Ng, 2011

Diagnosis. Carapace subhexagonal, smooth; anterolateral margin distinct from posterolateral margin, both smooth. Pereopod 2 more slender and shorter than pereopods 3, 4. Maximum cl. 10 mm (Pl. 66k). Intertidal–shelf (0–90 m; coral reefs). Indo-West Pacific, Temperate Southern Africa, Temperate Australia. 8 species (Guinot 1976: species descriptions; Serène 1984: key to 6 species).

Diagnosis. Carapace subovate, smooth; anterolateral margin continuous with posterolateral margin, both denticulate. Pereopod 2 similar to pereopods 3, 4. Maximum cl. 4 mm. Shelf (74–137 m). Temperate Northern W Pacific, Central Indo-Pacific. 2 species. At least one species carries nudibranchs as well as sea anemones (Mendoza and Ng 2011).

Polydectus H. Milne Edwards, 1837 Diagnosis. Carapace ovate, heavily tomentose, extending on to all pereopods; anterolateral margin continuous with posterolateral margin, both smooth under setae. Pereopod 2 similar to pereopods 3, 4. Maximum cl. 12 mm. Intertidal, shallow subtidal (0–6 m; under coral rubble). IndoWest Pacific. 1 species (Guinot 1976: ecology, description).

Subfamily Xanthinae MacLeay, 1838 Xanthines are generally less prominently ornamented than other xanthids, typically having lesser defined carapace regions without dense carapace setation or spination, comparatively elongated thoracic sternites 3–4, and an unspecialised dactylo-propodal articulation on pereopods 2–5, lacking the ‘lock’. As such, xanthines have the most ‘generalised’

Fig. 14.163.  Xanthidae, Polydectinae. Carapace, pereopods: a, b, Lybia spp.; c, Polydectus. d, Tunebia hatagumoana (Sakai, 1961). Chela: e, Lybia; f, Polydectus; g, Tunebia.

14 – Brachyura – crabs

morphology of the xanthids, and not surprisingly, the subfamily is probably not monophyletic (Lai et al. 2011). Xanthinae includes some of the most common tropical shore crabs, such as Leptodius and Macromedaeus, as well as some that are highly toxic to humans if consumed [e.g. Demania; (Garth and Alcala 1977; Yasumoto et  al. 1983)]. Most xanthines occur on shallow temperate and tropical reefs but some reach slope depths. The composition of Xanthinae employed here largely follows Ng et al. (2008) with the following changes: Eucratodes, Garthiope and Gonopanope, having wide thoracic sternites 3–4, are included in Pseudorhombilidae; Nectopanope is considered a euryplacid following Ng et al. (2019); Jacforus is moved to Euxanthinae because the anterolateral carapace margins descend below the orbits instead of meeting the exorbital angle (Ng and Clark 2003); Paraxanthodes is included in Euxanthinae following Mendoza and Ng (2010);

777

and Ectaesthesius is tentatively transferred to Trapeziidae owing to its similarity to other trapeziids. Diagnosis. Carapace regions not or well defined, usually smooth or minimally granular, glabrous or with scattered setae, occasionally densely granulate, finely, evenly setose; when present, surface granulation usually confined largely to anterior half, rarely over entire surface. Male thoracic sternites 3–4 length greater than half maximum width. Antennules folding transversely. Cheliped and pereopods 2–5 usually smooth or weakly granular, sometimes densely tuberculate or spinous; chelipeds with finger tips acute or expanded, spoonshaped; pereopods 2–5 without dactylo-propodal ‘lock’. Implicit generic characters. Front not extending prominently beyond supraorbital angles; eyes, when extended forwards, close to level of frontal margin. Chelipeds unequal, often grossly so; larger cheliped finger tips pointed, partially crossing when closed. Male pleonites 3–5 fused.

Key to genera of Xanthinae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. – 9. –

Carapace front extending prominently beyond supraorbital angles; eyes, when extended forwards, well behind level of frontal margin (e.g. Fig. 14.165e, f)����������������������������������������������������������������������������������������������������������������2 Carapace front not extending prominently beyond supraorbital angles; eyes, when extended forwards, close to level of frontal margin��������������������������������������������������������������������������������������������������������������������������������������������������8 Carapace front quadrate or approximately rounded, medially at most with subtle shallow concavity, without median fissure (Fig. 14.165f)����������������������������������������������������������������������������������������������������������������� Megametope Carapace front variously shaped, with distinct median fissure������������������������������������������������������������������������������������������������3 Pereopods 2–5 slender, pereopod 5 merus more than twice as long as wide. Male pleonite 6 about as long as wide, appearing square (Fig. 14.167k)��������������������������������������������������������������������������������������������������������������������������������������4 Pereopods 2–5 stout, pereopod 5 merus less than twice as wide as long. Male pleonite 6 longer than wide, longitudinally rectangular���������������������������������������������������������������������������������������������������������������������������������������������������������7 Carapace regions and margins of anterolateral teeth coarsely granular (Fig. 14.165e)����������������������������������������� Marratha Carapace regions punctate or not indicated, smooth; margins of anterolateral teeth smooth��������������������������������������������5 Carapace anterolateral margin behind exorbital angle with 4 well developed teeth: anterior 3 large, triangular, projecting, separated by large V-shaped incisions; fourth tooth small, acute (Fig. 14.165g)���������������������������������������������������������������������������������������������������������������������������������������������������������Metaxanthops Carapace anterolateral margins behind exorbital angles with 4 or 5 low teeth, rounded or obtusely angular lobes��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������6 Carapace regions smooth or rugose, with or without regions indicated. Cheliped carpus pitted and irregular, palm surface with irregular longitudinal ridges (Fig. 14.165k)���������������������������������������������Neoxanthops Carapace surface smooth, regions not indicated. Cheliped carpus and palm surface smooth, without ridges, tubercles or rugosity (Fig. 14.166l)�����������������������������������������������������������������������������������������������������Wardoxanthops Carapace with 3 small anterolateral teeth behind exorbital angle (Fig. 14.166b)���������������������������������������������� Paraxanthus Carapace with 8 prominent anterolateral teeth behind exorbital angle (Fig. 14.166g)��������������������������������� Cycloxanthops Larger cheliped finger tips rounded, hollowed on occlusal surface, meeting when closed (Fig. 14.167l)����������������������������9 Larger cheliped finger tips pointed, partially crossing when closed��������������������������������������������������������������������������������������13 Cheliped finger tips rounded and hollowed but not expanded laterally (Fig. 14.165d). Antennal basal article oriented almost parallel to carapace midline (Fig. 14.167a)�����������������������������������������������������������������������������������������������10 Cheliped finger tips laterally expanded, hoof or spoon-like (Fig. 14.165a). Antennal basal article oriented obliquely to carapace midline (Fig. 14.167b, c)���������������������������������������������������������������������������������������������������������������������11

778

Marine Decapod Crustacea

10.

Carapace with 3 low anterolateral teeth (excluding exorbital angle), all placed near lateral extremities (Fig. 14.166i)��������������������������������������������������������������������������������������������������������������������������������������������������������� Gaudichaudia – Carapace with 4 or more prominent anterolateral teeth (excluding exorbital angle) evenly spaced along margin (Fig. 14.165d) or 2 small teeth at lateral extremities (M. demani (Odhner, 1925))������������Macromedaeus 11. Gonopod 1 with broad petaloid tip; subdistal margin with dense brush of long setae (Fig. 14.167u)���������������Xanthodius – Gonopod 1 with tapering tip; subdistal margin lined with short stiff setae, no longer than adjacent gonopod width (Fig. 14.167o, s)���������������������������������������������������������������������������������������������������������������������������������������������� 12

Fig. 14.164.  Xanthidae, Xanthinae. a, Aldrovandiopanope taylori (Garth, 1986); b, Aristotelopanope ashcrafti (Garth, 1986); c, Bottoxanthodes insculptus (Stimpson, 1871); d, Camilohelleria manteri (Garth, 1986); e, Cataleptodius floridanus (Gibbes, 1850); f, Coralliope parvula (A. Milne-Edwards, 1869); g, Demania reynaudii (H. Milne Edwards, 1834); h, Euryxanthops cepros Davie, 1997; i, Guitonia leimomi Lasley, Mendoza & Ng, 2010; j, Juxtaxanthias lividus (Latreille in Milbert, 1812); k, Lachnopodus subacutus (Stimpson, 1858).

14 – Brachyura – crabs

779

12. Male pleonite 6 as long as or longer than wide. Gonopod 1 with minute, subdistal spines and elongated apical lobe, bordered by anchor-shaped projections (Fig. 14.167s)�����������������������������������������������������������������������Leptodius – Male pleonite 6 wider than long (Fig. 14.167j). Gonopod 1 with row of long subdistal setae on one margin and some minute subdistal spines on other (Fig. 14.167o)�������������������������������������������������������������� Cataleptodius 13. Carapace with 9 subequal anterolateral teeth behind exorbital angle (Fig. 14.166d)���������������������������� Williamstimpsonia – Carapace with no more than 5 anterolateral teeth behind exorbital angle���������������������������������������������������������������������������14 14. Carapace branchial regions strongly inflated, bulging, smooth (Fig. 14.165l)���������������������������������������������Orphnoxanthus

Fig. 14.165.  Xanthidae, Xanthinae. a, Leptodius affinis (De Haan, 1835); b, Liagore rubromaculata (De Haan, 1835); c, Lioxanthodes alcocki Calman, 1909; d, Macromedaeus crassimanus (A. Milne-Edwards, 1867); e, Marratha angusta (Rathbun, 1906); f, Megametope rotundifrons (H. Milne Edwards, 1834); g, Metaxanthops acutus Serène, 1984; h, Microcassiope xantusii (Stimpson, 1871); i, Nanocassiope alcocki (Rathbun, 1902); j, Neoxanthias impressus (Latreille in Milbert, 1812); k, Neoxanthops quadrilobatus (Sakai, 1939); l, Orphnoxanthus microps (Alcock & Anderson, 1894);

780

– 15.

– 16. – 17. –

Marine Decapod Crustacea

Carapace branchial regions flat or gently convex, not inflated, surface smooth, granular or grooved���������������������������15 Carapace branchial and gastric regions separated by deep narrow grooves, anterior part of mesogastric triangle distinct, short median groove from tip of mesogastric triangle extending to frontal margin and crossing transverse postfrontal groove; postfrontal groove curving laterally behind orbit and continuing parallel to anterolateral margin, terminating near last anterolateral tooth; regions in posterior half of carapace not indicated (Fig. 14.166c)���������������������������������������������������������������������������������Pestoxanthodes Carapace regionalisation, if present, not as above��������������������������������������������������������������������������������������������������������������������16 Male chelipeds subequal or feebly unequal. Gonopod 1 elongate, slightly curved, with row of long subdistal setae on one margin and some short subdistal spines on other (Fig. 14.167q, r, t) (except Demania splendida Laurie, 1906)������������������������������������������������������������������������������������������������������������������������������������������������������������17 Male chelipeds subequal or distinctly unequal and sometimes strongly dissimilar. Gonopod 1 otherwise��������������������� 23 Carapace regions generally weakly indicated to obsolete over entire surface, or weakly indicated in anterior half, finely granular or smooth�����������������������������������������������������������������������������������������������������������������������������������������������18 Carapace regions well demarcated over almost entire surface, surface smooth to coarsely granular������������������������������21

Fig. 14.166.  Xanthidae, Xanthinae. a, Paraxanthias notatus (Dana, 1852); b, Paraxanthus barbiger (Poeppig, 1836); c, Pestoxanthodes sulcatus (Faxon, 1893); d, Williamstimpsonia denticulatus (White, 1848); e, Pseudomedaeus africanus (Monod, 1956); f, Xanthias lamarckii (H. Milne Edwards, 1834). Carapace: g, Cycloxanthops; h, Epixanthops casellatoi Serène, 1984; i, Gaudichaudia; j, Neolioxantho; k, Ovatis; l, Wardoxanthops; m, Xantho; n, Xanthodius.

14 – Brachyura – crabs

18. – 19. – 20. – 21. – 22. – 23. – 24. – 25. – 26. – 27. – 28. – 29. – 30. – 31. – 32. –

781

Carapace anterolateral teeth unequal and dissimilar: posterior one or two teeth pointed and angular, preceding anterolateral teeth rounded����������������������������������������������������������������������������������������������������������������������������������19 Carapace anterolateral teeth absent or with similar apices as pointed, angular or rounded lobes���������������������������������� 20 Carapace posterior 1–3 anterolateral teeth pointed, preceding tooth/teeth rounded, prominent. Pereopods 2–5 setose (Fig. 14.164j)�������������������������������������������������������������������������������������������������������������������Juxtaxanthias Carapace posterior two anterolateral teeth bluntly angular, anterior two teeth low rounded. Pereopods 2–5 glabrous (Fig. 14.166j)���������������������������������������������������������������������������������������������������������������� Neolioxantho Carapace front slightly anterior to anterolateral margins, breaking general outline of carapace. Pereopods 2–5 meri, carpi and propodi upper margins sharply crested (Fig. 14.164h)��������������������������� Euryxanthops Carapace front more-or-less in line with anterolateral margins, not breaking general outline of carapace. Pereopods 2–5 meri, carpi and propodi upper margins bluntly rounded in cross-section, without crest (Fig. 14.166f)�������������������������������������������������������������������������������������������������������������������������������������������������������������������Xanthias Carapace regions upraised, projecting, areolate, smooth or pitted giving eroded aspect (Fig. 14.165j)����������� Neoxanthias Carapace regions well demarcated, but not projecting and areolate, usually covered in low nodules, small tubercles or low rugosities������������������������������������������������������������������������������������������������������������������������������������������������������ 22 Carapace 1.1–1.5 times as wide as long; front unimarginate (Fig. 14.164g)������������������������������������������������������������� Demania Carapace 1.6–1.7 times as wide as long; front bimarginate (Fig. 14.166e)�����������������������������������������������������Pseudomedaeus Carapace regions mostly well defined, surface smooth or granular; with 3 or more anterolateral teeth behind exorbital angle������������������������������������������������������������������������������������������������������������������������������������������������������������� 24 Carapace regions weakly defined, surface smooth; with 4 or fewer anterolateral teeth behind exorbital angle�������������� 33 Cheliped palm smooth or weakly eroded, not granular or spinose������������������������������������������������������������������������������Xantho Cheliped palm granular and/or spinose������������������������������������������������������������������������������������������������������������������������������������ 25 Carapace anterolateral margin with posterior two teeth stronger and more acute than anterior two; regions and front smooth (Fig. 14.166b)�������������������������������������������������������������������������������������������������������������������������� Paraxanthias Carapace with last anterolateral tooth smaller than preceding two teeth; regions regularly granular, at least in anterior half of carapace����������������������������������������������������������������������������������������������������������������������������������������������������� 26 Carapace regions indicated anteriorly, obsolete posteriorly, surface granular on anterior half, posterior half smooth��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 27 Carapace regions indicated anteriorly and posteriorly, all granular��������������������������������������������������������������������������������������31 Maxilliped 3 exopod outer margin serrated (Fig. 14.167d)����������������������������������������������������������������������������������������������������� 28 Maxilliped 3 exopod outer margin smooth������������������������������������������������������������������������������������������������������������������������������ 29 Maxilliped 3 merus distolateral margin produced (Fig. 14.167d). Male pleonites 4–5 fused, pleonite 3 free (Fig. 14.167h). Gonopod 1 tip dilated, distinctly bent laterally (Fig. 14.167n)�������������������������������������������Camilohelleria Maxilliped 3 merus distolateral margin rounded, not produced. Male pleonites 3–5 fused (Fig. 14.167i). Gonopod 1 tip tapering, not bent laterally (Fig. 14.167p)�������������������������������������������������������������������������������������Coralliope Pereopod 5 merus 4 times as long as wide (Fig. 14.164i)���������������������������������������������������������������������������������������������� Guitonia Pereopod 5 merus 3 times or less as long as wide�������������������������������������������������������������������������������������������������������������������� 30 Carapace granulation arranged into numerous transverse or oblique rows (Fig. 14.165h)��������������������������� Microcassiope Carapace granulation regular or with only few transverse or oblique granular rows (Fig. 14.165i)�������������Nanocassiope Carapace surface glabrous. Pereopod 5 shorter than carapace width; merus upper margin unarmed (Fig. 14.164c)������������������������������������������������������������������������������������������������������������������������������������������������������ Bottoxanthodes Carapace surface setose. Pereopod 5 longer than carapace width; merus upper margin dentate or spinous�����������������32 Carapace frontal lobes straight, bimarginate, margins serrated. Pereopods 2–4 meri, carpi and propodi upper margins spinose (Fig. 14.164a). Male pleonite 6 wider proximally than distally (Fig. 14.167f)������������������������������������������������������������������������������������������������������������������������������������������������ Aldrovandiopanope Carapace frontal lobes slightly curved or sinuous, unimarginate, margins minutely granular. Pereopods 2–4 meri, carpi and propodi upper margins with acute granules (Fig. 14.164b). Male pleonite 6 wider distally than proximally (Fig. 14.167g)��������������������������������������������������������������������������������������������������������Aristotelopanope

782

Marine Decapod Crustacea

Fig. 14.167.  Xanthidae, Xanthinae. Antennule, antenna, orbit (ventral view): a, Cycloxanthops; b, Xanthodius. c, Leptodius. d, maxilliped 3, Camilohelleria. e, male sternites 1–4, telson, Leptodius. Male pleon: f, Aldrovandiopanope; g, Aristotelopanope; h, Camilohelleria; i, Coralliope. Male pleonite 6, telson: j, Cataleptodius; k, Marratha. Chela fingers: l, Cataleptodius (ventral view); m, Lioxanthodes (outer view, minor). Gonopod 1 apex: n, Camilohelleria; o, Cataleptodius; p, Coralliope; q, Demania; r, Euryxanthops; s, Leptodius; t, Neolioxantho; u, Xanthodius.

33. – 34. – 35.

Carapace anterolateral margin smooth, rounded, without teeth or lobes behind exorbital angles��������������������������������� 34 Carapace anterolateral margin interrupted, with teeth or lobes behind exorbital angles��������������������������������������������������35 Carapace 1.3–1.4 times as wide as long, widest anterior to midlength (Fig. 14.165b)������������������������������������������������Liagore Carapace 1.5–1.7 times as wide as long, widest near midlength (Fig. 14.166k)������������������������������������������������������������ Ovatis Carapace subcircular-narrowly ovate, 1.2 times as wide as long; anterolateral margins, behind exorbital angles, with 4 teeth or lobes of which second and third well defined, larger than first and fourth (Fig. 14.166h)������������������������������������������������������������������������������������������������������������������������������������������������������������Epixanthops – Carapace transversely ovate, distinctly wider than long (1.5–1.9 times as wide as long); anterolateral margins, behind exorbital angles, with 2 feebly rounded lobes followed by 2 low angular or pointed teeth, or all lobes inconspicuous�������������������������������������������������������������������������������������������������������������������������������������������� 36 36. Carapace anterolateral teeth incipient, indicated by undulation on margin (Fig. 14.165c). Chelipeds strongly unequal, dissimilar; smaller cheliped narrower, more slender than larger cheliped; finger tips rounded and hollowed; larger cheliped finger tips bluntly pointed (Fig. 14.165c)��������������������������������������������������������Lioxanthodes – Carapace anterolateral teeth with first 2 low rounded, poorly defined lobes, third and fourth bluntly angular or pointed. Chelipeds unequal, similar; finger tips bluntly pointed (Fig. 14.164k)��������������������� Lachnopodus Aldrovandiopanope Števčić, 2011

Aristotelopanope Števčić, 2011

Diagnosis. Carapace regions indicated, granular; front lobes straight, bimarginate, margins serrated; anterolateral margin with last tooth smaller than preceding two teeth. Pereopods 2–4 meri, carpi and propodi upper margins spinose. Pereopod 5 longer than carapace width; merus upper margin dentate or spinous. Male pleonite 6 wider proximally than distally. Maximum cl. 6.3 mm. Intertidal, subtidal (0–3 m). Tropical Eastern Pacific (Panama). 1 species (Garth 1986: description of only species as Micropanope taylori Garth, 1986).

Diagnosis. Carapace regions indicated, granular; front lobes slightly curved or sinuous, unimarginate, margins minutely granular; anterolateral margin with last tooth smaller than preceding two teeth. Pereopods 2–4 meri, carpi and propodi upper margins with acute granules. Pereopod 5 longer than carapace width; merus upper margin dentate or spinous. Male pleonite 6 wider distally than proximally. Maximum cl. 6 mm. Shelf (55–64 m). Tropical Eastern Pacific (Panama). 1 species (Garth 1986: description of only species as Micropanope ashcrafti Garth, 1986).

14 – Brachyura – crabs

Bottoxanthodes Števčić, 2011 Diagnosis. Carapace regions indicated anteriorly and posteriorly, smooth; anterolateral margin with last tooth smaller than preceding two teeth. Pereopod 5 shorter than carapace width; merus upper margin unarmed. Maximum cl. 8 mm. Slope (280–335 m). Tropical Eastern Pacific. 1 species (Rathbun 1930: description of only species as Paraxanthias insculptus (Stimpson, 1871)).

Camilohelleria Števčić, 2011 Diagnosis. Carapace regions indicated anteriorly, obsolete posteriorly, surface granular on anterior half, posterior half smooth; anterolateral margin with last tooth smaller than preceding two teeth. Maxilliped 3 3 merus distolateral margin auricular, produced; exopod outer margin serrated. Male pleonites 4–5 fused, pleonite 3 free. Gonopod 1 tip dilated, distinctly bent laterally. Maximum cl. 4.3 mm. Shelf (65–133 m). Tropical Eastern Pacific (Galapagos). 1 species (Garth 1986: description of only species as Micropanope manteri Garth, 1986).

Cataleptodius Guinot, 1968 Diagnosis. Carapace regions indicated anteriorly, obsolete posteriorly, surface granulation regular; anterolateral margin with 4 teeth directed anteriorly. Antennal basal article oriented obliquely to carapace midline. Larger cheliped finger tips rounded, hollowed on occlusal surface, laterally expanded, hoof or spoon-like, meeting when closed. Male pleonite 6 wider than long. Gonopod 1 with tapering tip, with row of long subdistal setae on one margin and some minute subdistal spines on other. Maximum cl. 25 mm. Intertidal–shelf (0–69 m). Tropical Atlantic, Tropical Eastern Pacific. 5 species (Guinot 1968; Manning and Chace 1990).

Coralliope Guinot, 1967 Diagnosis. Carapace regions indicated anteriorly, obsolete posteriorly, surface granular on anterior half, posterior half smooth; anterolateral margin with last tooth smaller than preceding two teeth. Maxilliped 3 distolateral margin rounded; exopod outer margin serrated. Gonopod 1 tip tapering, not bent laterally. Maximum cl. 4 mm. Subtidal–slope (6–355 m). Tropical Atlantic, Tropical Eastern Pacific. 2 species (Guinot 1967a; Monod 1956: figure of type species as Micropanope parvula (A. Milne-Edwards, 1869)).

Cycloxanthops Rathbun, 1897 Diagnosis. Carapace regions indicated anteriorly and posteriorly, smooth; front extending prominently beyond supraorbital angles, with distinct median fissure; eyes, when extended forwards, well behind level of frontal margin; anterolateral margin with 8 prominent teeth. Pereopods 2–5 stout, pereopod 5 merus less than twice as long as wide. Male pleonite 6 longitudinally rectangular. Maximum cl. 31 mm.

783

Intertidal–shelf (0–73 m). Tropical Eastern Pacific. 4 species (Rathbun 1930: figures). Cycloxanthops bocki Garth, 1957 and C. truncatus (De Haan, 1837) are included in Neoxanthops.

Demania Laurie, 1906 Diagnosis. Carapace 1.1–1.5 times as wide as long; regions well demarcated over almost entire surface, surface smooth to coarsely granular; front unimarginate; anterolateral margin with 4 broad lobes or 4 teeth. Chelipeds of male subequal or feebly unequal. Gonopod 1 elongate, slightly curved, with row of long subdistal setae on one margin and some short subdistal spines on other. Maximum cl. 75 mm (Pl. 67a). Intertidal–slope (0–780 m). Indo-West Pacific. 17 species (Ng and Yang 1989: key to 16 species). Species of Demania are relatively common on shallow coral reefs but can be fatal if eaten (Alcala et al. 1988).

Epixanthops Serène, 1984 Diagnosis. Carapace subcircular-narrowly ovate, 1.2 times as wide as long; regions weakly defined, surface smooth; anterolateral margin with 4 teeth or lobes, second and third well defined, larger than first and fourth. Maximum cl. 7.7 mm. Shelf (150 m). Western Indo-Pacific (Madagascar). 1 species (Serène 1984).

Euryxanthops Garth & Kim, 1983 Diagnosis. Carapace regions generally weakly indicated to obsolete over entire surface; front slightly anterior to anterolateral margins, breaking general outline of carapace; anterolateral margin with 4 broad lobes. Chelipeds of male subequal or feebly unequal. Pereopods 2–5 meri, carpi and propodi upper margins sharply crested. Gonopod 1 elongate, slightly curved, with row of long subdistal setae on one margin and some short subdistal spines on other. Maximum cl. 29 mm (Pl. 67b). Slope (85–550 m). Temperate Northern W Pacific, Indo-West Pacific, Temperate Australasia (Kermadec Islands). 6 species (Garth and Kim 1983; Ng and McLay 2007: key to species).

Gaudichaudia Rathbun, 1930 Diagnosis. Carapace regions indicated anteriorly and posteriorly, smooth; anterolateral margin with 3 rounded teeth all placed near lateral extremities. Antennal basal article oriented almost parallel to carapace midline. Larger cheliped finger tips rounded, hollowed on occlusal surface, meeting when closed. Maximum cl. 33 mm. Intertidal, subtidal. Tropical Eastern Pacific (Peru). 2 species (Moscoso 2013: key to species; Rathbun 1930: figure).

Guitonia Garth & Iliffe, 1992 Diagnosis. Carapace regions indicated anteriorly, obsolete posteriorly, surface granular on anterior half, posterior half smooth;

784

Marine Decapod Crustacea

anterolateral margin with last tooth smaller than preceding two teeth. Pereopod 5 merus 4 times as long as wide. Maximum cl. 10 mm. Subtidal, marine anchialine caves. Central and Eastern IndoPacific, Tropical Eastern Pacific. 3 species (Lasley et  al. 2010: rediagnosis, key to species).

Juxtaxanthias Ward, 1942 Diagnosis. Carapace regions generally weakly indicated to obsolete over entire surface; anterolateral margin with posterior 1–3 teeth pointed, preceding tooth/teeth rounded, prominent. Chelipeds of male subequal or feebly unequal. Pereopods 2–5 setose. Gonopod 1 elongate, slightly curved, with row of long subdistal setae on one margin and some short subdistal spines on other. Maximum cl. 42 mm. Intertidal. Western and Central Indo-Pacific. 3 species (Devi et al. 2019: figure; Ward 1942).

Lachnopodus Stimpson, 1858 Diagnosis. Carapace transversely ovate, distinctly wider than long (1.5–1.9 times as wide as long); regions weakly defined, surface smooth; anterolateral margin with 2 low rounded, poorly defined lobes, third and fourth bluntly lobes angular or pointed. Chelipeds unequal, similar; finger tips bluntly pointed. Maximum cl. 40 mm (Pl. 67c). Intertidal, subtidal (0–8 m). Western and Central Indo-Pacific, Temperate SW Australia. 6 species (Serène 1984: key to species).

Leptodius A. Milne-Edwards, 1863 Diagnosis. Carapace regions weakly defined, surface smooth; anterolateral margin with 4 broad lobes. Antennal basal article oriented obliquely to carapace midline. Chelipeds subequal. Larger cheliped finger tips rounded, hollowed on occlusal surface, laterally expanded, hoof or spoon-like, meeting when closed. Male pleonite 6 as long as or longer than wide. Gonopod 1 with tapering tip, with minute, subdistal spines and elongated apical lobe, bordered by anchor-shaped projections. Maximum cl. 25 mm (Pl. 67d). Intertidal, subtidal (0–6 m). Tropical W Atlantic, Temperate Northern W Pacific, Indo-West Pacific, Temperate Southern Africa, Temperate South America (Peru), Temperate Australasia. 11 species (Lee et al. 2013: taxonomy of common species; Rathbun 1930: 9 species; Serène 1984: key to 7 species).

Liagore De Haan, 1833 Diagnosis. Carapace 1.3–1.4 times as wide as long, widest anterior to midlength; regions weakly defined, surface smooth; anterolateral margin smooth, rounded, without teeth or lobes behind exorbital angles. Maximum cl. 34 mm. Shelf (81–109 m). Western and Central Indo-Pacific. 3 species (Ng and Chen 2004: systematic placement; Ng and Naruse 2007: remarks, species comparison).

Lioxanthodes Calman, 1909 Diagnosis. Carapace transversely ovate, distinctly wider than long (1.5–1.9 times as wide as long); regions weakly defined, surface smooth; anterolateral margin with 4 low undulating lobes. Chelipeds strongly unequal, dissimilar; smaller cheliped narrower, more slender than larger cheliped; finger tips rounded and hollowed; larger cheliped finger tips bluntly pointed. Maximum cl. 8 mm. Intertidal. Western and Central Indo-Pacific. 3 species (Serène 1984: descriptions of 2 species).

Macromedaeus Ward, 1942 Diagnosis. Carapace regions well defined; anterolateral margin with 4 or more prominent evenly spaced teeth or 2 small teeth at lateral extremities (M. demani (Odhner, 1925)). Antennal basal article oriented almost parallel to carapace midline. Chelipeds subequal. Larger cheliped finger tips rounded, hollowed on occlusal surface, meeting when closed. Maximum cl. 23 mm (Pl. 67e). Intertidal, subtidal (0–10 m). Indo-West Pacific. 7 species (Serène 1984: key to 6 species).

Marratha Ng & Clark, 2003 Diagnosis. Carapace regions and margins of anterolateral teeth coarsely granular; front extending prominently beyond supraorbital angles, with distinct median fissure; eyes, when extended forwards, well behind level of frontal margin; anterolateral margin with 4 teeth with tuberculate margins. Pereopods 2–5 slender, pereopod 5 merus more than twice as long as wide. Male pleonite 6 about as long as wide, appearing square. Maximum cl. 9 mm. Subtidal, shelf (18–140 m). Central Indo-Pacific and Eastern Indo-Pacific. 1 species (Ng and Clark 2003).

Megametope Filhol, 1886 Diagnosis. Carapace regions weakly defined; front extending prominently beyond supraorbital angles; eyes, when extended forwards, well behind level of frontal margin; anterolateral margin with obscure lobes if any. Maximum cl. 20 mm. Subtidal. Temperate Northern W Pacific, Temperate Australasia. 4 species (Poore 2004: key to 3 Australian species).

Metaxanthops Serène, 1984 Diagnosis. Carapace regions punctate or not indicated, smooth; front extending prominently beyond supraorbital angles, with distinct median fissure; eyes, when extended forwards, well behind level of frontal margin; anterolateral margin with 4 well developed teeth: anterior 3 large, triangular, projecting, separated by large V-shaped incisions; fourth tooth small, acute. Pereopods 2–5 slender, pereopod 5 merus more than twice as long as wide. Male pleonite 6 about as long as wide, appearing square. Maximum cl. 7.2 mm. Intertidal. Western Indo-Pacific. 1 species (Serène 1984).

14 – Brachyura – crabs

Microcassiope Guinot, 1967 Diagnosis. Carapace regions indicated anteriorly, obsolete posteriorly, surface granulation arranged into numerous transverse or oblique rows; anterolateral margin with last tooth smaller than preceding two teeth. Pereopod 5 merus 3 times as long as wide or less. Maximum cl. 6.4 mm. Intertidal–slope (0–220 m). Tropical Atlantic, Temperate Northern Pacific, Tropical Eastern Pacific. 5 species (Guinot 1967a; Rathbun 1930: 3 species as Micropanope; Takeda and Miyake 1969: rediagnosis).

Nanocassiope Guinot, 1967 Diagnosis. Carapace regions indicated anteriorly, obsolete posteriorly, surface granulation regular; anterolateral margin with last tooth smaller than preceding two teeth. Pereopod 5 merus 3 times or less as long as wide. Maximum cl. 5.2 mm (Pl. 67f). Intertidal–slope (0–220 m). Tropical Atlantic, Temperate Northern W Pacific, Western and Central Indo-Pacific, Tropical Eastern Pacific. 7 species (Davie 1995: rediagnosis, descriptions of 2 species).

785

workers (Guinot 1968; Serène 1984) included species now in other genera; we now include Cycloxanthops bocki Garth, 1957 and C. truncatus (De Haan, 1837).

Orphnoxanthus Alcock, 1898 Diagnosis. Carapace branchial regions strongly inflated, bulging, smooth; anterolateral margin with 4 teeth. Pereopods 2–5 elongate, slender; pereopod 5 merus more than 3 times as long as wide. Maximum cl. 11 mm. Shelf, slope (190–637 m). Western Indo-Pacific (Bay of Bengal). 1 species (Alcock 1898, 1899).

Ovatis Ng & Chen, 2004 Diagnosis. Carapace 1.5–1.7 times as wide as long, widest near midlength; regions weakly defined, surface smooth; anterolateral margin smooth, rounded, without teeth or lobes behind exorbital angles. Maximum cl. 6 mm. Shelf (83–92 m). Central Indo-Pacific (South China Sea). 1 species (Ng and Chen 2004).

Neolioxantho Garth & Kim, 1983

Paraxanthias Odhner, 1925

Diagnosis. Carapace regions generally weakly indicated to obsolete over entire surface; anterolateral margin with posterior two teeth bluntly angular, anterior two teeth low, rounded. Chelipeds of male subequal or feebly unequal. Gonopod 1 elongate, slightly curved, with row of long subdistal setae on one margin and some short subdistal spines on other. Maximum cl. 7.5 mm. Subtidal–shelf (6–27 m). Western and Central Indo-Pacific. 2 species (Garth and Kim 1983).

Diagnosis. Carapace regions mostly well defined, surface smooth; anterolateral margin with posterior two teeth stronger and more acute than anterior two. Cheliped palm granular and/or spinose; larger cheliped finger tips pointed, partially crossing when closed. Maximum cl. 32 mm (Pl. 67i). Shelf–slope (89–320 m). Tropical W Atlantic, Temperate Northern Pacific (California), Western and Central Indo-Pacific, Temperate Australasia. 4 species (Rathbun 1930: 3 species; Takeda and Miyake 1969: rediagnosis).

Neoxanthias Ward, 1932 Diagnosis. Carapace regions upraised, projecting, areolate, smooth or pitted giving eroded aspect; anterolateral margin with 4 broad lobes. Chelipeds of male subequal or feebly unequal. Gonopod 1 elongate, slightly curved, with row of long subdistal setae on one margin and some short subdistal spines on other. Maximum cl. 39 mm (Pl. 67g). Intertidal–slope (0–220 m). Indo-West Pacific, Temperate SE South Africa. 3 species (Serène 1984: key to species).

Neoxanthops Guinot, 1968 Diagnosis. Carapace regions smooth or rugose, without regions indicated; front extending prominently beyond supraorbital angles, with distinct median fissure; eyes, when extended forwards, well behind level of frontal margin; anterolateral margin with 4 low smooth teeth. Cheliped carpus pitted, irregular, palm surface with irregular longitudinal ridges. Pereopods 2–5 slender, pereopod 5 merus more than twice as long as wide. Male pleonite 6 about as long as wide, appearing square. Maximum cl. 33 mm (Pl. 67h). Intertidal. Western and Central Indo-Pacific, Temperate South America, Pacific coast, Temperate SW Australia. 4 species. Earlier

Paraxanthus Lucas, 1844 Diagnosis. Carapace regions indicated anteriorly and posteriorly, smooth; front extending prominently beyond supraorbital angles, with distinct median fissure; eyes, when extended forwards, well behind level of frontal margin; anterolateral margin with 3 small teeth. Pereopods 2–5 stout, pereopod 5 merus less than twice as long as wide. Male pleonite 6 longitudinally rectangular. Maximum cl. 57 mm. Intertidal–slope (0–27 m). Tropical Eastern Pacific (Peru, Chile). 1 species (Rathbun 1930: figure).

Pestoxanthodes Števčić, 2011 Diagnosis. Carapace branchial and gastric regions separated by deep narrow grooves, anterior part of mesogastric triangle distinct, short median groove from tip of mesogastric triangle extending to frontal margin and crossing transverse postfrontal groove; postfrontal groove curving laterally behind orbit and continuing parallel to anterolateral margin, terminating near last anterolateral tooth; regions in posterior half of carapace not indicated; anterolateral margin with 4 teeth. Pereopods 2–5 elongate,

786

Marine Decapod Crustacea

slender; pereopod 5 merus more than 3 times as long as wide. Maximum cl. 8 mm. Slope (280–333 m). Tropical Eastern Pacific (Panama). 1 species (Rathbun 1930: description as Parathanthias sulcatus (Faxon, 1893)).

Pseudomedaeus Guinot, 1968 Diagnosis. Carapace 1.6–1.7 times as wide as long; regions well demarcated, coarsely granular; front bimarginate; anterolateral margin with 4 teeth. Chelipeds of male subequal or feebly unequal. Gonopod 1 elongate, slightly curved, with row of long subdistal setae on one margin and some short subdistal spines on other. Maximum cl. 11 mm. Subtidal, shelf (1–185 m). Tropical Atlantic. 3 species (Guinot 1968; Monod 1956: figures).

Wardoxanthops Števčić, 2011 Diagnosis. Carapace regions punctate or not indicated, smooth; front extending prominently beyond supraorbital angles, with distinct median fissure; eyes, when extended forwards, well behind level of frontal margin; anterolateral margin with 4 low smooth teeth. Cheliped carpus and palm surface smooth. Pereopods 2–5 slender, pereopod 5 merus more than twice as long as wide. Male pleonite 6 about as long as wide, appearing square. Maximum cl. 11 mm. Intertidal. Central Indo-Pacific (NW Australia). 1 species (Guinot 1968: description of type species as Neoxanthops rotundus Guinot, 1968).

Williamstimpsonia Števčić, 2011 Diagnosis. Carapace regions mostly well defined, surface smooth; anterolateral margin with 9 subequal teeth. Maximum cl. 20 mm. Intertidal–shelf (0–21 m). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Rathbun 1930: descriptions of species as Xanthodius).

Xanthias Rathbun, 1897 Diagnosis. Carapace regions generally weakly indicated to obsolete over entire surface; anterolateral margin with 4 broad lobes or 4 teeth with 4 broad lobes or 4 teeth. Chelipeds of male subequal or feebly unequal. Gonopod 1 elongate, slightly curved, with row of long subdistal setae on one margin and some short subdistal spines on other. Maximum cl. 42 mm (Pl. 67j). Intertidal–shelf (0–110 m). Indo-West Pacific, Tropical Eastern Pacific (1 species only). 19 species (Mendoza 2013: remarks, species description; Serène 1984: key to 15 species).

Xantho Leach, 1814 Diagnosis. Carapace regions indicated, granular; anterolateral margin with 4 broad lobes or 4 teeth. Cheliped palm smooth or weakly eroded, not granular or spinose. Maximum cl. 45 mm. Intertidal–shelf (0–100 m). Temperate Northern E Atlantic, including Mediterranean. 4 species (Drach and Forest 1953: key to 4 European species with junior synonyms).

Xanthodius Stimpson, 1859 Diagnosis. Carapace regions indicated anteriorly and posteriorly, smooth; anterolateral margin with 4 broad, sometimes indistinct lobes. Antennal basal article oriented obliquely to carapace midline. Larger cheliped finger tips rounded, hollowed on occlusal surface, laterally expanded, hoof or spoon-like, meeting when closed. Gonopod 1 with broad petaloid tip; subdistal margin with dense brush of long setae. Maximum cl. 19 mm. Intertidal, subtidal (0–18 m). Tropical W Atlantic, Tropical Eastern Pacific. 4 species (Rathbun 1930: species as Xanthiodius and Leptodius).

Subfamily Zosiminae Alcock, 1898 Zosimines are shore and mostly shallow water crabs, common on tropical coral and rocky reefs, predominantly in the Indo-West Pacific. They often closely resemble xanthines and actaeines but have thin crested margins variously or always present on the chelipeds, walking legs and around the carapace. Although these groups are currently taxonomically separated, phylogenetic analyses show the subfamilies are probably not monophyletic and require revision (Lai et al. 2011). As with several xanthine genera, several classes of water soluble toxins have been isolated from the tissues of zosimines that have led to human fatalities following consumption (Garth and Alcala 1977; Tsai et al. 2006). Diagnosis. Anterolateral margin with thin crested crimped rim or 4 broad, flat rimmed lobes or teeth behind exorbital angle, or margins rounded, only subtly interrupted. Antennules folding transversely. Chelipeds usually with thin, inwardly inclined crest on upper margin of palm, fitting closely against carapace when folded; pereopods 2–5 meri, carpi and propodi upper margins usually with thin crest; pereopods 2–5 without dactylo-propodal ‘lock’. Implicit generic character. Pereopods 2–5 short, stout, merus about twice as long as wide, flattened.

Key to genera of Zosiminae 1. –

Carapace anterolateral margin strongly convex, sometimes gently sinuous but without distinct teeth or lobes, with or without narrow crested rim������������������������������������������������������������������������������������������������������������������������������2 Carapace anterolateral margin lobate or dentate, but margined with more or less broad, continuous crest or crimped rim���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������5

14 – Brachyura – crabs

2. – 3. – 4. – 5. – 6.

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Pereopods 2–5 short, stout, merus about twice as long as wide������������������������������������������������������������������������������������������������3 Pereopods 2–5 long, thin, merus ~3 times as long as wide��������������������������������������������������������������������������������������������������������4 Carapace anterolateral margin rimmed with narrow thin crest marked by shallow groove slightly inward of margin. Pereopods 2–5 flattened, upper margins of meri, carpi and propodi with thin crest (Fig. 14.168a)������������������������������������������������������������������������������������������������������������������������������������������������������������������Atergatis Carapace anterolateral margin blunt, without narrow crest around rim not marked by groove inward of margin. Pereopods 2–5 subovate in cross-section, upper margins of meri, carpi and propodi usually bluntly rounded, rarely with thin crest (Fig. 14.168b)��������������������������������������������������������������������������������������� Atergatopsis Carapace ~1.4–1.5 times as wide as long (Fig. 14.168g). Cheliped palm outer surface with shallow reticulated pitting���������������������������������������������������������������������������������������������������������������������������������������������������������� Pulcratis Carapace ~1.7–1.8 times as wide as long. Cheliped palm outer surface smooth (Fig. 14.168d)������������������������Paratergatis Pereopods 2–5 carpi upper, outer surfaces with 2 high, smooth, subparallel crests, united distally and usually proximally, forming more-or-less complete elongated cup or deep hollow (Fig. 14.168i)���������������Zozymodes Pereopods 2–5 carpi upper surfaces usually with single high crest, sometimes also with independent second low carina on outer surface������������������������������������������������������������������������������������������������������������������������������������������6 Carapace anterolateral margin with well delineated smooth thin crest, more or less broadened and indistinctly divided into 4 contiguous rounded (rarely obtusely angular) lobes separated by short, narrow, fissure�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������7

Fig. 14.168.  Xanthidae, Zosiminae. a, Atergatis integerrimus (Lamarck, 1818) ; b, Atergatopsis amoyensis De Man, 1879; c, Lophozozymus sp.; d, Paratergatis longimanus Sakai, 1964; e, Platypodia pseudogranulosa Serène, 1984; f, Platypodiella spectabilis (Herbst, 1794); g, Pulcratis reticulatus Ng & Huang, 1997; h, Zosimus aeneus (Linnaeus, 1758); i, Zozymodes cavipes (Dana, 1852).

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– 7. – 8. –

Marine Decapod Crustacea

Carapace anterolateral marginal crest divided into 4 distinct, usually angular, lobes or teeth, clearly demarcated�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������8 Carapace regions well defined; 3M undivided (Fig. 14.168e)�����������������������������������������������������������������������������������Platypodia Carapace regions usually well defined; 3M, if indicated, divided into slender anterior portion and broad posterior portion (Fig. 14.168f)����������������������������������������������������������������������������������������������������������������������������Platypodiella Carapace regions well defined, glabrous, covered with flat, confluent tubercles, regions 3M always separated from 2M; anterolateral margin with 3 broad blunt anterior lobes and narrow posterior triangular tooth (Fig. 14.168h)�������������������������������������������������������������������������������������������������������������������������������������Zosimus Carapace regions absent or weakly indicated, surface smooth or finely rugose, with regularly distributed elevations, often anteriorly setose, regions 3M and 2M confluent; anterolateral margin with 2 broad blunt anterior lobes and 2 narrower angular teeth (Fig. 14.168c)��������������������������������������������������������������� Lophozozymus Atergatis De Haan, 1833

Diagnosis. Carapace regions weakly indicated, surface smooth or punctate; anterolateral margin rimmed with narrow thin crest marked by shallow supramarginal groove. Pereopods 2–5 meri, carpi and propodi upper margins with thin crest. Maximum cl. 75 mm (Pl. 67k). Intertidal, subtidal (0–20 m). Western and Central IndoPacific, Temperate SE South Africa, Temperate Australia. 16 species (Ng and Davie 2007: species resolution; Serène 1984: key to 11 species). Atergatis rosea (Rüppell, 1830) is a Lessepian migrant to Mediterranean (Mavidis et al. 2008).

Atergatopsis A. Milne-Edwards, 1862 Diagnosis. Carapace regions more or less indicated; anterolateral margin blunt, without narrow crest around rim not marked by groove inward of margin. Pereopods 2–5 short, stout, merus about twice as long as wide, subovate in cross-section; meri, carpi and propodi upper margins usually bluntly rounded, rarely with thin crest. Maximum cl. 75 mm (Pl. 67l). Intertidal–shelf (0–166 m). Western and Central Indo-Pacific. 10 species (Serène 1984: key to 8 species).

Lophozozymus A. Milne-Edwards, 1863 Diagnosis. Carapace regions absent or weakly indicated, surface smooth or finely rugose, with regularly distributed elevations, often anteriorly setose, regions 3M and 2M confluent; anterolateral margin with 2 broad blunt anterior lobes and 2 narrower angular teeth. Pereopods 2–5 usually with single high crest. Maximum cl. 70 mm (Pl. 67r). Intertidal–shelf (0–120 m). Temperate Northern E Atlantic, Indo-West Pacific, Temperate SE South Africa, Temperate, SW Australia. 15 species (Ng and Chia 1997: remarks, descriptions of 4 species; Serène 1984: key to 13 species). At least some species of Lophozozymus are toxic to humans if consumed (Garth and Alcala 1977; Tsai et al. 2006).

Paratergatis Sakai, 1965 Diagnosis. Carapace ~1.7–1.8 times as wide as long; regions not indicated; anterolateral margin with 3 equal low lobes. Cheliped

palm outer surface smooth. Pereopods 2–5 long, thin, merus ~3 times as long as wide. Pereopods 2–5 meri upper margins slightly crested. Maximum cl. 12 mm (Pl. 67q). Shelf (60–127 m). Temperate Northern W Pacific, Western and Central Indo-Pacific. 1 species (Jeng and Ng 1998: redescription, figures; Ng and Chen 2004: figure).

Platypodia Bell, 1835 Diagnosis. Carapace regions well defined; 3M undivided; anterolateral margin with well delineated smooth thin crest, more or less broadened and indistinctly divided into 4 contiguous rounded (rarely obtusely angular) lobes separated by short, narrow, fissure. Pereopods 2–5 usually with single high crest. Maximum cl. 32 mm. Intertidal–shelf (0–132 m). Indo-West Pacific. 11 species (Serène 1984: key to species).

Platypodiella Guinot, 1967 Diagnosis. Carapace regions usually well defined; 3M, if indicated, divided into slender anterior portion and broad posterior portion; anterolateral margin with well delineated smooth thin crest, more or less broadened and indistinctly divided into 4 contiguous rounded (rarely obtusely angular) lobes separated by short, narrow, fissure. Pereopods 2–5 usually with single high crest. Maximum cl. 13 mm (Pl. 67n). Intertidal–shelf (0–37 m; associated with zoantharian, Palythoa spp. (den Hartog 1991)). Tropical Atlantic. 5 species (Guinot 1967b: diagnosis, 4 species; Martin and Zimmerman 2007: colour variation).

Pulcratis Ng & Huang, 1997 Diagnosis. Carapace ~1.4–1.5 times as wide as long; regions weakly defined, surface smooth; anterolateral margin with 4 broad lobes. Cheliped palm outer surface with shallow reticulated pitting. Pereopods 2–5 long, thin, merus ~3 times as long as wide. Pereopods 2–5 meri, carpi and propodi upper margins sharply crested. Maximum cl. 14 mm (Pl. 67m). Shelf, slope (50–400 m). Western and Central Indo-Pacific. 2 species (Mendoza et al. 2020; Ng and Huang 1997).

14 – Brachyura – crabs

Zosimus Leach, 1823 Diagnosis. Carapace regions well defined, glabrous, covered with flat, confluent tubercles, regions 3M always separated from 2M; anterolateral margin cristate, with 3 broad blunt anterior lobes and narrow posterior triangular tooth. Pereopods 2–5 usually with single high crest. Maximum cl. 59 mm (Pl. 67o). Intertidal, shallow subtidal (0–8 m). Indo-West Pacific, Temperate SE South Africa. 7 species (Hsueh 2020: remarks, species redescription).

Zozymodes Heller, 1861 Diagnosis. Carapace regions well defined; anterolateral margin with 4 teeth or lobes. Pereopods 2–5 carpi upper, outer surfaces with 2 high, smooth, subparallel crests, united distally and usually proximally and forming more–or-less complete elongated cup or deep hollow. Maximum cl. 15 mm (Pl. 67p). Intertidal, shallow subtidal (0–1 m). Indo-West Pacific, Temperate SE South Africa. 4 species (Lasley and Ng 2013: remarks, fourth species; Serène 1984: key to 3 species). References Alcala AC, Alcala LC, Garth JS, Yasumura D, Yasumoto T (1988) Human fatality due to ingestion of the crab Demania reynaudii that contained a palytoxin-like toxin. Toxicon 26, 105–107. doi:10.1016/0041-0101​ (88)90142-0 Alcock A (1898) Materials for a carcinological fauna of India. No. 3. The Brachyura Cyclometopa. Part I. The family Xanthidae. Journal of the Asiatic Society of Bengal 67, 67–233. Alcock A (1899) Illustrations of the zoology of the Royal Indian marine surveying steamer, Investigator, under the command of Commander T.H. Heming, R.N. Crustacea. – Part VII, Plates XXXVI–XLV. Indian Museum, Calcutta. Clark PF, Galil BS (1993) A revision of the xanthid genus Pilodius Dana, 1851 (Crustacea: Brachyura: Xanthoidea). Journal of Natural History 27, 1119–1206. doi:10.1080/00222939300770691 Clark PF, Ng PKL (1999) The identity of Chlorodius miliaris A. Milne Edwards, 1873, and the establishment of a new genus of Chlorodiinae (Crustacea, Decapoda, Brachyura, Xanthoidea, Xanthidae) from New Caledonia. Zoosystema 21, 353–365. Crosnier A (1991) Hypocolpus mararae, espèce nouvelle de Polynésie française (Crustacea Decapoda Brachyura Xanthidae). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 13, 423–428. Crosnier A (1995) Pleurocolpus boileaui, genre nouveau et espèce nouvelle de Polynésie française (Crustacea Decapoda Brachyura Xanthidae). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série 17, 245–251. Dai AY, Yang SL (1998) A new species of the genus Tweedieia from Nansha Islands (Crustacea: Decapoda: Brachyura: Xanthidae). Studies on Marine Fauna and Flora and Biogeography of the Nansha Islands and Neighbouring Waters 3, 317–320. Davie PJF (1993) Deepwater xanthid crabs from French Polynesia (Crustacea, Decapoda, Xanthoidea). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 14, 501–561. Davie PJF (1995) Two new species of Nanocassiope from the Western Pacific (Crustacea, Brachyura, Xanthidae). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 17, 201–210. Davie PJF (1997) Crustacea Decapoda: deep water Xanthoidea from the south-western Pacific and the Indian Ocean. In: Crosnier A (Ed.),

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Marine Decapod Crustacea

Guinot D (1967c) Recherches préliminaires sur les groupements naturels chez les Crustacés. Décapodes Brachyoures I. Les affinités des genres Aethra, Osachila, Hepatus, Hepatella et Actaeomorpha (suite et fin). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 38, 828–845. Guinot D (1968) Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures. IV. Observations sur quelques genres de Xanthidae. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 39, 695–727. Guinot D (1969) Sur divers Xanthidae, notamment sur Actaea de Haan et Paractaea gen. nov. (Crustacea Decapoda Brachyra). Cahiers du Pacifique 13, 222–267. Guinot D (1976) Constitution de quelques groupes naturels chez les Crustacés Décapodes Brachyoures: I. La superfamille des Bellioidea et trois sous-familles de Xanthidae (Polydectinae Dana, Trichiinae de Haan, Actaeinae Alcock). Mémoires du Muséum National d’Histoire Naturelle. Nouvelle Série. Série A. Zoologie 97, 1–308. Guinot D (1978) Principes d’une classification évolutive des Crustacés Décapodes Brachyoures. Bulletin Biologique de la France et de la Belgique 112, 211–292. Guinot D (1979) Données nouvelles sur la morphologie, la phylogenèse et la taxonomie des Crustacés Décapodes Brachyoures. Mémoires du Muséum National d’Histoire Naturelle, Paris, ser. Zoologie 112, 1–354. Guinot D, Macpherson E (1988) Remarques sur le genre Monodaeus Guinot, 1967, avec la description de deux espèces nouvelles (Crustacea Decapoda Brachyura). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 10, 731–757, pls 1–3. Ho PH, Ng PKL (2005) On a new species of coral-symbiont crab of the genus Cymo De Haan, 1833 (Crustacea: Decapoda: Brachyura: Xanthidae), from the South China Sea. Zootaxa 1029, 31–38. doi:10.11646/ zootaxa.1029.1.2 Hsueh PW (2020) New species and record of xanthid crabs (Crustacean, Decapoda) from Taiwan. Zootaxa 4809, 535–546. doi:10.11646/ zootaxa.4809.3.7 Jeng M-S, Ng PKL (1998) Notes on Paratergatis longimanus Sakai, 1965 (Decapoda, Brachyura, Xanthidae), a new record for Taiwan. Crustaceana 71, 827–832. doi:10.1163/156854098X00095 Karplus I, Fielder GC, Ramcharan P (1998) The Intraspecific fighting behavior of the Hawaiian boxer crab, Lybia edmondsoni – fighting with dangerous weapons? Symbiosis 24, 287–302. Komatsu H, Takeda M (2011) A new species of the genus Meractaea (Crustacea, Decapoda, Xanthidae) from Japan. Bulletin of National Science Museum, Tokyo, series A 37, 79–85. Lai JC, Mendoza JCE, Guinot D, Clark PF, Ng PKL (2011) Xanthidae MacLeay, 1838 (Decapoda: Brachyura: Xanthoidea) systematics: A multi-gene approach with support from adult and zoeal morphology. Zoologischer Anzeiger 250, 407–448. doi:10.1016/j.jcz.2011.07.002 Lasley RM, Ng PKL (2013) A new species of the crab genus Zozymodes (Crustacea: Decapoda: Xanthidae) from Guam. Micronesica 2013–03, 1–12. Lasley RM, Mendoza JCE, Ng PKL (2010) Two new species of Guitonia Garth & Iliffe, 1992 (Crustacea: Decapoda: Brachyura: Xanthidae) from the central and western Pacific Ocean. Zootaxa 2684, 1–13. doi:10.11646/​ zootaxa.2684.1.1 Lasley RM, Lai JCY, Thoma BP (2013) A new genus for Chlorodiella longimana (H. Milne Edwards) supported by morphology and molecular data, with a preliminary phylogeny of the Chlorodiellinae (Crustacea: Decapoda: Xanthidae). Invertebrate Systematics 27, 379–390. doi:10.1071/IS12075 Lasley RM, Klaus S, Ng P (2015) Phylogenetic relationships of the ubiquitous coral reef crab subfamily Chlorodiellinae (Decapoda, Brachyura, Xanthidae). Zoologica Scripta 44, 165–178. doi:10.1111/zsc.12094

Lasley RM Jr, Smith A, Paulay G, Ng PKL (2022) Revision of the coral reef crab genus Tweedieia Ward, 1935 (Decapoda: Brachyura: Xanthidae). Journal of Crustacean Biology 42, ruab081. Lee S-K, Mendoza JCE, Ng PKL, Kim W (2013) On the identity of the Indo-West Pacific littoral xanthid crab, Leptodius exaratus (H. Milne Edwards, 1834) (Crustacea: Decapoda: Brachyura: Xanthidae). Raffles Bulletin of Zoology 61, 189–204. Llewellyn LE, Dodd MJ, Robertson A, Ericson G, de Koning C, et  al. (2002) Post-mortem analysis of samples from a human victim of a fatal poisoning caused by the xanthid crab, Zosimus aeneus. Toxicon 40, 1463–1469. doi:10.1016/S0041-0101(02)00164-2 Manning RB, Chace FA (1990) Decapod and stomatopod Crustacea from Ascension Island, South Atlantic Ocean. Smithsonian Contributions to Zoology 503, 1–91. doi:10.5479/si.00810282.503 Manuel-Santos MR, Ng PKL (2007) On the genus Ladomedaeus Števčić, 2005, from the Philippines and Japan, and the status of the Ladomedaeidae Števčić, 2005 (Decapoda: Brachyura: Xanthoidea). Raffles Bulletin of Zoology Supplement 16, 177–185. Martin JW, Zimmerman TL (2007) Color variation in the Caribbean crab Platypodiella spectabilis (Herbst, 1794) (Decapoda, Brachyura, Xanthidae). Gulf and Caribbean Research 19, 59–63. doi:10.18785/gcr.1901.08 Mavidis M, Türkay M, Koukouras A (2008) The genera Atergatis, Microcassiope, Monodaeus, Paractea, Paragalene, and Xantho (Decapoda, Xanthidae) in the Mediterranean Sea. Crustaceana 81, 1035–1053. doi:10.1163/156854008X360798 Mendoza JCE (2013) A new species of ocellated Xanthias Rathbun, 1897 (Crustacea: Decapoda: Brachyura: Xanthidae) from the Bohol Sea, Philippines. Zootaxa 3636, 374–384. doi:10.11646/zootaxa.3636.2.8 Mendoza JCE, Guinot D (2011) Revision of the genus Glyptoxanthus A. Milne-Edwards, 1879, and establishment of Glyptoxanthinae nov. subfam. (Crustacea: Decapoda: Brachyura: Xanthidae). Zootaxa 3015, 29–51. doi:10.11646/zootaxa.3015.1.4 Mendoza JCE, Manuel-Santos MR (2012) Revision of Garthiella Titgen, 1986 (Crustacea: Decapoda: Brachyura: Xanthidae), with description of a new subfamily and a new species from the central Philippines. Zootaxa 3446, 32–48. doi:10.11646/zootaxa.3446.1.2 Mendoza JCE, Ng PKL (2008a) New genera and species of euxanthine crabs (Crustacea: Decapoda: Brachyura: Xanthidae) from the Bohol Sea, the Philippines. Raffles Bulletin of Zoology 56, 385–404. Mendoza JCE, Ng PKL (2008b) A new species of Alainodaeus Davie, 1993 (Crustacea: Decapoda: Brachyura: Xanthidae) from Balicasag Island, Philippines, with a key to the genus. Zootaxa 1897, 53–63. doi:10.11646/ zootaxa.1897.1.5 Mendoza JCE, Ng PKL (2010) The euxanthine crabs (Crustacea: Brachyura: Xanthidae) of the Philippines. Raffles Bulletin of Zoology 58, 57–74. Mendoza JCE, Ng PKL (2011) The Polydectinae Dana, 1851, of the Philippines, with description of a new genus for Lybia hatagumoana Sakai, 1961 (Crustacea: Decapoda: Brachyura: Xanthidae). Zootaxa 3052, 51–61. doi:10.11646/zootaxa.3052.1.2 Mendoza JCE, Ng PKL (2012) A new genus and species of deep-water xanthid crab (Decapoda, Brachyura, Xanthidae) from the Philippines. In: Komatsu H, Okuno J, Fukuoka K (Eds) Studies on Eumalacostraca: a homage to Masatsune Takeda. Crustaceana Monographs 17, 197–210. doi:10.1163/9789004202894_017 Mendoza JCE, Chong VC, Ng PKL (2009a) A new xanthid crab of the genus Medaeops Guinot, 1967, from peninsular Malaysia, with a note on Leptodius granulosus Haswell, 1882 (Crustacea: Decapoda: Brachyura: Xanthidae). Zootaxa 2297, 44–54. doi:10.11646/zootaxa.2297.1.4 Mendoza JCE, Manuel-Santos MR, Ng PKL (2009b) Rediscovery of the rare euxanthine crab, Guinotellus melvillensis Sèrene, 1971 (Decapoda, Brachyura, Xanthidae) in the Philippines. Crustaceana 82, 39–51. doi:10.1163/156854008X363687

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Mendoza JCE, Kazmi QB, Moazzam M (2012) First report of two rare xanthid crabs (Crustacea: Decapoda: Brachyura: Xanthidae) from Pakistan. Marine Biodiversity Records 5, e51. doi:10.1017/ S1755267211001084 Mendoza JCE, Mani P, Ravichandran S (2020) First record of the deep-water xanthid crab genus, Pulcratis Ng & Huang, 1997, from the Indian Ocean, with description of a new species (Crustacea: Brachyura: Xanthidae). Nauplius 28, e2020010. doi:10.1590/2358-2936e2020010 Monod T (1956) Hippidea et Brachyura ouest-africains. Mémoires de I’Institut Français d’Afrique Noire 45, 1–674. Moscoso V (2013) Clave para identificación de crustáceos decápodos y estomatópodos del Perú. Boletín Instituto de Mar del Perú 28, 8–135. Naruse T, Forges MRd, Forges BRD, Uyeno D (2020) Records of Cyrtocarcinus truncatus (Rathbun, 1906) (Decapoda: Xanthidae) from the Loyalty Islands, New Caledonia, and the Ryukyu Islands, Japan. Crustacean Research 49, 33–39. doi:10.18353/crustacea.49.0_33 Naruse T, Fujita Y (2015) Lipkemera iejima, a new cavernicolous crab (Brachyura: Xanthidae) from a submarine cave at Ie Island, central Ryukyu Islands, Japan. Crustacean Research 44, 21–27. doi:10.18353/ crustacea.44.0_21 Naruse T, Maenosono T, Ng PKL (2021) Remarkable bilaterally asymmetrical gonopores and gonopods in a new genus and species of brachyuran crab from the Ryukyu Islands, Japan (Decapoda: Brachyura: Xanthidae). Journal of Crustacean Biology 41, doi:10.1093/jcbiol/ruab022. Ng PKL (1993a) Kraussiinae, a new subfamily for the genera Kraussia Dana, 1852, Palpedia, new genus, and Garthasia new genus (Crustacea: Decapoda: Brachyura: Xanthidae), with descriptions of two new species from Singapore and the Philippines. Raffles Bulletin of Zoology 41, 133–157. Ng PKL (1993b) On a new genus and species of xanthid crab (Crustacea: Decapoda: Brachyura) from Chesterfield Island, Coral Sea. Proceedings of the Biological Society of Washington 106, 705–713. Ng PKL (2002a) Olenothus, a new genus of euxanthine crab (Crustacea: Decapoda: Brachyura: Xanthidae) from Guam. Micronesica 34, 201–208. Ng PKL (2002b) On a new species of cavernicolous Neolimera (Crustacea: Decapoda: Brachyura: Xanthidae) from Christmas Island and Ryukyus, Japan. Raffles Bulletin of Zoology 50, 95–99. Ng PKL (2014) A new deep-water species of Forestiana (Crustacea: Decapoda: Brachyura: Xanthidae) from Taiwan, with a clarification of the name F. depressa (White, 1848). Bishop Museum Bulletin in Zoology 9, 115–122. Ng PKL, Chen HL (2004) On a new genus and new species of xanthid crab (Crustacea, Decapoda, Brachyura, Xanthinae) from the South China Sea, with notes on the genus Liagore De Haan, 1833. Journal of Natural History 38, 2345–2360. doi:10.1080/00222930310001647352 Ng PKL, Chia DGB (1994) The genus Glyptocarcinus Takeda, 1973, with descriptions of a new subfamily, two new genera and two new species from New Caledonia (Crustacea: Decapoda: Brachyura: Xanthidae). Raffles Bulletin of Zoology 42, 701–731. Ng PKL, Chia DGB (1997) Lophozozymus erinnyes, a new species of poisonous crab from Australia, with notes on L. pictor (Fabricius, 1798), L. incisus (H. Milne Edwards, 1834) and L. edwardsi (Odhner, 1925) (Crustacea: Decapoda: Brachyura: Xanthidae). Raffles Bulletin of Zoology 45, 419–443. Ng PKL, Clark PF (2003) Three new genera of Indo-West Pacific Xanthidae (Crustacea, Decapoda, Brachyura, Xanthoidea). Zoosystema 25, 131–147. Ng PKL, Davie PJF (2007) On the identity of Atergatis floridus (Linnaeus, 1767) and recognition of Atergatis ocyroe (Herbst, 1901) as a valid species from the Indian Ocean (Crustacea: Brachyura: Xanthidae). Raffles Bulletin of Zoology Supplement 16, 169–175. Ng PKL, Huang JF (1997) Unrecorded crabs (Crustacea: Decapoda: Brachyura) from Taiwan and Tungsha Islands, with description of a new genus and species of Xanthidae. Zoological Studies 36, 261–276.

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Ng PKL, McLay CL (2007) Two new species of deep-water xanthid crabs of the genera Euryxanthops Garth & Kim, 1983, and Medaeops Guinot, 1967 (Crustacea: Decapoda: Brachyura: Xanthidae) from New Zealand. Zootaxa 1505, 37–50. doi:10.11646/zootaxa.1505.1.3 Ng PKL, Naruse T (2007) Liagore pulchella, a new species of xanthid crab (Crustacea: Decapoda: Brachyura) From Vanuatu. Zootaxa 1665, 53–60. Ng PKL, Tan LWH (1985) ‘Right handedness’ in heterochelous calappoid and xanthoid crabs: suggestion for a functional advantage. Crustaceana 49, 98–100. doi:10.1163/156854085X00288 Ng PKL, Yang CM (1989) On some species of Demania Laurie, 1906 (Crustacea: Decapoda: Brachyura: Xanthidae) from Malaysia, Singapore and the Philippines, with a key for the genus. Raffles Bulletin of Zoology 37, 37–50. Ng PKL, Yang SL (1998) Description of a new genus for the xanthid crab Pilodius etisoides Takeda and Miyake, 1968 (Crustacea: Decapoda: Brachyura: Xanthoidea). Journal of Natural History 32, 1685–1696. doi:10.1080/00222939800771201 Ng PKL, Chen HL, Boyko CB (2005) On two species of euxanthine crabs from the South China Sea, including a description of a new species of Crosnierius (Crustacea: Decapoda: Brachyura: Xanthidae). Proceedings of the Biological Society of Washington 118, 319–325. doi:10.2988/0006324X​(2005)118[319:OTSOEC]2.0.CO;2 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Ng PKL, Priyaja P, Kumar AB, Devi SS (2019) A collection of crabs (Crustacea, Brachyura) from the southwestern coast of India, with a discussion of the systematic position of Nectopanope Wood-Mason in Wood-Mason & Alcock, 1891 (Euryplacidae). ZooKeys 818, 1–24. doi:10.3897/zookeys.818.32108 Odhner T (1925) Monographierte Gattungen der Krabbenfamilie Xanthidae. I. Göteborgs Kungliga Vetenskaps-och Vitterhets-samhälles Handlingar, Series 4 24, 1–92, pls 1–5. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Poupin J, Starmer J (2013) Neoliomera moana, a new cavernicolous species of xanthid crab from the Marquesas Islands (Crustacea: Decapoda: Brachyura). Zootaxa 3737, 585–592. doi:10.11646/zootaxa.3737.5.5 Rathbun MJ (1930) The cancroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae. Bulletin of the United States National Museum 152, 1–609, pls 1–230. Sakai T (1976) Crabs of Japan and adjacent seas. Kodansha Ltd, Tokyo. Sakai T (1983) Description of new genera and species of Japanese crabs together with systematically and biogeographically interesting species. (I). Researches on Crustacea 12, 1–44, pls 1–8. Serène R (1972) Observations on the Indo-Pacific species of Kraussia Dana 1852 (Decapoda: Brachyura). Journal of the Royal Society of Western Australia 55, 41–62. Serène R (1984) Crustacés Décapodes Brachyoures de l’Océan Indien Occidental et de la Mer Rouge, Xanthoidea: Xanthidae et Trapeziidae. Avec un addendum par Crosnier, A.: Carpiliidae et Menippidae. Faune Tropicale 24, 1–400. Serène R, Vadon C (1981) Crustacés Décapodes: Brachyoures. Liste préliminaire, description de formes nouvelles et remarques taxonomiques. In: Forest J (Ed.) Résultats des campagnes MUSORSTOM: I. Philippines (18–28 Mars 1976). Mémoires du Muséum National d’Histoire Naturelle. Série A, Zoologie 91, 117–140. Soto LA (1980) Allactaea lithostrota Williams, 1979: new records and range extension (Decapoda: Xanthidae). Florida Scientist 43, 125–128. Števčić Z (2005) The reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Natura Croatica 14(Supplement 1), 1–159.

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Takeda M (1989) Shallow-water crabs from the Oshima Passage between Amami-Oshima and Kakeroma-jima Islands, the Northern Ryukyu Islands. Memoirs of The National Science Museum, Tokyo 22, 135–184, pl. 4. Takeda M, Komatsu H (2010) A new xanthid crab (Decapoda, Brachyura) from a submarine cave in the Philippines. In: Fransen C, De Grave S, Ng P (Eds) Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume. Crustaceana Monographs 14, 677–683. Takeda M, Komatsu H (2011) A new genus and species of euxanthine crab (Crustacea: Decapoda: Xanthidae) from submarine banks off the Izu Islands, Central Japan. Bulletin of the National Science Museum, Tokyo, Series A (Zoology) 5, 119–126. Takeda M, Marumura M (2002) The genus Pseudactea Seréne, 1962 (Crustacea, Decapoda), with description of a new species from central Japan. Bulletin of the National Science Museum, Tokyo (ser. A). Zoology 28, 101–107. Takeda M, Miyake S (1968) Two new xanthid crabs inhabiting coral reefs of the Ryukyu Islands. Ohmu 1, 183–189, pl. 8. Takeda M, Miyake S (1969) On two species of the family Xanthidae (Crustacea, Brachyura) from southern Japan. Ohmu 2, 195–206. Thoma BP, Guinot D, Felder DL (2014) Evolutionary relationships among American mud crabs (Crustacea: Decapoda: Brachyura: Xanthoidea) inferred from nuclear and mitochondrial markers, with comments on adult morphology. Zoological Journal of the Linnean Society 170, 86–109. doi:10.1111/zoj.12093 Titgen RH (1986) Hawaiian Xanthidae (Decapoda: Brachyura) II. Description of Garthiella, new genus, with a redescription of G. aberrans (Rathbun, 1906). Proceedings of the Biological Society of Washington 99, 56–60. Tsai Y-H, Ho P-H, Hwang C-C, Hwang P-A, Cheng C-A, et  al. (2006) Tetrodotoxin in several species of xanthid crabs in southern Taiwan. Food Chemistry 95, 205–212. doi:10.1016/j.foodchem.2004.12.032 Vázquez-Bader AR, Gracia A (2004) A new genus and new species of crab of the family Xanthidae MacLeay, 1838 (Crustacea: Decapoda: Brachyura) from the southwestern Gulf of Mexico. Proceedings of the Biological Society of Washington 117, 505–513. Ward M (1933) New genera and species of marine Decapoda Brachyura from the coasts of New South Wales and Queensland. Australian Zoologist 7, 377–394, pls 21–23. Ward M (1942) Notes on the Crustacea of the Desjardins Museum, Mauritius Institute, with descriptions of new genera and species. Mauritius Institute Bulletin 2, 49–113, pls 5, 6. Williams AB (1974) Allactaea lithostrota, a new genus and species of crab (Decapoda: Xanthidae) from North Carolina, U.S.A. Proceedings of the Biological Society of Washington 87, 19–26. Williams AB (1984) Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C. Yasumoto T, Oshima Y, Tajiri M, Kotaki Y (1983) Paralytic shellfish toxins in previously unrecorded species of coral reef crabs. Nippon Suisan Gakkaishi 49, 633–636. doi:10.2331/suisan.49.633

Subsection Thoracotremata Guinot, 1977 Guinot (1977) introduced Thoracotremata for crabs with the ‘highest’ level of structural organisation, exemplified in the presence of gonopores on the thoracic sternum in both sexes. Thoracotremes are primarily shore and land crabs and include the most ‘terrestrialised’ brachyurans. Whereas monophyly of Thoracotremata is well corroborated, its

largest constituent superfamilies, Grapsoidea and Ocypodoidea, are unlikely to be natural groups (Ma et  al. 2019; Schubart et al. 2006; Tsang et al. 2018; Tsang et al. 2014). Diagnosis. Gonopores of both sexes on thoracic stermum, on sternite 8 in males, on sternite 6 in females. References Guinot D (1977) Propositions pour une novelle classification des Crustacés, Décapodes, Brachyoures. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 285, 1049–1052. Ma KY, Qin J, Lin C-W, Chan T-Y, Ng PKL, et al. (2019) Phylogenomic analyses of brachyuran crabs support early divergence of primary freshwater crabs. Molecular Phylogenetics and Evolution 135, 62–66. doi:10.1016/j.ympev.2019.02.001 Schubart CD, Cannicci S, Vannini M, Fratini S (2006) Molecular phylogeny of grapsoid crabs (Decapoda, Brachyura) and allies based on two mitochondrial genes and a proposal for refraining from current superfamily classification. Journal of Zoological Systematics and Evolutionary Research 44, 193–199. doi:10.1111/j.1439-0469.2006.00354.x Tsang LM, Schubart CD, Ahyong ST, Lai JCY, Au EYC, et al. (2014) Evolutionary history of true crabs (Crustacea: Decapoda: Brachyura) and the origin of freshwater crabs. Molecular Biology and Evolution 31, 1173–1187. doi:10.1093/molbev/msu068 Tsang LM, Ahyong ST, Shih H-T, Ng PKL (2018) Further polyphyly of pinnotheroid crabs: the molecular phylogenetic position of the polychaete-associated Aphanodactylidae. Invertebrate Systematics 32, 92–99. doi:10.1071/IS17038

Aphanodactyloidea Ahyong & Ng, 2009 All members of the only family, Aphanodactylidae, are associated with tube building polychaete worms (Terebellidae and Chaetopteridae). All have short, stout, ambulatory legs with one or more spines lining the flexor margins of the ischiomerus of at least some legs, and one or more short spines on the ambulatory propodus opposing a short, clawlike dactylus. All are sexually dimorphic, the female being distinctly wider than the male. Only 13 species are known Their morphological similarities and association with other invertebrates have led to their association with pea crabs of the family Pinnotheridae (Ahyong and Ng 2009; Davie et al. 2015; Ng et al. 2008), most recently in a superfamily, Pinnotheroidea. The most recent molecular study has shown that these similarities are convergent and the closest aphanodactylid relatives are ocypodoid shore crabs of the family Macrophthalmidae (Tsang et al. 2018). Aphanodactylids are not readily accommodated among the ocypodoids as currently understood. The high level placement of Aphanodactylidae may be clarified when the current polyphyly of Ocypodoidea and Grapsoidea is resolved; for the time being, however, we treat the group in its own superfamily. Diagnosis. Carapace transversely ovate, squarish or rounded, never pyriform or subpyriform; antennular fossae

14 – Brachyura – crabs

wider than long, antennules folding transversely or obliquely. Antennal gland opening on surface of epistome, not on margin of buccal cavity. Maxilliped 3 merus smaller than ischium; palp articles clearly demarcated, not enlarged. Pereopods 2–5 usually with 1 or more small spines along flexor margin of merus; propodus usually with 1 or small distal spines or spiniform setae opposing a short dactylus.

793

Aphanodactylidae Ahyong & Ng, 2009 Figure 14.169, Plate 68a Implicit generic characters. Orbit laterally rounded. Maxilliped 3 palp entirely visible from ventral view. Pereopods 2–5 meri as long as or slightly longer than propodus and carpus combined. Male pleonites and telson free.

Key to genera of Aphanodactylidae 1. – 2. – 3.

Maxilliped 3 ischium triangular, inner margin produced to rounded mesiodistal lobe (Fig. 14.169j). Male pleonites 4–6 fused�������������������������������������������������������������������������������������������������������������������������������������������� Gustavus Maxilliped 3 ischium subquadrate-trapezoid, mesiodistal angle not produced. Male pleonites free��������������������������������2 Pereopod 5 merus less than half as long as pereopod 4 merus (Fig. 14.169e)�������������������������������������������������������������������������3 Pereopod 5 merus more than half as long as pereopod 4 merus (Fig. 14.169p, r)������������������������������������������������������������������4 Pereopods 2–5 dactyli short, broad, without tooth on flexor margin (Fig. 14.169x). Maxilliped 3 ischium subquadrate, inner margin convex, little shorter than distal margin (Fig. 14.169o)���������������������������������������������Uruma

Fig. 14.169.  Aphanodactylidae. Carapace, female: a, Ankerius; b, Gustavus; c, Lichtylus; d, Selwynia. Habitus, female: e, Takedactylus compressus Naruse & Yoshida, 2018. Right orbit: f, Gandoa; g, Selwynia. Maxilliped 3: h, Ankerius; i, Gandoa; j, Gustavus; k, l, Lichtylus (with inside view); m, Selwynia; n, Takedactylus; o, Uruma. Pereopod 4: p, Ankerius; q, Gandoa. Pereopod 5: r, Ankerius. Pereopod dactylus: s, Ankerius; t, Gustavus; u, Lichtylus; v, Selwynia; w, Takedactylus; x, Uruma.

794

– 4. – 5. – 6. –

Marine Decapod Crustacea

Pereopods 2–5 dactyli longer than wide, with tooth on flexor margin (Fig. 14.169w). Maxilliped 3 ischium wider than long, with convex inner margin half as long as distal margin (Fig. 14.169n)���������������������������Takedactylus Carapace of female trapezoidal (Fig. 14.169c). Maxilliped 3 palp with lamellar carpus, propodus and carpus concealed in external view (Fig. 14.169k, l)��������������������������������������������������������������������������������������������������Lichtylus Carapace of female subquadrate or ovoid (Fig. 14.169a, d). Maxilliped 3 palp fully visible in external view�������������������5 Carapace of female transversely subquadrate; surface distinctly punctate. Orbit extended laterally to meet anterolateral margin (Fig. 14.169f)���������������������������������������������������������������������������������������������������������������������Gandoa Carapace of female transversely ovate; surface smooth, with few scattered pits. Orbit laterally rounded (Fig. 14.169g)��������������������������������������������������������������������������������������������������������������������������������������������������������������������������������6 Maxilliped 3 ischium inner margin straight, merus wider than long (Fig. 14.169m). Pereopods 2–5 dactyli about as long as wide (Fig. 14.169v)����������������������������������������������������������������������������������������������������������������������������Selwynia Maxilliped 3 ischium inner margin convex, merus longer than wide (Fig. 14.169h). Pereopods 2–5 dactyli longer than wide (Fig. 14.169s)������������������������������������������������������������������������������������������������������������������������������������Ankerius Ankerius Ng, Rahayu & Shih, 2016

Diagnosis. Carapace of female as long as wide, deeply vaulted, dome shaped; surface smooth. Maxilliped 3 ischium subtriangular, mesial margin convex. Pereopods 2–5 propodus with opposing teeth on distoflexor corners; dactylus elongate, longer than greatest width. Pereopod 5 merus two-thirds length of pereopod 4 merus. Maximum cl. 7 mm. Subtidal (10–20 m; associated wiith Chaetopterus (Chaetopteridae)). Western Indo-Pacific (Red Sea). 2 species (Naruse and Uyeno 2017; Ng et al. 2016).

Gandoa Kammerer, 2006 Diagnosis. Carapace of female transversely subquadrate; surface distinctly punctate. Orbit extended laterally to meet anterolateral margin. Maxilliped 3 ischium subquadrate, inner margin convex, longer than distal margin. Pereopods 2–5 propodus without opposing teeth on distoflexor corners; dactylus triangular, minute, about as long as wide. Pereopod 5 merus two-thirds length of pereopod 4 merus. Maximum cl. 5.3 mm. Subtidal (?). Western Indo-Pacific (Zanzibar, Mayotte). 1 species (Ng and Rahayu 2016: rediagnosis).

Gustavus Ahyong & Ng, 2009 Diagnosis. Surface smooth, finely setose. Maxilliped 3 ischium triangular, mesiodistal angle produced to rounded lobe. Pereopods 2–5 propodus with opposing teeth on distoflexor corners; dactylus triangular, minute, about as long as wide. Pereopod 5 merus less than half as long as pereopod 4 merus. Male pleonites 4–6 fused. Maximum cl. 8.8 mm. Subtidal. Central Indo-Pacific (Guam). 1 species (Ahyong and Ng 2009).

Lichtylus Naruse & Yoshida, 2018 Diagnosis. Carapace of female trapezoidal; surface smooth. Maxilliped 3 ischium subquadrate, inner margin convex, little shorter than distal margin; palp with lamellar carpus, propodus and

carpus concealed in ventral view. Pereopods 2–5 propodus without opposing teeth on distoflexor corners; dactylus elongate, longer than greatest width. Pereopod 5 merus two-thirds length of pereopod 4 merus. Maximum cl. 2.3 mm. Subtidal (10 m; host Eupolymnia (Terebellidae)). Central IndoPacific (Ryukyu Is). 1 species (Naruse and Yoshida 2018).

Selwynia Borradaile, 1903 Diagnosis. Carapace of female transversely ovate; surface smooth, with few scattered pits. Maxilliped 3 ischium subquadrate, inner margin convex, little shorter than distal margin. Pereopods 2–5 propodus with opposing teeth on distoflexor corners; dactylus triangular, minute, about as long as wide. Pereopod 5 merus more than half as long as pereopod 4 merus. Maximum cl. 9.5 mm (Pl. 68a). Intertidal–shelf (0–36 m). Indo-West Pacific. 7 species (Ng et  al. 2010; Ng and Rahayu 2016: rediagnosis, key to males and females). Aphanodactylus Tesch, 1918, which gives the family its name, is a junior synonym (Ng and Clark 2014).

Takedactylus Naruse & Maenosono, 2012 Diagnosis. Carapace of female subrectangular, wider than long; surface smooth. Maxilliped 3 ischium wider than long, with convex inner margin half as long as distal margin. Pereopods 2–5 propodus with opposing teeth on distoflexor corners; dactylus longer than greatest width, with tooth on flexor margin. Pereopod 5 merus less than half as long as pereopod 4 merus. Maximum cl. 2.9 mm. Subtidal (1–15 m; associated with Loimia (Terebellidae)). Central Indo-Pacific (Ryukyu Is). 2 species (Naruse and Maenosono 2012; Naruse and Yoshida 2018).

Uruma Naruse, Fujita & Ng, 2009 Diagnosis. Carapace of female transversely subquadrate; surface smooth. Maxilliped 3 ischium subquadrate, inner margin convex, little shorter than distal margin. Pereopods 2–5 meri ~1.5 times as long as propodus and carpus combined; propodus with opposing teeth on distoflexor corners; dactylus triangular, minute, about as

14 – Brachyura – crabs

long as wide. Pereopod 5 merus less than half as long as pereopod 4 merus. Maximum cl. 4.2 mm. Subtidal (8 m; host unknown). Central Indo-Pacific (Ryukyu Ryuku Is). 1 species (Naruse et al. 2009). References Ahyong ST, Ng PKL (2009) Aphanodactylidae, a new family of thoracotreme crabs (Crustacea: Brachyura) symbiotic with polychaete worms. Zootaxa 2289, 33–47. doi:10.11646/zootaxa.2289.1.3 Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. Naruse T, Maenosono T (2012) A new genus and species of Aphanodactylidae Ahyong & Ng, 2009 (Decapoda, Brachyura, Pinnotheroidea) from the Ryukyu Islands, Japan. In: Komatsu H, Okuno J, Fukuoka K (Eds) Studies on Eumalacostraca: a homage to Masatsune Takeda. Crustaceana Monographs 17, 225–231. doi:10.1163/9789004202894_019 Naruse T, Uyeno D (2017) Ankerius grusocurare, a new species of Aphanodactylidae (Decapoda, Brachyura) from Iriomote Island, Ryukyu Islands, Japan. Crustaceana 90, 1055–1060. doi:10.1163/15685403-00003668 Naruse T, Yoshida R (2018) Two new species of Aphanodactylidae (Crustacea: Decapoda: Brachyura) from the Ryukyu and Ogasawara Islands, Japan, with the establishment of a new genus. Tropical Zoology 31, 21–33. doi:10.1080/03946975.2017.1395163 Naruse T, Fujita Y, Ng PKL (2009) A new genus and new species of symbiotic crab (Crustacea: Brachyura: Pinnotheroidea) from Okinawa. Zootaxa 2053, 59–68. doi:10.11646/zootaxa.2053.1.4 Ng PKL, Clark PF (2014) Selwynia, a senior synonym of Aphanodactylus, with a redescription of Selwynia laevis (Crustacea: Brachyura: Pinnotheroidea: Aphanodactylidae). Marine Biodiversity Records 7, e113. doi:10.1017/S1755267214001134 Ng PKL, Rahayu DL (2016) On the genera Selwynia Borradaile, 1903, and Gandoa Kammerer, 2006, with descriptions of two new species from Papua New Guinea and French Polynesia (Crustacea: Decapoda: Brachyura: Aphanodactylidae). Zootaxa 4092, 339–370. doi:10.11646/ zootaxa.4092.3.2 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Ng PKL, Ahyong ST, Komai T (2010) On the male of Aphanodactylus loimiae Nonishi & Noda, 1999 (Crustacea: Brachyura: Pinnotheroidea: Aphanodactylidae). Raffles Bulletin of Zoology 58, 75–78. Ng PKL, Rahayu DL, Shih HT (2016) Ankerius aenigmaticus, a new genus and new species of aphanodactylid crab symbiotic with polychaetes from the Red Sea coast of Saudi Arabia (Crustacea: Decapoda: Brachyura: Aphanodactylidae). Zootaxa 4179, 271–278. doi:10.11646/zootaxa.4179.2.7 Tsang LM, Ahyong ST, Shih H-T, Ng PKL (2018) Further polyphyly of pinnotheroid crabs: the molecular phylogenetic position of the polychaete-associated Aphanodactylidae. Invertebrate Systematics 32, 92–99. doi:10.1071/IS17038

Cryptochiroidea Paulson, 1875 coral gall crabs Gall crabs inhabit cavities in living scleractinian corals, an environment in which the females become trapped for their

795

entire adult life. Males are more mobile. The crabs are obligate associates of corals residing in tunnels or pits in the coral skeleton and feed on the host coral mucus and tissues (Chan et  al. 2020; Kropp 1986). They induce the coral to form a protective gall around them. Most are from shallow coral reefs in the photic zone but species of a few are known only from deep-water corals. Gall crabs are typically less than 5 mm long, the largest reaching not more than twice this length. There is little doubt that Cryptochiroidea are monophyletic (van der Meij and Schubart 2014) but their relationship to other thoracotreme superfamilies is uncertain. Three clades of genera within Cryptochiridae have been repeatedly recovered by molecular data (van der Meij and N ­ ieman 2016) but morphological support for these groups has not been sought. The groups of genera are more or less correlated with their host corals (Chan et al. 2020; van der Meij 2015a). Gall crabs inhabit a wide range of scleractinian families. Kropp (1990) listed host families for most species in his taxonomic review. Host specificity has been long debated and thought for many years to be low (Hoeksema et  al. 2012). van der Meij (2015b) on the other hand has shown that sibling species may be differentiated on the basis of coral host. van der Meij’s (2015a) thesis dealt with phylogeny, taxonomy, host specifity, coevolution, biogeography, distribution and reproduction. Several of the chapters included there have been subsequently published. There is one family. Diagnosis. Carapace longer than wide, pyriform to longitudinally ovate or rectangular, transversely arched, weakly to well calcified. Orbit ill-defined, eye mostly exposed. Antennules folding longitudinally or almost so. Maxilliped 3 ischium distinctly smaller than merus, both free. Pereopods 2–5 stout, pereopod 5 shorter than carapace length.

Cryptochiridae Paulson, 1875 Figures 14.170–14.172, Plate 68b–j Kropp (1990) rediagnosed, or diagnosed as new, genera from the Indo-West Pacific. All species then known were listed; few have been described since. Three genera have been added since 1990 (Badaro et  al. 2012; Kropp and Manning 1996). Kropp and Manning (1987) dealt with the four genera occurring in the Atlantic. Kropp’s (1990) key relied in part on some characters difficult to interpret including the degree of fusion of the pterygostomian region to the carapace. The latter may not be a reliable feature (Zayasu et  al. 2013), at least for Fizesereneia. Males are infrequently caught and may not be reliably identified with this key.

796

Marine Decapod Crustacea

Implicit generic attributes. Carapace longer than wide; mesogastric region not inflated, with isolated tubercles and granules; pterygostomian region fused to carapace.

Thoracic sternite 7 with median suture. Maxilliped 3 with exopod. Pereopod 2 merus with distomesial and distolateral margins similar, neither overlapping base of carapace.

Key to of genera of Cryptochiridae (females) l. – 2. –

Carapace deflected anteriorly (Figs 14.170c, f, h, s, 14.171e, g, k)���������������������������������������������������������������������������������������������2 Carapace not deflected anteriorly (Fig. 14.170j, m)�������������������������������������������������������������������������������������������������������������������13 Carapace widest anterior to midlength (Figs 14.170d, g, 14.171a, j)�����������������������������������������������������������������������������������������3 Carapace widest at or posterior to midlength������������������������������������������������������������������������������������������������������������������������������6

Fig. 14.170.  Cryptochiridae. Carapace, dorsal and lateral views: a, Cecidocarcinus; b, c, Cryptochirus; d, Dacryomaia; e, f, Detocarcinus; g, h, Fizesereneia; i, j, Fungicola; k, Hapalocarcinus; l, m, Hiroia; n, Kroppcarcinus; o, Lithoscaptus; p, Luciades; q, Neotroglocarcinus; r, s, Opecarcinus.

14 – Brachyura – crabs

3. – 4. – 5. – 6. – 7. – 8. – 9. – 10.

797

Anterior carapace with 2 bowl-shaped concavities (Figs 14.170g, 14.171a)�����������������������������������������������������������������������������4 Anterior carapace without concavities�����������������������������������������������������������������������������������������������������������������������������������������5 Anterior concavities divided and surrounded by 1 or 2 rows of small spines (Fig. 14.170g). Pereopods without or with few small spines on carpus and propodus. Thoracic sternite 4 anterior extension smooth������������������������������������������������������������������������������������������������������������������������������������������������������������������������ Fizeserenia Anterior concavities divided and surrounded by clusters of stout spines (Fig. 14.171a). Pereopods with stout spines on carpus and propodus (Fig. 14.172o). Thoracic sternite 4 anterior extension granular������������������ Pelycomaia Carapace pterygostomial region fused to remaining carapace. Anterior carapace with deep V-shaped groove (Fig. 14.170d)�����������������������������������������������������������������������������������������������������������������������������������������������Dacryomaia Carapace pterygostomial region separate from remaining carapace (Fig. 14.171k). Anterior carapace without V-shaped groove (Fig. 14.171j)�����������������������������������������������������������������������������������������������������������������������Zibrovia Carapace vase-shaped (Figs 14.170p–r, 14.171f)��������������������������������������������������������������������������������������������������������������������������7 Carapace widest at midlength or not vase-shaped (Figs 14.170b, n, o, 14.171d)�������������������������������������������������������������������10 Carapace mesogastric region slightly inflated, flanked by longitudinal grooves (Fig. 14.170q)�����������Neotroglocarcinus Carapace mesogastric region ill-defined��������������������������������������������������������������������������������������������������������������������������������������8 Carapace pterygostomial region separate from remaining carapace (Fig. 14.171g). Thoracic sternite 4 anterior extension smooth (Fig. 14.172g)��������������������������������������������������������������������������������������������������������Troglocarcinus Carapace pterygostomial region fused to remaining carapace (Fig. 14.170c). Thoracic sternite 4 anterior extension tuberculate (Fig. 14.172b)�����������������������������������������������������������������������������������������������������������������������������������������9 Pereopods 2–5 meri with row of 4 or 5 spines along extensor margin proximal to distal tooth (Fig. 14.172n). Slope depths������������������������������������������������������������������������������������������������������������������������������������������������������������������� Luciades Pereopods 2–5 meri smooth or with denticles proximal to distal tooth or angle. Subtidal�����������������������������Opecarcinus Thoracic sternite 4 anterior extension broader than long, with tuberculate margin (Fig. 14.172d). Carapace sparsely tuberculate, setose (Fig. 14.170n)������������������������������������������������������������������������������������������������������� Kroppcarcinus

Fig. 14.171.  Cryptochiridae. Carapace, dorsal and lateral views: a, Pelycomaia; b, Pseudocryptochirus; c, Pseudohapalocarcinus; d, e, Sphenomaia; f, g, Troglocarcinus; h, Utinomiella; i, Xynomaia; j, k, Zibrovia.

798

Marine Decapod Crustacea



Thoracic sternite 4 anterior extension prominent, longer than wide (Fig. 14.172b). Carapace coarsely tuberculate or spinose���������������������������������������������������������������������������������������������������������������������������������������������������������������11 11. Carapace mesogastric region inflated, densely tuberculate (Fig. 14.170b, c)�����������������������������������������������������Cryptochirus – Carapace mesogastric region not inflated, with isolated tubercles (Figs 14.170o, 14.171d, e)������������������������������������������� 12 12. Thoracic sternite 4 anterior extension smooth (Fig. 14.172e)��������������������������������������������������������������������������������Lithoscaptus – Thoracic sternite 4 anterior extension tuberculate�������������������������������������������������������������������������������������������������Sphenomaia 13. Carapace widest anterior to midlength (Fig. 14.170i)������������������������������������������������������������������������������������������������� Fungicola – Carapace widest at or posterior to midlength����������������������������������������������������������������������������������������������������������������������������14 14. Carapace less than 1.1 times as long as wide (Figs 14.170e, k, 14.171c). Thoracic sternite 4 anterior extension smooth����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������15 – Carapace more than 1.3 times as long as wide (Figs 14.170a, l, 14.171b, h, i). Thoracic sternite 4 anterior extension tuberculate or smooth��������������������������������������������������������������������������������������������������������������������������������������������17

Fig. 14.172.  Cryptochiridae. Thoracic sternum: a, Cecidocarcinus (sternite 4); b, Cryptochirus; c, Fungicola; d, Kroppcarcinus (sternite 4); e, Lithoscaptus; f, Pseudocryptochirus; g, Troglocarcinus (sternite 4). Maxilliped 3: h, Cryptochirus; i, Utinomiella. Pereopod 2: j, Cecidocarcinus; k, Cryptochirus; l, Hapalocarcinus; m, Neotroglocarcinus; n, Luciades; o, Pelycomaia; p, Pseudocryptochirus; q, Pseudohapalocarcinus; r, Sphenomaia; s, Xynomaia.

14 – Brachyura – crabs

799

15. – 16. – 17. – 18.

Carapace tuberculate (Fig. 14.170e). Maxilliped 3 without exopod���������������������������������������������������������������������Detocarcinus Carapace smooth (Figs 14.170k, 14.171c). Maxilliped 3 with exopod������������������������������������������������������������������������������������16 Pereopod 2 merus little longer than wide, tuberculate (Fig. 14.172q)�������������������������������������������������Pseudohapalocarcinus Pereopod 2 merus 4 times as long as wide, smooth (Fig. 14.172l)������������������������������������������������������������������ Hapalocarcinus Maxilliped 3 without exopod (Fig. 14.172i). Carapace pyriform (Fig. 14.171h)��������������������������������������������������Utinomiella Maxilliped 3 with exopod. Carapace oval (Figs 14.170a, l, 14.171b, i)������������������������������������������������������������������������������������18 Thoracic sternite 4 anterior extension tuberculate; sternite 7 with median suture. Carapace cardiac region prominent (Fig. 14.170l, m)���������������������������������������������������������������������������������������������������������������������������������������������� Hiroia – Thoracic sternite 4 anterior extension smooth (Fig. 14.172a) or with few granules; sternite 7 without (Fig. 14.172c) or with incomplete median suture (Fig. 14.172f). Carapace evenly convex in profile���������������������������19 19. Pereopod 2 merus flexor margin convex, without distal angle, extensor margin without distomesial lobe overlapping carpus (Fig. 14.172s)������������������������������������������������������������������������������������������������������������������������������Xynomaia – Pereopod 2 merus flexor margin expanded, with sharp distal angle, extensor margin with distomesial lobe overlapping carpus (Fig. 14.172j, p)��������������������������������������������������������������������������������������������������������������������������������������� 20 20. Thoracic sternite 4 anterior extension slightly granular (Fig. 14.172f)��������������������������������������������������� Pseudocryptochirus – Thoracic sternite 4 anterior extension smooth, setose (Fig. 14.172a)�������������������������������������������������������������� Cecidocarcinus Cecidocarcinus Kropp & Manning, 1987 Diagnosis. Carapace widest posterior to midlength, lateral margins convex, not deflected anteriorly; pterygostomian region separate from carapace. Thoracic sternite 4 anterior extension smooth, setose; 7 with incomplete median suture. Pereopod 2 merus extensor margin expanded, with sharp distal angle. Pereopod 2 merus with distomesial margin expanded, overlapping base of carapace. Maximum cl. 5.2 mm. Slope (425–512 m). Central Indo-Pacific (New Caledonia), Temperate Southern Africa (Valdivia Ridge). 2 species (Kropp and Manning 1987; Manning 1991).

Cryptochirus Heller, 1861 Diagnosis. Carapace widest at midlength, deflected anteriorly; mesogastric region inflated, densely tuberculate. Thoracic sternite 4 anterior extension tuberculate. Maximum cl. 4 mm. Intertidal, shelf (0–30 m). Indo-West Pacific. 2 species (Kropp 1990).

Dacryomaia Kropp, 1990 Diagnosis. Carapace widest anterior to midlength, deflected anteriorly; anterior carapace with deep V-shaped groove. Thoracic sternite 4 anterior extension tuberculate. Maximum cl. 4.5 mm (Pl. 68b). Subtidal, shelf (1–60 m). Central Indo-Pacific. 2 species (Kropp 1990).

Detocarcinus Kropp & Manning, 1987 Diagnosis. Carapace about as long as wide, widest posterior to midlength, lateral margins convex, not deflected anteriorly. Thoracic sternite 4 anterior extension with few granules; 7 without median suture. Maxilliped 3 without exopod. Maximum cl. 4.7 mm.

Subtidal, shelf (3–98 m). Tropical W Atlantic. 1 species (Kropp and Manning 1987; van der Meij and Nieman 2016: phylogenetic position).

Fizesereneia Takeda & Tamura, 1980 Diagnosis. Carapace widest anterior to midlength, deflected anteriorly; anterior carapace with 2 bowl-shaped concavities divided and surrounded by 1 or 2 rows of small spines; pterygostomian region separate from or fused to carapace. Thoracic sternite 4 anterior extension smooth. Maximum cl. 8.3 mm (Pl. 68c). Intertidal, subtidal (0–15 m). Western and Central IndoPacific. 7 species (Kropp 1990, 1994; van Der Meij et al. 2015: species compared).

Fungicola Serène, 1966 Diagnosis. Carapace widest anterior to midlength, not deflected anteriorly. Thoracic sternite 4 anterior extension with few granules. Maximum cl. 5.5 mm (Pl. 68d, e). Subtidal (1–15 m). Central Indo-Pacific. 3 species (Kropp 1990; van der Meij 2015b: cryptic species).

Hapalocarcinus Stimpson, 1859 Diagnosis. Carapace about as long as wide, widest posterior to midlength, lateral margins convex, not deflected anteriorly; mesogastric region not defined, almost smooth. Thoracic sternite 4 anterior extension smooth; 7 without median suture. Pereopod 2 merus 4 times as long as wide, smooth. Maximum cl. 4.5 mm (Pl. 68i). Intertidal–shelf (0–35 m). Indo-West Pacific, Tropical Eastern Pacific. 1 species (Kropp 1990; Bähr et  al. 2021: species complex).

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Marine Decapod Crustacea

Hiroia Takeda & Tamura, 1981 Diagnosis. Carapace widest at midlength, not deflected anteriorly; cardiac region prominent. Thoracic sternite 4 anterior extension tuberculate. Maximum cl. 3.7 mm. Subtidal (1–19 m). Central Indo-Pacific. 1 species (Kropp 1990).

Kroppcarcinus Badaro, Neves, Castro & Johnsson, 2012 Diagnosis. Carapace widest at midlength, deflected anteriorly; regions ill-defined, sparsely tuberculate, setose; pterygostomian region separate from carapace. Thoracic sternite 4 anterior extension broader than long, with tuberculate margin. Maximum cl. 3.5 mm. Subtidal, shelf (1–40 m). Tropical Atlantic (Brazil). 1 species (Badaro et al. 2012).

Intertidal–shelf (0–82 m). Tropical Atlantic, Indo-West Pacific. 9 species (Chan et  al. 2020: biogeography; Kropp 1989: key to 8 species; Kropp and Manning 1996; van Der Meij 2014).

Pelycomaia Kropp, 1990 Diagnosis. Carapace widest anterior to midlength, deflected anteriorly; anterior carapace with 2 bowl-shaped concavities concavities divided and surrounded by clusters of stout spines. Thoracic sternite 4 anterior extension tuberculate. Maximum cl. 4 mm. Subtidal (2 m). Central and Eastern Indo-Pacific. 1 species (Kropp 1990).

Pseudocryptochirus Hiro, 1938

Diagnosis. Carapace widest at midlength, deflected anteriorly. Thoracic sternite 4 anterior extension smooth. Maximum cl. 4.4 mm (Pl. 68h). Intertidal–shelf (0–25 m). Western and Central Indo-Pacific. 10 species (Kropp 1990; van der Meij 2017: species compared).

Diagnosis. Carapace widest posterior to midlength, lateral margins convex, not deflected anteriorly; mesogastric region not defined, almost smooth; pterygostomian region separate from carapace. Thoracic sternite 4 anterior extension tuberculate; 7 with incomplete median suture. Pereopod 2 merus extensor margin expanded, with sharp distal angle. Pereopod 2 merus with distomesial margin expanded, overlapping base of carapace. Maximum cl. 4 mm (Pl. 68g). Subtidal, shelf (1–30 m). Central Indo-Pacific. 1 species (Kropp 1990).

Luciades Kropp & Manning, 1996

Pseudohapalocarcinus Fize & Serène, 1956

Diagnosis. Carapace widest posterior to midlength, vase-shaped, deflected anteriorly; anterior carapace with broad W-shaped depression. Thoracic sternite 4 anterior extension tuberculate. Pereopods 2–5 meri with row of 4 or 5 spines along extensor margin proximal to distal tooth. Pereopod 2 merus with distomesial margin expanded, overlapping base of carapace. Maximum cl. 2.3 mm. Shelf (128–137 m). Central Indo-Pacific (Guam). 1 species (Kropp and Manning 1996).

Diagnosis. Carapace about as long as wide, widest posterior to midlength, lateral margins convex, not deflected anteriorly; mesogastric region not defined, almost smooth. Thoracic sternite 4 anterior extension smooth; 7 without median suture. Pereopod 2 merus little longer than wide, tuberculate. Maximum cl. 3.3 mm. Subtidal (1–21 m). Central Indo-Pacific. 1 species (Kropp 1990).

Lithoscaptus A. Milne-Edwards, 1862

Sphenomaia Kropp, 1990 Neotroglocarcinus Takeda & Tamura, 1980 Diagnosis. Carapace widest posterior to midlength, vase-shaped, deflected anteriorly; mesogastric region slightly inflated, flanked by longitudinal grooves. Thoracic sternite 4 anterior extension tuberculate. Pereopod 2 merus with distomesial margin expanded, overlapping base of carapace. Maximum cl. 3.5 mm. Subtidal (1–13 m). Central Indo-Pacific. 2 species (Kropp 1990).

Opecarcinus Kropp & Manning, 1987 Diagnosis. Carapace widest posterior to midlength, vase-shaped, deflected anteriorly. Thoracic sternite 4 anterior extension tuberculate. Pereopods 2–5 meri with or without denticles proximal to distal tooth or angle. Pereopod 2 merus with distomesial margin expanded, overlapping base of carapace, or with distomesial and distolateral margins similar, neither overlapping base of carapace. Maximum cl. 4 mm (Pl. 68f).

Diagnosis. Carapace widest at midlength, deflected anteriorly. Thoracic sternite 4 anterior extension tuberculate; 7 without median suture. Subtidal (1–5 m). Eastern Indo-Pacific (Kiribati). 1 species (Kropp 1990).

Troglocarcinus Verrill, 1908 Diagnosis. Carapace widest posterior to midlength, vase-shaped, deflected anteriorly; pterygostomian region separate from carapace. Thoracic sternite 4 anterior extension smooth. Maximum cl. 5.2 mm. Subtidal, shelf (1–75 m). Tropical Atlantic. 2 species (Canário et al. 2015: rediagnosis; Kropp and Manning 1987).

Utinomiella Kropp & Takeda, 1988 Diagnosis. Carapace widest posterior to midlength, pear shaped, not deflected anteriorly. Thoracic sternite 4 anterior extension

14 – Brachyura – crabs

tuberculate; 7 with median suture. Maxilliped 3 without exopod. Maximum cl. 4.5 mm (Pl. 68j). Intertidal–shelf (0–29 m). Indo-West Pacific. 1 species (Kropp 1990).

Xynomaia Kropp, 1990 Diagnosis. Carapace widest posterior to midlength, lateral margins convex, not deflected anteriorly; anterior carapace with broad W-shaped depression; pterygostomian region separate from carapace. Thoracic sternite 4 anterior extension smooth; 7 without median suture. Pereopod 2 merus extensor margin convex, without distal angle. Maximum cl. 5 mm. Intertidal–shelf (0–25 m). Central Indo-Pacific. 3 species (Kropp 1990).

Zibrovia Kropp & Manning, 1996 Diagnosis. Carapace widest anterior to midlength, deflected anteriorly; pterygostomian region separate from carapace. Thoracic sternite 4 anterior extension tuberculate. Maximum cl. 3.5 mm. Shelf (81–100 m). Western and Central Indo-Pacific. 1 species (Kropp and Manning 1996). References Badaro MFS, Neves EG, Castro P, Johnsson R (2012) Description of a new genus of Cryptochiridae (Decapoda: Brachyura) associated with Siderastrea (Anthozoa: Scleractinia), with notes on feeding habits. Scientia Marina 76, 517–526. doi:10.3989/scimar.03538.02E Bähr S, Johnson ML, Berumen ML, Hardenstine RS, Rich WA, et  al. (2021) Morphology and reproduction in the Hapalocarcinus marsupialis Stimpson, 1859 species complex (Decapoda: Brachyura: Cryptochiridae). Journal of Crustacean Biology 41, ruab052. doi:10.1093/ jcbiol/ruab052 Canário R, Badaró MFS, Johnsson R, Neves EG (2015) A new species of Troglocarcinus (Decapoda: Brachyura: Cryptochiridae) symbiotic with the Brazilian endemic coral Mussismilia (Anthozoa: Scleractinia: Mussidae). Marine Biology Research 11, 76–85. doi:10.1080/17451000.2 014.894243 Chan B, Wong KJH, Cheng Y-R (2020) Biogeography and host usage of coral-associated crustaceans: barnacles, copepods, and gall crabs as model organisms. In The Natural History of the Crustacea. (Eds Poore GCB, Thiel M) pp. 183–215. Oxford University Press, New York. Hoeksema BW, van der Meij SET, Fransen CHJM (2012) The mushroom coral as a habitat. Journal of the Marine Biological Association of the United Kingdom 92, 647–663. doi:10.1017/S0025315411001445 Kropp RK (1986) Feeding biology and mouthpart morphology of three species of coral gall crabs (Decapoda: Cryptochiridae). Journal of Crustacean Biology 6, 377–384. doi:10.2307/1548178 Kropp RK (1989) A revision of the Pacific species of gall crabs, genus Opecarcinus (Crustacea: Cryptochiridae). Bulletin of Marine Science 45, 98–129. Kropp RK (1990) Revision of the genera of gall crabs (Crustacea: Cryptochiridae) occurring in the Pacific Ocean. Pacific Science 44, 417–448. Kropp RK (1994) The gall crabs (Crustacea: Decapoda: Brachyura: Cryptochiridae) of the Rumphius Expeditions revisited, with descriptions of three new species. Raffles Bulletin of Zoology 42, 521–538. Kropp RK, Manning RB (1987) The Atlantic gall crabs, family Cryptochiridae (Crustacea: Decapoda: Brachyura). Smithsonian Contributions to Zoology 462, 1–21. doi:10.5479/si.00810282.462

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Kropp RK, Manning RB (1996) Crustacea Decapoda: two new genera and species of deep water gall crabs from the Indo-west Pacific (Cryptochiridae). In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 15. Mémoires du Muséum National d’Histoire Naturelle, Paris 168, 531–539. Manning RB (1991) Crustacea Decapoda: Cecidocarcinus zibrowii, a new deep water gall crab (Cryptochiridae) from New Caledonia. In: Crosnier A (Ed.), Résultats des Campagnes MUSORSTOM, vol. 9. Mémoires du Muséum National d’Histoire Naturelle, Paris 152, 515–520. van Der Meij SE (2014) A new species of Opecarcinus Kropp & Manning, 1987 (Crustacea: Brachyura: Cryptochiridae) associated with the stony corals Pavona clavus (Dana, 1846) and P. bipartita Nemenzo, 1980 (Scleractinia: Agariciidae). Zootaxa 3869, 44–52. doi:10.11646/ zootaxa.3869.1.4 van der Meij SET (2015a). Evolutionary diversification of coral-dwelling gall crabs (Cryptochiridae). PhD thesis. Leiden University, Leiden. van der Meij SET (2015b) Host relations and DNA reveal a cryptic gall crab species (Crustacea: Decapoda: Cryptochiridae) associated with mushroom corals (Scleractinia: Fungiidae). Contributions to Zoology 84, 39–57. doi:10.1163/18759866-08401004 van der Meij SET (2017) The coral genus Caulastraea Dana, 1846 (Scleractinia, Merulinidae) as a new host for gall crabs (Decapoda, Cryptochiridae), with the description of Lithoscaptus tuerkayi sp.nov. Crustaceana 90, 1027–1038. doi:10.1163/15685403-00003607 van der Meij SET, Nieman AM (2016) Old and new DNA unweave the phylogenetic position of the eastern Atlantic gall crab Detocarcinus balssi (Monod, 1956) (Decapoda: Cryptochiridae). Journal of Zoological Systematics and Evolutionary Research 54, 189–196. doi:10.1111/jzs.12130 van der Meij SET, Schubart CD (2014) Monophyly and phylogenetic origin of the gall crab family Cryptochiridae (Decapoda : Brachyura). Invertebrate Systematics 28, 491–500. doi:10.1071/IS13064 van Der Meij SE, Berumen ML, Paulay G (2015) A new species of Fizesereneia Takeda & Tamura, 1980 (Crustacea: Brachyura: Cryptochiridae) from the Red Sea and Oman. Zootaxa 3931, 585–595. doi:10.11646/ zootaxa.3931.4.8 Zayasu Y, Nomura K, Seno K, Asakura A (2013) A new species of Fizesereneia Takeda & Tamura, 1980 (Crustacea: Decapoda: Brachyura: Cryptochiridae) from Japan. Zootaxa 3681, 257–269. doi:10.11646/ zootaxa.3681.3.5

Grapsoidea MacLeay, 1838 Common shore crabs are possibly the most familiar of all crabs. Davie et al. (2015a) reviewed the taxonomic history of various family names and conflicting adult and larval morphological, molecular evidence. Davie et  al. (2015b) diagnosed the superfamily and eight families. One family has been added since by Guinot et al. (2018) who diagnosed this and some other families, and discussed similarities and relationships extensively. The two major thoracotreme families, Grapsoidea and Ocypodoidea, are currently acknowledged as not monophyletic. Most phylogenetic studies reveal significant polyphyly of both superfamilies (Schubart et al. 2006; Tsang et al. 2018; Tsang et al. 2014), although studies to date are yet to be sufficiently comprehensive to provide a large enough framework on which a new classification could be based. As such, both superfamilies are used herein pending phylogenetic revision.

802

Marine Decapod Crustacea

The superfamily Grapsoidea includes nine families of which seven are almost exclusively marine, especially in intertidal or supralittoral environments but are known too from much deeper waters or damp forests. The remaining two families, Gecarcinidae and Glyptograpsidae, which are diverse in subtropical and tropical freshwater streams and in forests (Schubart et  al. 2000), are not dealt with further here. The key to families of Brachyura includes Gecarcinidae and Glyptograpsidae but they are not in the key below. Gecarcinidae MacLeay, 1838. Guinot et al. (2018) recognised two clades, and diagnosed and provided a key to three genera of the first clade: Cardisoma Latreille, 1828, Discoplax A. Milne-Edwards, 1867 and Tuerkayana Guinot, N.K.  Ng & Rodrígues Moreno, 2018. The second clade includes Epigrapsus HelIer, 1862, Gecarcinus Leach, 1814, Gecarcoidea H. Milne Edwards, 1837 and Johngarthia ­Türkay, 1970, each diagnosed by Guinot et al. (2018). All are

essentially pantropical terrestrial crabs that migrate towards the sea to release zoeae. Glyptograpsidae Schubart, Cuesta & Felder, 2002. Two genera of freshwater crabs from streams in Central America are included, Glyptograpsus Smith, 1870 and Platychirograpsus De Man, 1896. Some species range into brackish environments at stream mouths. Schubart et al. (2002) diagnosed the family on adult and larval morphological characters, and molecular differences. The genera and species were differentiated by Rathbun (1918). The seven remaining grapsoid families include 37 primarily marine genera, some of which include semiterrestrial species or others ranging into tropical forests. Grapsodius Holmes, 1900 is a poorly known monotypic genus, included by Rathbun (1918) in her synthesis of American grapsoids. We do not include it here as it has not been seen since and may be based on an aberrant specimen (Cuesta and Schubart 1999).

Key to marine families of Grapsoidea 1. – 2. – 3. – 4. – 5. – 6. –

Carapace frontal usually margin with deep cleft on each side of midline to receive antennules. Carapace distinctly subcircular to square, usually about as long as wide (Figs 14.175, 14.176)�������������������������������������������������������2 Carapace frontal margin without deep cleft to receive antennules. Carapace ovate to subquadrate���������������������������������3 Maxilliped 3 merus as wide as or slightly wider than ischium (Fig. 14.176k, l). Male pleonites 3–6 fused��������Plagusiidae Maxilliped 3 merus narrower than ischium (Fig. 14.175b). Male pleonites 3–5 fused������������������������������������������ Percnidae Orbit totally closed. Maxillipeds 3 with mesial margins meeting when closed; ischium and merus together subrectangular (Fig. 14.190). Associated with hydrothermal vents������������������������������������������� Xenograpsidae Orbit laterally open. Maxillipeds 3 separated by rhomboidal or small gap when closed; ischium and merus longer than wide. Supratidal to subtidal���������������������������������������������������������������������������������������������������������������������4 Carapace ovate, with well marked cervical grooves oblique to midline, strongly converging posteriorly, terminating at cardiac region. Setose pouches between coxae of pereopods 2–4 (Fig. 14.174g); pereopods 2–5 dactyli with strong, fixed chitinous spines (Fig. 14.174e)��������������������������������������������Leptograpsodidae Carapace subquadrate–polygonal, usually smooth. Without setose pouches between coxae of pereopods 2–4. Pereopods 2–5 dactyli without strong, fixed chitinous spines������������������������������������������������������������������������������������5 Suborbital crest granulate or dentate, stridulatory (Fig. 14.183) (except Pinnotherelinae, Fig. 14.185)�������������Varunidae Suborbital crest smooth or with few granules�����������������������������������������������������������������������������������������������������������������������������6 Carapace pterygostomial region covered in reticulated network of short, hooked, setae (Fig. 14.179r)��������� Sesarmidae Carapace pterygostomial region sparingly setose, without pattern of reticulated setae (Fig. 14.173)����������������Grapsidae

References Cuesta JA, Schubart CD (1999) First zoeal stages of Geograpsus lividus and Goniopsis pulchra from Panama confirm consistent larval characters for the subfamily Grapsinae (Crustacea: Brachyura: Grapsidae). Ophelia 51, 163–176. doi:10.1080/00785326.1999.10409406 Davie PJF, Guinot D, Ng PKL (2015a) Chapter 71–16. Phylogeny of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 921–979. Brill, Leiden. Davie PJF, Guinot D, Ng PKL (2015b) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy,

Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. Guinot D, Ng NK, Rodrígues Moreno PA (2018) Review of grapsoid families for the establishment of a new family for Leptograpsodes Montgomery, 1931, and a new genus of Gecarcinidae H. Milne Edwards, 1837 (Crustacea, Decapoda, Brachyura, Grapsoidea MacLeay, 1838). Zoosystema 40, 547–604. doi:10.5252/zoosystema2018v40a26 Rathbun MJ (1918) The grapsoid crabs of America. Bulletin of the United States National Museum 97, 1–461, pls 1–161. Schubart CD, Cuesta JA, Diesel R, Felder D (2000) Molecular phylogeny, taxonomy, and evolution of nonmarine lineages within the American

14 – Brachyura – crabs

grapsoid crabs (Crustacea: Brachyura). Molecular Phylogenetics and Evolution 15, 179–190. doi:10.1006/mpev.1999.0754 Schubart CD, Cuesta JA, Felder DL (2002) Glyptograpsidae, a new brachyuran family from Central America: larval and adult morphology, and a molecular phylogeny of the Grapsoidea. Journal of Crustacean Biology 22, 28–44. doi:10.1163/20021975-99990206 Schubart CD, Cannicci S, Vannini M, Fratini S (2006) Molecular phylogeny of grapsoid crabs (Decapoda, Brachyura) and allies based on two mitochondrial genes and a proposal for refraining from current superfamily classification. Journal of Zoological Systematics and Evolutionary Research 44, 193–199. doi:10.1111/j.1439-0469.2006.00354.x Tsang LM, Schubart CD, Ahyong ST, Lai JCY, Au EYC, et al. (2014) Evolutionary history of true crabs (Crustacea: Decapoda: Brachyura) and the origin of freshwater crabs. Molecular Biology and Evolution 31, 1173–1187. doi:10.1093/molbev/msu068 Tsang LM, Ahyong ST, Shih H-T, Ng PKL (2018) Further polyphyly of pinnotheroid crabs: the molecular phylogenetic position of the polychaete-associated Aphanodactylidae. Invertebrate Systematics 32, 92–99. doi:10.1071/IS17038

Grapsidae MacLeay, 1838 Figure 14.173, Plate 69a–e Grapsids are the archetypal shore crab of tropical to temperate environments. Several inhabit rocky or boulder shores exposed to wave action where they are very active scavengers or herbivores. Some spend considerable time out of the water and can move extremely quickly, especially when evading capture! Others burrow in soft sediments supralittorally or

803

intertidally. Grapsids can be recognised by features of the oral area in the combination of a minimally setose pterygostomial region, a distinct rhomboidal gap between the maxillipeds 3 of which the ischium lacks an oblique setose ridge. The once large family has been restricted to just seven genera after reappraisal of adult and larval morphology, and phylogenetics (Schubart 2011; Schubart et al. 2006a). Multilocus molecular phylogenetic analyses of the family recovered most genera as monophyletic, but in the case of the morphologically heterogeneous Pachygrapsus, earlier suspicions of polyphyly were borne out (Ip et al. 2015). Crosnier (1965) provided a key to five Indo-West Pacific genera. Rathbun (1918) dealt with the American genera. Diagnosis. Carapace subquadrate to rounded-trapezoid, smooth; front entire; orbit with lateral opening; suborbital crest smooth, without granules; pterygostomial region sparingly setose, without pattern of reticulated setae. Maxilliped 3 merus and ischium without oblique setose ridge; distinct rhomboidal gap between closed maxillipeds 3. Mandibles usually visible when mouthparts closed. Chelipeds similar. Male pleonites 1–6 and telson free. Implicit generic attributes. Carapace trapezoid, almost flat, laterraly with numerous transverse lines; width of front greater than half of carapace width. Antenna included in orbit. Chelipeds with spoon-shaped finger tips. Pereopods 2–5 flat, spinose.

Key to genera of Grapsidae 1. – 2. – 3. – 4. – 5. – 6. –

Front less than half width of carapace (Fig. 14.173a–c)��������������������������������������������������������������������������������������������������������������2 Front half width of carapace or more (Fig. 14.173d–f)���������������������������������������������������������������������������������������������������������������4 Carapace with 2 anterolateral teeth behind exorbital angle (Fig. 14.173c)���������������������������������������������������������Leptograpsus Carapace with 1 anterolateral tooth behind exorbital angle (Fig. 14.173a, b)�������������������������������������������������������������������������3 Chelipeds with spoon-shaped finger tips (Fig. 14.173m)�����������������������������������������������������������������������������������������Geograpsus Chelipeds with acute finger tips���������������������������������������������������������������������������������������������������������������������������������������� Grapsus Antenna excluded from orbit by tooth at lower inner angle (Fig. 14.173h)�����������������������������������������������������������������������������5 Antenna included in orbit (Fig. 14.173i)���������������������������������������������������������������������������������������������������������������������������������������6 Maxilliped 3 merus wider than long. Indo-West Pacific���������������������������������������������������������������������������������� Metopograpsus Maxilliped 3 merus longer than wide (Fig. 14.173k). Atlantic, East Pacific�������������������������������������������������������������Goniopsis Carapace subcircular or subquadrate, surface convex, smooth, with weak striations (Fig. 14.173f). Pereopods 2–5 propodus with dense fringe of setae along upper margin (Fig. 14.173p). Pelagic������������������������ Planes Carapace trapezoidal or subquadrate, surface almost flat, rough, with marked striations (Fig. 14.173e). Pereopods 2–5 propodus with scattered setae on upper margin or thick patch on surface (Fig. 14.173o), not a fringe along upper margin. Benthic��������������������������������������������������������������������������������������������������������� Pachygrapsus Geograpsus Stimpson, 1858

Diagnosis. Carapace subcircular to rounded-subquadrate; front narrower than half carapace width; with 1 anterolateral tooth behind exorbital angle. Maxilliped 3 merus longer than wide. Chelipeds massive, unequal, fingers with acute tips. Maximum cl. 40 mm.

Terrestrial above shore line, nocturnal. Indo-West Pacific. 6 species (Banerjee 1960: diagnosis, key to 3 Indo-West Pacific species; Rathbun 1918: American species). One species has probably gone extinct in Hawaii (Paulay and Starmer 2011).

804

Marine Decapod Crustacea

Fig. 14.173.  Grapsidae. Carapace: a, Geograpsus; b, Grapsus; c, Leptograpsus; d, Metopograpsus; e, Pachygrapsus; f, Planes. Front, orbit, antenna: g, Leptograpsus; h, Metopograpsus; i, Pachygrapsus. Maxilliped 3: j, Geograpsus; k, Goniopsis; l, Pachygrapsus. m, cheliped fingers, Pachygrapsus. Pereopod: n, Geograpsus; o, Pachygrapsus; p, Planes.

Goniopsis De Haan, 1833 mangrove crab Diagnosis. Carapace trapezoidal or subquadrate, almost flat; front wider than half carapace width. Antenna excluded from orbit by tooth at lower inner angle. Maxilliped 3 merus longer than wide. Maximum cl. 50 mm (Pl. 69a). Estuarine, mangroves, muddy sediments. Tropical Atlantic, Tropical Eastern Pacific. 3 species (Manning and Holthuis 1981; von Sternberg 1994). Mangrove crabs are more terrestrial than marine (viz e.g. Pereira Rodrigues de Lira et al. 2013).

Grapsus Lamarck, 1801 Diagnosis. Carapace subcircular to rounded-subquadrate; front narrower than half carapace width; with 1 anterolateral tooth behind exorbital angle. Maxilliped 3 merus longer than wide. Maximum cl. 56 mm (Pl. 69b, c). Intertidal, rocky shores. Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific. 8 species (Banerjee 1960: key to 4 Indo-West Pacific species; Rathbun 1918: American species). The genus includes the very active Sally-Lightfoot, Grapsus grapsus (Linnaeus, 1758), seen running on American rocky shores (Teschima et al. 2016).

Leptograpsus H. Milne Edwards, 1853 Diagnosis. Carapace subcircular to rounded-subquadrate, frontal region granulate, narrower than half carapace width; gastric region with 2 parallel longitudinal grooves, cardiac region with 1

transverse groove, 8–10 oblique ridges laterally; with 2 anterolateral teeth behind exorbital angle. Maxilliped 3 merus wider than long. Maximum cl. 70 mm. Intertidal. Western and Central Indo-Pacific, Temperate South America, Temperate Southern Africa, Temperate Australasia. 1 species (Griffin 1973).

Metopograpsus H. Milne Edwards, 1853 Diagnosis. Carapace trapezoidal, almost flat; front wider than half carapace width. Antenna excluded from orbit by tooth at lower inner angle. Maxilliped 3 merus wider than long. Maximum cl. 39 mm. Intertidal, mangroves, above water-line. Indo-West Pacific. 6 species (Banerjee 1960: key to species; Fratini et al. 2018: distribution, phylogeny). Paulay (2007) suspected M. oceanicus (Hombron & Jacquinot, 1846) to be a recent invasive of Guam and Hawaii.

Pachygrapsus Randall, 1840 Diagnosis. Carapace trapezoid, sides converging posteriorly, almost flat, with numerous transverse striations; front wider than half carapace width. Antenna included in orbit. Maxilliped 3 merus wider than long. Pereopods 2–5 propodi with few scattered setae on upper margin or thick patch on surface (Fig. 14.173o), not a fringe along upper margin. Maximum cl. 37 mm (Pl. 69e). Intertidal, rubble, algae, rocks, mangroves. Cosmopolitan except polar seas. 14 species (Poupin et al. 2005: diagnosis, key to 13 species; Schubart et  al. 2006b: phlylogeography of American species).

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Planes Bowdich, 1825 Columbus crab Diagnosis. Carapace subcircular to subquadrate, very convex, smooth, with weak striations; front as wide as or wider than half carapace width. Antenna included in orbit. Maxilliped 3 merus wider than long. Pereopods 2–5 propodi, dactyli with dense fringe of setae along upper margin. Maximum cl. 25 mm (Pl. 69d). Widespread in all tropical and temperate oceans on pelagic seaweeds or crawling on floating objects. 2 species (Prado and de Melo 2002). The unusual pelagic behaviour of species of Planes has attracted considerable research interest. Species are typically found on drifting macroalgae such as Sargassum but are also associated with turtles (Pfaller et  al. 2019) and litter (Tutman et  al. 2017). References Banerjee SK (1960) Biological results of the Snellius Expedition. XVIII. The genera Grapsus, Geograpsus, and Metopograpsus. Temminckia 10, 132–199. Crosnier A (1965) Crustacés Décapodes Grapsidae et Ocypodidae. Faune de Madagascar 18, 1–143, pls 1–11. Fratini S, Cannicci S, Schubart CD (2018) Molecular phylogeny of the crab genus Metopograpsus H. Milne Edwards, 1853 (Decapoda : Brachyura : Grapsidae) reveals high intraspecific genetic variation and distinct evolutionarily significant units. Invertebrate Systematics 32, 215–223. doi:10.1071/IS17034 Griffin DJG (1973) A revision of the two southern temperate shore crabs Leptograpsus variegatus (Fabricius) and Plagusia chabrus (Linnaeus) (Crustacea, Decapoda, Grapsidae). Journal of the Royal Society of New Zealand 3, 415–440. doi:10.1080/03036758.1973.104 21866 Ip BHY, Schubart CD, Tsang LM, Chu KH (2015) Phylogeny of the shore crab family Grapsidae (Decapoda: Brachyura: Thoracotremata) based on a multilocus approach. Zoological Journal of the Linnean Society 174, 217–227. doi:10.1111/zoj.12235 Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Paulay G (2007) Metopograpsus oceanicus (Crustacea: Brachyura) in Hawai’i and Guam: another recent invasive? Pacific Science 61, 295–300. doi:10.2984/1534-6188(2007)61[295:MOCBIH]2.0.CO;2

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Paulay G, Starmer J (2011) Evolution, insular restriction, and extinction of oceanic land crabs, exemplified by the loss of an endemic Geograpsus in the Hawaiian Islands. PLoS One 6, e19916. doi:10.1371/journal. pone.0019916 Pereira Rodrigues de Lira JJ, dos Santos Calado TC, Leitão Câmara de Araújo Md S (2013) Breeding period in the mangrove crab Goniopsis cruentata (Decapoda: Grapsidae) in northeast Brazil. Revista de Biología Tropical 61, 29–38. doi:10.15517/rbt.v61i1.10878 Pfaller JB, Palau AN, Agamboué PD, Barret M, Ciccione S, et al. (2019) New records of Planes crabs associated with sea turtles in Africa and adjacent waters. African Journal of Marine Science 41, 331–336. doi:10. 2989/1814232X.2019.1651394 Poupin J, Davie PJF, Cexus JC (2005) A revision of the genus Pachygrapsus Randall, 1840 (Crustacea: Decapoda: Brachyura, Grapsidae), with special reference to the Southwest Pacific species. Zootaxa 1015, 1–66. doi:10.11646/zootaxa.1015.1.1 Prado A, de Melo GAS (2002) Le genre Planes Bowdich (Decapoda, Grapsidae) sur les côtes brésilliennes (the genus Planes Bowdich (Decapoda, Grapsidae) along the Brazilian coast). Crustaceana 75, 579–595. doi:10.1163/156854002760095615 Rathbun MJ (1918) The grapsoid crabs of America. Bulletin of the United States National Museum 97, 1–461, pls 1–161. Schubart CD (2011) Reconstruction of phylogenetic relationships within Grapsidae (Crustacea: Brachyura) and comparison of trans-isthmian versus amphi-atlantic gene flow based on mtDNA. Zoologischer Anzeiger 250, 472–478. doi:10.1016/j.jcz.2011.06.003 Schubart CD, Cannicci S, Vannini M, Fratini S (2006a) Molecular phylogeny of grapsoid crabs (Decapoda, Brachyura) and allies based on two mitochondrial genes and a proposal for refraining from current superfamily classification. Journal of Zoological Systematics and Evolutionary Research 44, 193–199. doi:10.1111/j.1439-0469.2006.00354.x Schubart CD, Cuesta JA, Felder DL (2006b) Phylogeography of Pachygrapsus transverses (Gibbes, 1850): The effect of the American continent and the Atlantic Ocean as gene flow barriers, and recognition of Pachygrapsus socius Stimpson 1871 as a valid species. Nauplius 13, 99–113. Teschima MM, Ströher PR, Firkowski CR, Pie MR, Freire AS (2016) Largescale connectivity of Grapsus grapsus (Decapoda) in the Southwestern Atlantic oceanic islands: integrating genetic and morphometric data. Marine Ecology 37, 1360–1372. doi:10.1111/maec.12347 Tutman P, Kapiris K, Kirinčić M, Pallaoro A (2017) Floating marine litter as a raft for drifting voyages for Planes minutus (Crustacea: Decapoda: Grapsidae) and Liocarcinus navigator (Crustacea: Decapoda: Polybiidae). Marine Pollution Bulletin 120, 217–221. doi:10.1016/j.marpolbul.​ 2017.04.063

Fig. 14.174.  Leptograpsodidae. Leptograpsodes octodentatus (H. Milne Edwards, 1837). a, carapace; b, orbit, eyestalk, antenna; c, maxilliped 3; d, cheliped; e, pereopod 2, propodus, dactylus; f, male pleon; g, left sternum, pereopods 2–4 coxae.

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von Sternberg R (1994) Systematic implications of color pattern polymorphism in Goniopsis pulchra (Decapoda: Brachyura: Grapsidae) from Ecuador. Proceedings of the Biological Society of Washington 107, 721–728.

Leptograpsodidae Guinot, Ng & Rodriguez Moreno, 2018 Figure 14.174, Plate 69f A separate family was erected for a single species of large semiterrestrial shore crab from southern Australia. It was differentiated from other grapsoids by the possession of pouches full of setae between the coxae of pereopods 2–4, evidence of its terrestrial adaptation (Guinot et  al. 2018), chitinous spines on dactyli of the walking legs and the narrow pleonite 6 of the male. Diagnosis. Carapace ovate, with well marked cervical grooves oblique to midline, strongly converging posteriorly, terminating at cardiac region; front entire; orbit with lateral opening; suborbital crest granulate or dentate, stridulatory. Maxilliped 3 merus and ischium without oblique setose ridge; distinct rhomboidal gap between closed maxillipeds 3. Mandibles usually visible when mouthparts closed. Chelipeds similar. Setose pouches between coxae of pereopods 2–4 (Fig. 14.174g); pereopods 2–5 dactyli with strong, fixed chitinous spines. Male pleonites 1–6 and telson free. Leptograpsodes Montgomery, 1931 burrowing shore crab Maximum cl. 70 mm (Pl. 69f). Supralittoral, rocky shores, often in nearshore freshwater seeps. Temperate Australasia (S, SW Australia). 1 species. The only species, Leptograpsodes octodentatus (H. Milne Edwards, 1837), was redescribed by Griffin (1969), Poore (2004) and Guinot et al. (2018). References Griffin DJG (1969) Notes on the taxonomy and zoogeography of the Tasmanian grapsid and ocypodid crabs (Crustacea, Brachyura). Records of the Australian Museum 27, 323–347. doi:10.3853/j.0067-1975.27.1969.452 Guinot D, Ng NK, Rodrígues Moreno PA (2018) Review of grapsoid families for the establishment of a new family for Leptograpsodes Montgomery, 1931, and a new genus of Gecarcinidae H. Milne Edwards, 1837 (Crustacea, Decapoda, Brachyura, Grapsoidea MacLeay, 1838). Zoosystema 40, 547–604. doi:10.5252/zoosystema2018v40a26 Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne.

Percnidae Števčić, 2005 Figure 14.175, Plate 69g Species of the only genus are rocky-shore crabs found widely in tropical and subtropical shores, more rarely in temperate

Fig. 14.175.  Percnidae. Percnon. a, carapace; b, maxilliped 3.

climes. The best studied species is Percon gibbesi (H. MilneEdwards, 1853) which is invasive and now widespread in the Mediterranean (Katsanevakis et  al. 2011; Sciberras and Schembri 2008). Like plagusiids, species of Percnon have an incised front but differ in having the merus of maxilliped 3 much narrower than the ischium (Schubart and Cuesta 2010). Diagnosis. Carapace distinctly subcircular to square, usually about as long as wide; front with deep cleft on each side of midline to receive antennules; suborbital crest smooth, without granules. Maxilliped 3 merus narrower than ischium; very small rhomboidal gap between closed maxillipeds 3. Mandibles not visible when mouthparts closed. Male cheliped swollen, smooth. Male pleonites 3–5 fused. Percnon Gistel, 1848 spray crabs Maximum cl. 28 mm (Pl. 69g). Intertidal, subtidal (0–15 m; rocky and coralline shores). Cosmopolitan. 7 species (Crosnier 1965: key to 3 species; Poupin 1994: differentiated 3 species; Retamal 2002). References Crosnier A (1965) Crustacés Décapodes Grapsidae et Ocypodidae. Faune de Madagascar 18, 1–143, pls 1–11. Katsanevakis S, Poursanidis D, Yokes MB, Ma ČIĆV, Beqiraj S, et  al. (2011) Twelve years after the first report of the crab Percnon gibbesi (H. Milne Edwards, 1853) in the Mediterranean: current distribution and invasion rates. Journal of Biological Research (Thessaloniki) 16, 224–236. Poupin J (1994) Quelques crustacés décapodes communs de Polynésie française. Rapport Scientifique du Service Mixte de Surveillance Radiologique et Biologique 1–86, figs 1–68, pls 1–8. Retamal MA (2002) Percnon pascuensis una nueva especie de isla de Pascua (Grapsidae, Plagusiinae). Investigaciones Marinas 30, 63–66. doi:10.4067/S0717-71782002000100005 Schubart CD, Cuesta JA (2010) Phylogenetic relationships of the Plagusiidae Dana, 1851 (Brachyura), with description of a new genus and recognition of Percnidae Števčić, 2005, as an independent family. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on

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Species of Plagusia inhabit rocky shores but also raft across oceans, while Euchirograpsus and Miersiograpsus are found in deep-sea environments. The phylogenetic relationships of the genera, and their relationship to the previously included genus Percnon and to other grapsoids, were investigated by Schubart and Plagusiidae Dana, 1851 Cuesta (2010) using adult and larval morphology and molecular data. Molecular phylogenetics places plaguisiids nearest Figure 14.176, Plate 68h–j to gecarcinids (Schubart and Cuesta 2010; Tsang et al. 2014). Plagusiids are differentiated from other grapsoids by the Diagnosis. Carapace distinctly subcircular to square, usucombination of partial pleonite fusion, the rounded-­ ally about as long as wide; front with or without deep cleft on quadrate carapace and distally unexpanded cheliped careach side of midline to receive antennules; suborbital crest pus. The fused male pleonites 3–6 lock on to a button on smooth, without granules. Maxilliped 3 merus as wide as or the thoracic sternite. A distinctive feature of some genera is slightly wider than ischium; very small rhomboidal gap the deeply incised carapace front permitting free vertical between closed maxillipeds 3. Mandibles not visible when movement of the antennules. Plagusiids occupy a range of mouthparts closed. Chelipeds small, similar. Male pleonites habitats and depths. Species of Guinusia are large active 3–6 fused. crabs on exposed rocky shores in the Southern Hemisphere.

Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 279–299. doi:10.1163/ej.9789004170865.i-366.171 Sciberras M, Schembri PJ (2008) Biology and interspecific interactions of the alien crab Percnon gibbesi in the Maltese Islands. Marine Biology Research 4, 321–332. doi:10.1080/17451000801964923

Key to genera of Plagusiidae 1. – 2.

Frontal sinuses deeply incised (Fig. 14.176c, e)����������������������������������������������������������������������������������������������������������������������������2 Frontal sinuses weakly incised, oblique (Fig. 14.176a, b, d)�������������������������������������������������������������������������������������������������������3 Carapace tuberculate (Fig. 14.176e). Pereopods 2–5 meri extensor margin with 1 subdistal spine (Fig. 14.176j)��������������������������������������������������������������������������������������������������������������������������������������������������������������������Plagusia

Fig. 14.176.  Plagusiidae. Carapace: a, Davusia glabra (Dana, 1852); b, Euchirograpsus; c, Guinusia chabrus (Linnaeus, 1758); d, Miersiograpsus australiensis (Türkay, 1978); e, Plagusia depressa (Fabricius, 1775). Front, antennule, antenna, eyestalk: f, Davusia; g, Plagusia. Chela: h, Davusia; i, Guinusia. j, pereopod 5, Plagusia. Maxilliped 3: k, Davusia; l, Plagusia. Male gonopod 1: m, Euchirograpsus; n, Miersiograpsus.

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Marine Decapod Crustacea

Carapace not tuberculate, tomentose. Pereopods 2–5 meri extensor margin with row or teeth (Fig. 14.176c)������������������������������������������������������������������������������������������������������������������������������������������������������������������ Guinusia Cheliped palm swollen (Fig. 14.176h). Carapace smooth, glabrous����������������������������������������������������������������������������� Davusia Cheliped palm cylindrical (Fig. 14.176d). Carapace tomentose or with scattered long setae�����������������������������������������������4 Carapace anterolateral margin with more than 1 tooth plus exorbital tooth (Fig. 14.176b). Carapace tomentose. Male gonopod 1 without narrow terminal tube (Fig. 14.176m)����������������������������������������������Euchirograpsus Carapace anterolateral margin with 1 tooth plus exorbital tooth (Fig. 14.176d). Carapace with scattered long setae. Male gonopod 1 with narrow terminal tube (Fig. 14.176n)����������������������������������������������������� Miersiograpsus Davusia Guinot, 2007

sourie, smooth bait crab Diagnosis. Carapace, limbs smooth; anterolateral margin with 3 teeth plus exorbital tooth; frontal sinuses weakly incised, oblique. Antennules extending into notch reaching surface of carapace. Cheliped with swollen palm, fingers acute. Pereopods 2–5 meri with 1 felted longitudinal groove, extensor margin with 1 subdistal tooth. Maximum cl. 50 mm. Intertidal, subtidal (0–5 m; exposed rocky shores). Temperate Australia. 1 species (Guinot 2007).

Euchirograpsus H. Milne Edwards, 1853 Diagnosis. Carapace, limbs tomentose; anterolateral margin with more than 1 tooth plus exorbital tooth; frontal sinuses weakly incised, oblique. Antennules transverse, in groove under front. Cheliped with cylindrical palm, fingers acute. Pereopods 2–5 meri extensor margin with 1 subdistal tooth. Male gonopod 1 with long-terminal tube. Maximum cl. 38 mm (Pl. 69h). Shelf, slope (30–508 m). Temperate North and Tropical Atlantic, Indo-West Pacific, Tropical Eastern Pacific. 10 species (Crosnier 2001; Ng and Martin 2010: rediagnosis, species listed, distributions; Türkay 1978: rediagnosis).

incised, oblique. Antennules transverse, in groove under front. Cheliped with cylindrical palm, fingers acute. Pereopods 2–5 meri extensor margin with 1 subdistal tooth. Male gonopod 1 with long-terminal tube. Maximum cl. 9.5 mm. Shelf, slope (171–825 m). Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 2 species (Türkay 1978).

Plagusia Latreille, 1804 rafting crab Diagnosis. Carapace, limbs tomentose; anterolateral margin with 3 teeth plus exorbital tooth; frontal sinuses deeply incised. Antennules extending vertically into notch reaching surface of carapace. Cheliped with cylindrical, ridged palm, fingers with spoonshaped tips. Pereopods 2–5 meri with more than 1 felted longitudinal groove, extensor margin with row of teeth (Pl. 69j). Intertidal, subtidal, rafting (0–5 m). Temperate Northern and Tropical Atlantic, Temperate Northern Pacific, Indo-West Pacific, Tropical Eastern Pacific. 5 species (Dawson 1987: key to species; Schubart and Cuesta 2010: diagnosis, phylogeny). Species of Plagusia are known to raft across oceans on algae and other flotsam (Schubart et al. 2001). References

Guinusia Schubart & Cuesta, 2010 red rock crab, red bait crab Diagnosis. Carapace, limbs tuberculate, tomentose; anterolateral margin with 3 teeth plus exorbital tooth; frontal sinuses deeply incised. Antennules extending vertically into notch reaching surface of carapace. Cheliped with cylindrical, ridged palm, fingers with spoon-shaped tips. Pereopods 2–5 meri with more than 1 felted longitudinal groove, extensor margin with subdistal tooth. Maximum cl. 73 mm (Pl. 69i). Intertidal, subtidal (0–8 m). Central and Eastern Indo-Pacific, Tropical Eastern Pacific, Temperate South America, Temperate Southern Africa, Temperate Australasia. 2 species (Dawson 1987: key to species as Plagusia; Schubart and Cuesta 2010: diagnosis, phylogeny).

Miersiograpsus Türkay, 1978 Diagnosis. Carapace, limbs with scattered long setae; anterolateral margin with 1 tooth plus exorbital tooth; frontal sinuses weakly

Crosnier A (2001) Grapsidae (Crustacea, Decapoda, Brachyura) d’eau profonde du Pacifique sud-ouest. Zoosystema 23, 783–796. Dawson EW (1987) A key to the world species of Plagusia (Crustacea: Brachyura), with a new record of P. depressa tuberculata Lamarck from New Zealand. National Museum of New Zealand Records 3, 37–45. Guinot D (2007) A new genus of the family Plagusiidae Dana, 1851, close to Plagusia Latreille, 1804 (Crustacea, Decapoda, Brachyura). Zootaxa 1498, 27–33. doi:10.11646/zootaxa.1498.1.2 Ng NK, Martin JW (2010) Two new species of grapsoid crabs of the genus Euchirograpsus (Decapoda, Brachyura, Grapsoidea, Plagusiidae) from the Philippines and the Hawaiian Islands. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 229–250. doi:10.1163/ ej.9789004170865.i-366.143 Schubart CD, Cuesta JA (2010) Phylogenetic relationships of the Plagusiidae Dana, 1851 (Brachyura), with description of a new genus and recognition of Percnidae Števčić, 2005, as an independent family. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 279–299. doi:10.1163/ej.9789004170865.i-366.171 Schubart CD, González-Gordillo JI, Reyns NB, Liu HC, Cuesta JA (2001) Are Atlantic and Indo-Pacific populations of the rafting crab, Plagusia

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depressa (Fabricius), distinct? New evidence from larval morphology and mtDNA. Raffles Bulletin of Zoology 49, 301–310. Tsang LM, Schubart CD, Ahyong ST, Lai JCY, Au EYC, et al. (2014) Evolutionary history of true crabs (Crustacea: Decapoda: Brachyura) and the origin of freshwater crabs. Molecular Biology and Evolution 31, 1173–1187. doi:10.1093/molbev/msu068 Türkay M (1978) Zwei neue Grapsiden-Arten aus dem Indopazifik (Euchirograpsus madagariensis n. sp.) mit Einführung von Miersiograpsus n. gen. (Crustacea: Decapoda: Grapsidae). Senckenbergiana Biologica 59, 133–141.

Sesarmidae Dana, 1851 mangrove crabs Figures 14.177–14.181, Plate 70 Sesarmid mangrove crabs have a conspicuous setose ridge on the outer surface of the ischium and merus of maxilliped 3, the maxillipeds themselves being widely separate. Frequently, the pterygostomial region is covered with a reticulate pattern of ridges. Most sesarmids have the characteristic square or trapezoid carapace with the eyes set at the edge of the carapace, and long compressed walking legs. With almost 50 genera and over 250 species, this diverse family of crabs is a significant component of tropical and subtropical mangrove forests (Davie 1994; Fratini et  al. 2005; Ragionieri et al. 2009). Most burrow in muddy intertidal sediments or live under rocks and logs where they feed on terrestrial detritus (Kawaida et  al. 2019; Miranda et  al. 2017). Many have adapted to spending considerable time out of water, spending much of their time climbing in the mangrove trees themselves (Fratini et al. 2005; Shanij et al. 2016). This facility is taken to the extreme in representatives of several genera that are essentially terrestrial or associated with fresh water in streams up to 1500 above sea level or water-filled tree holes and bromeliad axils in rainforests (Dole-Olivier et al. 2000). Many species of Sesarmidae have ridges or pectinate crests on the chelipeds, especially in males, that interact with the granulate surface at the front of the carapace to

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generate sound during fighting (Goh et  al. 2019) or other social interactions (Boon et al. 2009). The Indo-West Pacific is the centre of sesarmid diversity. The tropical Pacific and Atlantic coasts of the Americas include only five genera (Abele 1992; Shahdadi et  al. 2017) among which are trans-isthmus species pairs (Schubart 2011). Only one genus occurs in both the Americas and in the IndoWest Pacific. West Africa includes few genera from both centres plus endemic genera Guinearma and Metagrapsus (Ngo-Massou et al. 2018; Shahdadi and Schubart 2018). The taxonomy of the family was explored by Serène and Soh (1970) but not all their genera and subgenera are now accepted and some of their generic names were incorrect (Davie 2010). Several genera have been added since. The monophyly of Sesarmidae was confirmed using molecular data (Schubart et al. 2006) and by studies of larvae (Cuesta et  al. 2006b) but how they relate to other thoracotremes is the subject of ongoing study. Distinctions between many genera are subtle, and some, such as Geosesarma, ­Pseudosesarma and Sesarmops require further refinement in order better define their differences (Davie and Ng 2013; Schubart and Ng 2020). Diagnosis. Carapace subquadrate–trapezoid, smooth or rugose; front approximately transverse, with or without incisions; orbit with lateral opening; suborbital crest smooth, without granules; pterygostomial region covered in reticulated network of short, hooked, setae, without groove subparallel to lateral margin of buccal cavity. Maxilliped 3 merus and ischium with distinct oblique setose ridge; distinct rhomboidal gap between closed maxillipeds 3. Mandibles usually visible when mouthparts closed. Chelipeds similar. Male pleonites 1–6 and telson free. Implicit generic attributes. Carapace with epibranchial tooth/teeth. Cornea as wide or wider than peduncle. Antennular basal article slightly or not swollen, at least twice as wide as long. Antenna peduncle included in orbit. Ambulatory legs not especially long, narrow. Male gonopod with distal chitinous apical process.

Key to genera of Sesarmidae Metopaulias Rathbun, 1897, comprising one terrestrial species living in the leaf axils of bromeliads in Jamaica (Rathbun 1897, 1918), is not included. Orisarma, Pseudosesarma, Selatium and Sesarma appear twice because of variability in the number of epibranchial carapace teeth. Genera including species from fresh water or terrestrial environments marked with *

1. – 2. – 3.

Antenna peduncle excluded from orbit (Fig. 14.179t)�����������������������������������������������������������������������������������������������������������������2 Antenna peduncle included in orbit (Fig. 14.179s, v, w)�������������������������������������������������������������������������������������������������������������4 Carapace lateral margins strongly converging on convex posterior margin, appearing triangular. Male pleon subcircular (Fig. 14.181c)���������������������������������������������������������������������������������������������������������������������������������������Aratus Carapace quadrate. Male pleon triangular (Fig. 14.181h)����������������������������������������������������������������������������������������������������������3 Carapace with anterolateral teeth, dorsum uneven, with mat of short setae (Fig. 14.179b)�������������������������� Clistocoeloma

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– 4. – 5. – 6. – 7.

Marine Decapod Crustacea

Carapace without anterolateral tooth, dorsum smooth��������������������������������������������������������������������������������������� Metasesarma Pereopods 2–5 merus flexor distal margin with denticulate arc and larger spine (Fig. 14.181s). Maximum c. 8 mm�����������������������������������������������������������������������������������������������������������������������������������������������Nanosesarma Pereopods 2–5 with flexor distal margin of meri smooth or granulate, at most with minute spinule (Fig. 14.181o–r)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������5 Carapace without epibranchial tooth behind exorbital angle (e.g. Fig. 14.179c, g, h)�����������������������������������������������������������6 Carapace with 1 or 2 epibranchial teeth behind exorbital angle (e.g. Fig. 14.179k, m), rarely absent�������������������������������21 Carapace smooth, subcircular, anterolateral margin curving towards exorbital angle (Fig. 14.179c). Pereopods 2–5 meri–propodi expanded, wide (Fig. 14.181o)��������������������������������������������������������������������������������Cyclorma Carapace quadrate or trapezoidal (e.g. Fig. 14.179g, h). Pereopods 2–5 meri–propodi linear���������������������������������������������7 Carapace widest posteriorly, trapezoidal, margins diverging posteriorly (Fig. 14.179h)��������������������������������Lithoselatium

Fig. 14.177.  Sesarmidae. a, Armases cinereum Bosc, 1802; b, Circulium rotundatum (Hess, 1865); c, Episesarma singaporensis Tweedie, 1936; d, Guinearma huzardi (Desmarest, 1825); e, Karstarma guamense (Ng, 2002); f, Labuanium politum (De Man, 1887) (with front, ventral view).

14 – Brachyura – crabs

811

Fig. 14.178.  Sesarmidae. a, Metagrapsus curvatus (H. Milne Edwards, 1837); b, Miersarma granosimanum (Miers, 1880); c, Mindanium cruciatum (Bürger, 1893); d, Namlacium crepidatum (Calman, 1925); e, Sesarma crassipes Cano, 1889; f, Sesarmoides borneensis (Tweedie, 1950); g, Tiomanium indicum (H. Milne Edwards, 1837).

– 8. – 9.

Carapace quadrate, margins subparallel or trapezoidal, margins converging posteriorly���������������������������������������������������8 Pereopods 2, 3 subchelate, propodus expanded on flexor margin (Fig. 14.181q). Carapace front concave (Fig. 14.179g)�����������������������������������������������������������������������������������������������������������������������������������������������������������������Haberma Pereopods 2, 3 simple. Carapace front straight or medially concave���������������������������������������������������������������������������������������9 Cheliped palm upper surface with longitudinal granular or pectinated ridge (Figs 14.178e, 14.180t)�����������������������������10

812

Marine Decapod Crustacea

– 10. – 11. –

Cheliped palm upper surface without longitudinal granular ridge (Fig. 14.177a, 14.180a, c)������������������������������������������� 12 Carapace front medially straight (Fig. 14.179d)��������������������������������������������������������������������������������������������������������Cristarma* Carapace front medially concave�������������������������������������������������������������������������������������������������������������������������������������������������11 Cheliped dactylus upper margin smooth or irregularly granular (Fig. 14.180u)������������������������������������ Sesarma (in part)* Cheliped dactylus upper margin with pectinate ridge (Fig. 14.180t)����������������������������������������������������������������������������Selatium (in part, S. elongatum (A. Milne-Edwards, 1869)) 12. Male cheliped merus inner margin lamelliform (Fig. 14.180y)�����������������������������������������������������������������������������������������������13 – Male cheliped merus inner margin not expanded (Fig. 14.180b, j)����������������������������������������������������������������������������������������14 13. Carapace almost square, lateral margins parallel or almost so (Fig. 14.177a). Upper margin of male cheliped dactylus without stridulatory ridge (Fig. 14.180a)����������������������������������������������������������������������������������Armases* – Carapace distinctly trapezoidal, lateral margins posteriorly convergent. Upper margin of male cheliped dactylus with stridulatory ridge (Fig. 14.180y)���������������������������������������������������������������������������������������������������������Stelgistra 14. Carapace front sinuous, with rounded median concavity�������������������������������������������������������������������������������������������������������15 – Carapace front almost straight, weakly sinuous�����������������������������������������������������������������������������������������������������������������������17

Fig. 14.179.  Sesarmidae. Carapace: a, Andamanium; b, Clistocoeloma; c, Cyclorma; d, Cristarma; e, Eneosesarma; f, Geosesarma; g, Haberma; h, Lithoselatium; i, Nanosesarma; j, Neosesarma; k, Parasesarma; l, Pseudosesarma; m, Sarmatium; n, Scandarma; o, Selatium; p, Sesarmops; q, Shinobium. r, front of carapace, Parasesarma. Orbit, antennule, antenna: s, Armases; t, Clistocoeloma; u, Danarma; v, Geosesarma; w, Sesarmoides (with inner face of cheliped merus).

14 – Brachyura – crabs

15. – 16. – 17. – 18. – 19. – 20. – 21. – 22. – 23. – 24. – 25. – 26. – 27. – 28. –

813

Cheliped palm outer surface smooth. Male pleon lateral margins gently concave; pleonite 6 half as wide as pleonite 3 (Fig. 14.181e)������������������������������������������������������������������������������������������������������������������������� Chiromantes* Cheliped palm outer surface tuberculate. Male pleon lateral margins straight to gently convex; pleonite 6 at least 0.6 times as wide as pleonite 3 (Fig. 14.181j)������������������������������������������������������������������������������������������16 Male pleonite 6 0.6 times as wide as pleonite 3 (Fig. 14.181j)������������������������������������������������������������������������Orisarma (part)* Male pleonite 6 0.7 times as wide as pleonite 3 (Fig. 14.181l)���������� Pseudosesarma (part, P. boulengeri (Calman, 1920))* Cheliped dactylus upper margin with proximal row of minute corneous granules preceding row of widely spaced tubercles on central portion of margin (Fig. 14.180w, x)����������������������������������������������������Sinosesarma Cheliped dactylus upper margin smooth evenly or irregularly granulate����������������������������������������������������������������������������18 Carapace ovate-trapezoidal; front downcurved, obscuring antennules and antennae from anterior view (Fig. 14.179u)������������������������������������������������������������������������������������������������������������������������������������������������������Danarma* Carapace trapezoidal or quadrate; front horizonal, antennules and antennae visible in anterior view (Fig. 14.179v, w)����������������������������������������������������������������������������������������������������������������������������������������������������������������19 Carapace square, lateral margins parallel; postfronal and epigastric lobes not forming near-continuous transverse crest. Cheliped palm outer surface strongly convex (Fig. 14.178b)�����������������Miersarma* Carapace trapezoidal, lateral margins convergent posteriorly; postfrontal and epigastric lobes merged, forming near-continuous transverse crest. Cheliped palm outer surface flat (Fig. 14.180p)��������������������������������������� 20 Male telson slightly wider than long, margins divergent (Fig. 14.181k). Gonopod 1 chitinous tip elongated, more than twice as long as wide (Fig. 14.181y)���������������������������������������������������������������������������Platychirarma* Male telson slightly longer than wide, margins parallel (Fig. 14.181m). Gonopod 1 chitinous tip short, about as long wide (Fig. 14.181z)���������������������������������������������������������������������������������������������������������Trapezarma* Cheliped palm upper surface with 4–7 wide transverse grooves separating wide ridges, some pectinate; dactylus in male with 1 or more large wide chitinous tubercles (Fig. 14.180q). Ocular peduncle swollen basally, cornea reduced (Fig. 14.179m)����������������������������������������������������������������������������������������������������Sarmatium Cheliped palm upper surface without transverse grooves, or flat, or with longitudinal-oblique granular or pectinate ridges. Ocular peduncle not swollen, cornea wider than peduncle������������������������������������������� 22 Male cheliped palm upper surface with oblique or curved pectinate ridges (Fig. 14.180d–f, h, l–o)������������������������������� 23 Male cheliped palm upper surface smooth to irregularly granular, OR with simple longitudinal pectinate ridge�������������������������������������������������������������������������������������������������������������������������������������������������������������������������� 27 Male cheliped palm upper surface with 1 oblique or curved pectinate crest (Fig. 14.180e, f, h)�������������������������������������� 24 Male cheliped palm upper surface with 2 oblique or curved pectinate crests (Fig. 14.180d, l–o)������������������������������������ 25 Carapace with lateral margins convex, widest at midlength, strongly convergent posteriorly (Fig. 14.178a)����������������������������������������������������������������������������������������������������������������������������������������������������������� Metagrapsus Carapace with lateral margins straight or convex, but widest across anterior (Fig. 14.177d)��������������������������� Guinearma Male cheliped palm upper surface with 2 oblique or nearly longitudinal ridges of chitinous tubercles; dactylus with 2 low tubercles (Fig. 14.180d)����������������������������������������������������������������������������������������������������������� Fasciarma Male cheliped palm upper surface with usually 2 transverse pectinate crests; dactylus tuberculate������������������������������ 26 Male cheliped palm upper surface with 1–3 (usually 2) transverse pectinate crests of tall chitinous teeth flanked by large tubercle on inside (Fig. 14.180l, m)������������������������������������������������������������������������������ Parasesarma Male cheliped palm upper surface with 2 transverse pectinate crests of tall chitinous teeth flanked by 2 small tubercles on inside (Fig. 14.180n, o)��������������������������������������������������������������������������������������������������� Perisesarma Male cheliped palm upper surface with simple longitudinal pectinate or granular ridge along part or all of surface, or parallel to inner margin (Fig. 14.180c, k, r, s, u)������������������������������������������������������������������������������������ 28 Male cheliped palm upper surface smooth or irregularly granular, or with unornamented ridge�����������������������������������33 Cheliped palm upper surface with continuous pectinate or granular ridge; dactylus with similar ridge (Fig. 14.180c, q, r)��������������������������������������������������������������������������������������������������������������������������������������������������������������������� 29 Cheliped palm upper margin with 1 longitudinal ridge of even close-set granules or short pectinate ridge; dactylus upper margin with irregular row of larger tubercles or granules (Fig. 14.180k, s, t, u)��������������������������������� 30

814

Marine Decapod Crustacea

29.

Carapace with 1 prominent epibranchial tooth (Fig. 14.177c). Cheliped palm outer surface tuberculate proximally transitioning to smooth distally, without median protuberance (Fig. 14.180c)������������������������ Episesarma – Carapace epibranchial teeth weakly indicated or absent (Fig. 14.179n). Cheliped palm outer surface distinctly and uniformly tuberculate except for sharply demarcated smooth surface and median protuberance, near base of fingers (Fig. 14.180r, s)��������������������������������������������������������������������������������������������Scandarma* 30. Pereopods 2–5 dactyli two-thirds as long as propodi or longer (Fig. 14.178e)���������������������������������������������������������������������31 – Pereopods 2–5 dactyli one-third–half propodi length�������������������������������������������������������������������������������������������������������������32 31. Cheliped palm upper surface with pectinate crest limited to ‘limited to median part, proximally and distally continued by a smooth rim’ (not illustrated)�������������������������������������������������������������������������������������������Muradium – Cheliped palm upper surface with full-length granular crest (Fig. 14.180u)������������������������������������������� Sesarma (in part)* 32. Carapace front width much more than half exorbital width, without median concavity (Fig. 14.179j). Cheliped palm with fully pectinate crest clearly separate from upper inner margin (Fig. 14.180k). Pereopods 2–5 dactyli about half length of propodi����������������������������������������������������������������������������������������� Neosesarma – Carapace front width little more than half exorbital width, with median concavity (Fig. 14.179o). Cheliped palm upper inner surface with longitudinal partly-pectinate crest (Fig. 14.180t). Pereopods 2–5 dactyli one-third length of propodi�������������������������������������������������������������������������� Selatium (in part, S. brockii (De Man, 1887))

Fig. 14.180.  Sesarmidae. Male cheliped. a, Armases; b, Chiromantes; c, Episesarma; d, Fasciarma; e, f, Guinearma; g, Labuanium; h, Metagrapsus; i, j, Neosarmatium; k, Neosesarma; l, m, Parasesarma; n, o, Perisesarma; p, Platychirarma; q, Sarmatium; r, s, Scandarma; t, Selatium; u, Sesarma; v, Shinobium; w, x, Sinosesarma; y, Stelgistra; z, Tiomanium. a, q, r, v, chela outer views; d, e, h, l, n, t, palm upper surface; f, m, o, outer view of crest on palm; i, w, chela inner view; b, c, g, j, k, s, carpus, chela, upper view; b, z, merus–chela upper view; x, dactylus upper margin

14 – Brachyura – crabs

815

33. Carapace front wider than half exorbital width (e.g. Figs 14.177f, 14.178g, 14.179p, q)����������������������������������������������������� 34 – Carapace front half or less exorbital width������������������������������������������������������������������������������������������������������������������������������� 44 34. Carapace wider longer than wide; front deflexed more than 90°, postfrontal lobes produced forwards as 4 quadrate lobes, overhanging true frontal margin (Figs 14.177f, 14.179q)������������������������������������������������������������������������35 – Carapace wider than long; front horizontal or slightly deflexed, directed more-or-less anteriorly, margin concave medially in dorsal view�������������������������������������������������������������������������������������������������������������������������������������������� 36 35. Carapace almost parallel-sided (Fig. 14.177f). Male cheliped palm upper surface with incomplete lines of granules (Fig. 14.180g)������������������������������������������������������������������������������������������������������������������������������� Labuanium – Carapace widest posteriorly (Fig. 14.179q). Male cheliped palm upper surface with oblique lines of granules (Fig. 14.180v)���������������������������������������������������������������������������������������������������������������������������������������������Shinobium* 36. Cheliped palm upper surface sharp, ending in distal tooth; inner surface with denticles; carpus with long, acute inner angle (Figs 14.178g, 14.180z)����������������������������������������������������������������������������������������������������Tiomanium – Cheliped palm upper surface flat, without distal tooth; inner surface with or without vertical row of granules; carpus without long, acute inner angle����������������������������������������������������������������������������������������������������������������37 37. Cheliped palm inner surface with vertical row of prominent granules near base of fingers, reaching upper margin (Fig. 14.180i, j); upper and outer surfaces smooth, punctate, not tuberculate����������������� Neosarmatium

Fig. 14.181.  Sesarmidae. Maxilliped 3: a, Geosesarma; b, Selatium. Male pleon: c, Aratus; d, Bresedium; e, Chiromantes; f, Contusarma; g, Geosesarma; h, Metasesarma; i, Migmarma; j, Orisarma; k, Platychirarma; l, Pseudosesarma; m, Trapezarma. n, male thoracic sternum, gonopods, Pseudosesarma. Pereopod: o, Cyclorma; p, Geosesarma; q, Haberma; r, Labuanium; s, Nanosesarma. Pereopod 5: t, Orisarma; u, Manarma. Gonopod 1: v, Bresedium; w, Labuanium; x, Neosarmatium; y, Platychirarma; z, Trapezarma.

816

– 38. – 39. – 40. – 41. – 42. – 43. – 44. – 45. – 46. – 47. – 48. – 49. – 50. –

Marine Decapod Crustacea

Cheliped palm inner surface smooth or granulate, with or without vertical granular row at midlength, upper and usually outer surfaces tuberculate������������������������������������������������������������������������������������������������� 38 Carapace lateral margins divergent posteriorly, widest across posterior (Fig. 14.179p)��������������������������������������� Sesarmops Carapace lateral margins divergent anteriorly, widest across anterior��������������������������������������������������������������������������������� 39 Male telson deeply embedded into pleonite 6 (Fig. 14.181d). Male gonopod with elongate distal process (Fig. 14.181v)������������������������������������������������������������������������������������������������������������������������������������������������Bresedium* Male telson not embedded into pleonite 6, at most shallowly recessed. Male gonopod usually with short ‘stubby’ distal chitinous process, seldom elongated����������������������������������������������������������������������������������������������������������� 40 Pereopod 5 dactylus as long as or shorter than upper margin propodus (Fig. 14.181u)�������������������������������������� Manarma* Pereopod 5 dactylus longer than upper margin of propodus (Fig. 14.181t)��������������������������������������������������������������������������41 Male pleonite 6 0.6 times width of pleonite 3 (Fig. 14.181j)��������������������������������������������������������������������������������������Orisarma* Male pleonite 6 0.67 times or more width of pleonite 3 (Fig. 14.181f, i, l)���������������������������������������������������������������������������� 42 Male sternite 5 with excavation on each wall of sternopleonal cavity with excavation for reception of swollen gonopods 1 (Fig. 14.181n)������������������������������������������������������������������������������������������� Pseudosesarma* Male sternite 5 without excavation on each wall of sternopleonal cavity���������������������������������������������������������������������������� 43 Male telson as long as wide or slightly longer than wide (Fig. 14.181f)������������������������������������������������������������� Contusarma* Male telson slightly wider than long (Fig. 14.181i)�������������������������������������������������������������������������������������������������� Migmarma* Carapace longer than exorbital width (Fig. 14.177b)��������������������������������������������������������������������������������������������������Circulium Carapace as long as or shorter than exorbital width���������������������������������������������������������������������������������������������������������������� 45 Carapace almost circular, with strongly convex lateral margins (Fig. 14.179e)�������������������������������������������������Eneosesarma Carapace more or less rectangular, with straight or weakly convex anterolateral margins���������������������������������������������� 46 Carapace lateral margins strongly, widest near midlength; with 1 pair of postfrontal lobes, far from frontal margin. Pereopods 2–5 dactyli one-third length of propodi, densely setose (Fig. 14.178d)������������� Namlacium Carapace lateral margins straight or slightly curved. Pereopods 2–5 dactyli half or more as long as propodi���������������� 47 Pereopods 5 dactylus shorter than upper margin of propodus (Fig. 14.178c)�����������������������������������������������������Mindanium Pereopods 5 dactylus as long as or longer than upper margin of propodus������������������������������������������������������������������������ 48 Carapace square or trapezoidal; exorbital tooth prominent, flared anterolaterally (Fig. 14.179a, f). Male chelipeds equal��������������������������������������������������������������������������������������������������������������������������������������������������������������� 49 Carapace subtrapezoidal, widest posteriorly; exorbital tooth weakly developed, incurved (Fig. 14.177e). Male chelipeds unequal����������������������������������������������������������������������������������������������������������������������������������������������������������� 50 Carapace lateral margins subparallel; postfrontal lobes overhanging true frontal margin, appearing as 4 broad transverse lobes separated by fissures (Fig. 14.179a). Male chelipeds palm outer surface smooth, without tubercles�������������������������������������������������������������������������������������������������������������������������������� Andamanium* Carapace lateral margins subparallel, divergent or convergent; postfrontal lobes not overhanging true frontal margin; frontal margin sinuous in dorsal view with median concavity (Fig. 14.179f). Male chelipeds palm outer surface tuberculate�������������������������������������������������������������������������������������������������� Geosesarma Carapace trapezoidal, anterolateral margins divergent posteriorly (Fig. 14.178f). Cheliped fixed finger with longitudinal row or granules or ridge on outer surface; merus inner surface with short ridge forming stridulating organ with milled suborbital margin (Fig. 14.179w)���������������������������������������������������� Sesarmoides Carapace usually subquadrate (Fig. 14.177e). Cheliped fixed finger without ornamentation on outer surface; without stridulating ridge����������������������������������������������������������������������������Karstarma* Andamanium Naruse & Ng, 2020

Diagnosis. Carapace wider than long, quadrate, lateral margins subparallel; with 1 distinct, 2 weak epibranchial teeth; front about half exorbital width, front deflexed, distally recurved, directed anteroventrally; margin straight in anterior view; postfrontal lobes well developed, overhanging true front, appearing in dorsal

view as 4 broad, transverse lobes separated by fissures. Male chelipeds symmetrical; cheliped palm upper surface with oblique lines of granules, outer surface without prominent sloping smooth area; dactylus with row tubercles. Pereopods 5 dactylus longer than upper margin of propodus. Maximum cl. 17 mm. Freshwater, 300–400 m asl. Andaman Is. 1 species (Naruse and Ng 2020).

14 – Brachyura – crabs

Aratus H. Milne Edwards, 1853 Diagnosis. Carapace lateral margins strongly convergent on rounded posterior margin, appearing triangular; without epibranchial tooth. Antenna peduncle excluded from orbit. Male cheliped palm almost smooth. Pereopods 2–5 dactyli about onefifth length of propodi. Male pleon subcircular. Maximum cl. 25 mm (Pl. 70a). Estuarine, supralittoral (tree-climbing). Tropical Atlantic, Eastern Indo-Pacific. 2 species (Cuesta et al. 2006a: larvae, biology; Thiercelin and Schubart 2014: 2 species).

Armases Abele, 1992 Diagnosis. Carapace wider than long, dorsally almost flat; with weak epibranchial tooth; front with weak median concavity. Male cheliped palm upper surface with scattered granules. Maximum cl. 27 mm (Pl. 70b). Estuarine, intertidal (including mangroves, freshwater, terrestrial to 330 m asl, some in bromeliads). Temperate Northern Atlantic, Tropical Atlantic, Tropical Eastern Pacific. 13 species (Abele 1992: diagnosis, keys to species; Niem 1996: phylogeny).

Bresedium Serène & Soh, 1970 Diagnosis. Carapace little shorter than exorbital width, quadrate, dorsally almost flat; with prominent epibranchial tooth; front about half exorbital width, front with strong median concavity. Male cheliped palm upper margin without pectinate ridge; upper and outer surfaces tuberculate or variously granulate, inner surface with tubercles. Male telson deeply embedded into pleonite 6. Male gonopod with elongate distal process. Maximum cl. 17 mm. Freshwater, semiterrestrial. SE Asia, N Australia. 3 species (Li 2014: redescription, figure; Li et al. 2020; Schubart and Ng 2020: species comparisons).

817

sharp, outer surface without prominent sloping smooth are; dactylus with row tubercles. Pereopods 2–5 dactyli shorter than half length of propodi. Maximum cl. 44 mm. Supralittoral, tree-climbing. Western and Central IndoPacific. 4 species (Naruse and Ng 2020; Ng and Davie 2011).

Clistocoeloma A. Milne-Edwards, 1873 Diagnosis. Carapace quadrate, dorsally uneven, with mat of setae; with weak epibranchial tooth. Antenna peduncle excluded from orbit. Male cheliped palm upper surface with pectinate crest continuous, of numerous tubercles; dactylus with similar crest. Pereopods 2–5 dactyli third length of propodi. Maximum cl. 23 mm (Pl. 70c). Intertidal, supralittoral. Central Indo-Pacific. 9 species (Lee et al. 2013: list of species).

Contusarma Schubart & Ng, 2020 Diagnosis. Carapace quadrate, lateral margins parallel; front wider than half exorbital width; with epibranchial tooth. Male chelipeds asymmetrical; cheliped palm upper surface without pectinate ridge; upper and outer surfaces tuberculate. Male sternite 5 without excavation on each wall of sternopleonal cavity. Male pleon lateral margins straight to gently convex; pleonite 6 two-thirds width of pleonite 3; telson as long as wide or slightly longer than wide, not embedded into pleonite 6. Pereopod 5 dactylus longer than upper margin of propodus Maximum cl. 24 mm. Intertidal, supralittoral, estuarine, coastal freshwater swamps. Central Indo-West Pacific. 2 species (Schubart and Ng 2020).

Cristarma Schubart & Ng, 2020

Diagnosis. Carapace lateral margins convergent posteriorly; without epibranchial tooth. Male chelipeds asymmetrical; cheliped palm upper margin without longitudinal pectinate ridge; upper and outer surfaces smooth; inner surface without vertical row of granules. Male pleon lateral margins gently concave; pleonite 6 half as wide as pleonite 3. Maximum cl. 30 mm. Intertidal, supralittoral. Central Indo-West Pacific. 2 species (Schubart and Ng 2020).

Diagnosis. Carapace quadrate, lateral margins subparallel; front wider than half exorbital width, gently deflexed, not obscuring antennae from anterior view, straight in dorsal view; without epibranchial tooth. Male chelipeds asymmetrical; cheliped palm upper surface with pectinated ridge; dactylus with upper stridulatory ridge. Male pleon lateral margins gently concave; pleonite 6 0.5 width of pleonite 3; telson slightly longer than wide, not embedded into pleonite 6. Pereopod 5 dactylus as long as or longer than upper margin of propodus Maximum cl. 19 mm. Supralittoral. Western Indo-West Pacific. 2 species (Schubart and Ng 2020).

Circulium Naruse & Ng, 2020

Cyclorma Naruse & Ng, 2012

Diagnosis. Carapace longer than exorbital width, circular, laterally inflated on branchial regions; with 1 or 2 epibranchial teeth; front about half exorbital width, front deflexed, distally recurved, directed anteriorly; margin concave medially in dorsal view; lateral pair of postfrontal lobes exceeding mesial pair; neither reaching frontal margin in dorsal view. Antennular basal article swollen, little wider than long. Male cheliped palm upper surface

Diagnosis. Carapace smooth, subcircular, anterolateral margin curving towards exorbital angle; without epibranchial tooth. Male cheliped palm upper surface with line of small granules; dactylus with c. 8 granules. Pereopods 2–5 with meri-propodi flat, wide. Maximum cl. 15 mm. Intertidal. Western Indo-Pacific (Djibouti). 1 species (Naruse and Ng 2012).

Chiromantes Gistel, 1848

818

Marine Decapod Crustacea

Danarma Schubart & Ng, 2020 Diagnosis. Carapace ovate-trapezoidal, lateral margins convergent posteriorly; front wider than half exorbital width, downcurved, obscuring antennae from anterior view, straight in dorsal view; without epibranchial tooth. Male chelipeds asymmetrical; cheliped palm upper surface without pectinated ridge. Male sternite 5 without excavation on each wall of sternopleonal cavity. Male pleon lateral margins straight to gently convex; pleonite 6 0.6 width of pleonite 3; telson as long as wide or slightly longer than wide, not embedded into pleonite 6. Pereopod 5 dactylus as long as or longer than upper margin of propodus Maximum cl. 15 mm. Intertidal, supralittoral. Central Indo-West Pacific. 5 species (Schubart and Ng 2020).

Eneosesarma Brösing, Spiridonov, Al-Aidaroos & Türkay, 2014 Diagnosis. Carapace wider than long, subcircular, with strongly convex lateral margins, widest anterior to midpoint; without epibranchial tooth; front about half exorbital width, front nearly straight, without median concavity. Antennular basal article swollen, little wider than long. Male cheliped palm upper surface with few granules. Pereopods 2–5 dactyli about as long as propodi. Maximum cl. 12 mm. Supralittoral. Saso Island, Red Sea. 1 species (Brösing et  al. 2014).

Episesarma De Man, 1895 Diagnosis. Carapace quadrate; front with strong median concavity; postfrontal lobes well developed; with 1 epibranchial tooth behind exorbital angle. Male cheliped merus anterior border with subdistal triangular or lobiform process with denticulate margin, palm upper surface with pectinate crest continuous, of numerous tubercles; dactylus with similar crest of over 30 tubercles. Pereopods 2–5 dactyli longer than half length of propodi. Maximum cl. 50 mm (Pl. 70d). Estuarine, supralittoral. Central Indo-Pacific. 7 species (Lee et al. 2015: rediagnosis, key to 5 species).

Fasciarma Shahdadi & Schubart, 2017 Diagnosis. Carapace lateral margins parallel over anterior half, convergent posteriorly; with weak epibranchial tooth. Male cheliped palm upper surface with 2 oblique-nearly longitudinal ridges of chitinous tubercles; dactylus with 2 low tubercles. Maximum cl. 13 mm (Pl. 70e). Estuarine, intertidal. Central Indo-Pacific. 1 species (Shahdadi and Schubart 2018).

epibranchial teeth; front about half exorbital width, front sinuous, medially concavity; postfrontal lobes well developed. Antennular basal article swollen, little wider than long. Antenna peduncle included in orbit (longitudinal). Male cheliped palm upper surface with few incomplete lines of granules; dactylus with row tubercles. Pereopods 2–5 margins straight; dactyli about as long as propodi. Maximum cl. 14 mm (Pl. 70f). Supralittoral, terrestrial, forests. Central Indo-Pacific. SE Asia. 68 species. Ng and Davie (1995) listed 40 species, a figure now superseded, and opined that the genus has ‘no clear definition’. Geosesarma is essentially a terrestrial genus of very colourful crabs, some traded as pets. Ng and Wowor (2019) provided a key to 11 species from Java. Naruse and Ng (2020) remarked on species groups and redescribed two species.

Guinearma Shahdadi & Schubart, 2017 Diagnosis. Carapace lateral margins parallel over anterior half, convergent posteriorly; with 2 epibranchial teeth. Male cheliped palm upper surface with 1 straight or curved tuberculate crest, followed by proximal line of granules; dactylus with low tubercles. Maximum cl. 50 mm. Estuarine, intertidal. Tropical E Atlantic. 3 species (Shahdadi and Schubart 2018).

Haberma Ng & Schubart, 2002 Diagnosis. Carapace quadrate; without epibranchial tooth; front concave. Male cheliped palm upper surface with several short uneven ridges of small granules. Pereopods 2–5 1, 2, of adult male subchelate. Maximum cl. 8.2 mm. Intertidal (tree-climbing). Central Indo-Pacific. 3 species (Cannicci and Ng 2017: compared 3 species; Ng and Schubart 2002).

Karstarma Davie & Ng, 2007 Diagnosis. Carapace wider than long, usually more or less rectangular, with curved anterolateral margins, widest near posterior margin; exorbital tooth weakly developed, incurved; front about half exorbital width, front with weak median concavity; postfrontal lobes obsolete. Antennular basal article swollen, little wider than long. Male cheliped merus inner upper angle expanded to sublamelliform lobe; palm almost smooth. Pereopods 2–5 long, narrow; dactyli about as long as propodi. Maximum cl. 12 mm. Karst areas, caves, sometimes distant from sea. SE Asia, N Australia. 18 species (Davie and Ng 2007; Poupin et  al. 2018; Wowor and Ng 2018).

Labuanium Serène & Soh, 1970 Geosesarma De Man, 1892 vampire crabs Diagnosis. Carapace as wide as long, widest anteriorly or posteriorly; exorbital tooth prominent, flared anterolaterally; with 1 or 2

Diagnosis. Carapace longer than wide, longitudinally rectangular, with subparallel lateral margins; with 1 epibranchial tooth; front exceeding half exorbital width, front deflexed more than 90°, distally sloping ventroposteriorly; bilobed, recurved, directed anteriorly; mesial and lateral pairs of postfrontal lobes aligned; reaching

14 – Brachyura – crabs

beyond frontal margin in dorsal view, overhanging front. Antennular basal article swollen, little wider than long. Male cheliped palm upper surface sharp; dactylus with row tubercles. Pereopods 2–5 long, narrow; pereopods 2–4 dactyli shorter than half length of propodi; pereopod 5 dactylus about as long as propodus. Maximum cl. 36 mm (Pl. 70g). Estuarine, intertidal, associated with nipah palms, mangroves. Western and Central Indo-Pacific. 1 species (Naruse and Ng 2020: rediagnosis, limit of genus to 1 species; Ng et al. 2015: ecology).

Lithoselatium Schubart, Liu & Ng, 2009 Diagnosis. Carapace as wide as long, weakly trapezoidal, lateral margins weakly diverging posteriorly, without epibranchial tooth behind exorbital angle; front with weak median concavity; postfrontal lobes well developed. Male cheliped palm upper margin with 1 longitudinal crest of regular tubercles, inner surface convex without ridge; dactylus upper margin with c. 30 tubercles. Pereopods 2–5 dactyli longer than two-thirds length of propodi. Maximum cl. 27 mm. Intertidal, supralittoral. Central Indo-Pacific. 3 species (Schubart et al. 2009).

Manarma Schubart & Ng, 2020 Diagnosis. Carapace quadrate–trapezoid, lateral margins parallel or convergent posteriorly; front wider than half exorbital width; with epibranchial tooth. Male chelipeds asymmetrical; cheliped palm upper surface with short pectinate ridge; upper and outer surfaces tuberculate. Male sternite 5 without excavation on each wall of sternopleonal cavity. Male pleon lateral margins straight to gently convex; pleonite 6 6 0.6 times width of pleonite 3; telson as long as wide, not embedded into pleonite 6. Pereopod 5 dactylus as long as or shorter than upper margin of propodus Maximum cl. 18 mm (Pl. 70n). Intertidal, supralittoral, estuarine into fresh water. Central Indo-West Pacific. 2 species (Schubart and Ng 2020).

Metagrapsus H. Milne Edwards, 1853 Diagnosis. Carapace lateral margins convex, widest at midpoint, strongly convergent posteriorly, dorsally strongly convex; with prominent epibranchial tooth. Male cheliped palm upper outer surface with curved longitudinal pectinate ridge; dactylus upper margin with oblique rugose ridge. Maximum cl. 26 mm. Intertidal, supralittoral. Tropical E Atlantic. 1 species (Manning and Holthuis 1981; Shahdadi and Schubart 2018).

Metasesarma H. Milne Edwards, 1853 Diagnosis. Carapace quadrate, convex, smooth; without epibranchial tooth. Antenna peduncle excluded from orbit. Male cheliped palm upper surface with line of microscopic granules extending on to dactylus. Pereopods 2–5 dactyli longer than half length of propodi. Maximum cl. 24 mm (Pl. 70h). Supralittoral. Central Indo-Pacific. 2 species (Ng and Davie 1995: diagnosis).

819

Miersarma Schubart & Ng, 2020 Diagnosis. Carapace square, lateral margins subparallel; front wider than half exorbital width, gently deflexed, not obscuring antennae from anterior view, straight in dorsal view; postfrontal and epigastric lobes not forming near-continuous transverse crest; without epibranchial tooth. Male chelipeds asymmetrical; cheliped palm upper surface without longitudinal pectinated ridge; outer surface convex. Male sternite 5 without excavation on each wall of sternopleonal cavity. Male pleon lateral margins straight to gently convex; pleonite 6 0.7 width of pleonite 3; telson as long as wide, not embedded into pleonite 6. Pereopod 5 dactylus as long as or longer than upper margin of propodus Maximum cl. 15 mm. Intertidal, supralittoral, estuarine, freshwater swamps. Central Indo-West Pacific. 1 species (Schubart and Ng 2020).

Migmarma Schubart & Ng, 2020 Diagnosis. Carapace quadrate–trapezoid, lateral margins parallel or convergent posteriorly; front wider than half exorbital width; with epibranchial tooth. Male chelipeds asymmetrical; cheliped palm upper surface without pectinate ridge; upper and outer surfaces tuberculate. Male sternite 5 without excavation on each wall of sternopleonal cavity. Male pleon lateral margins straight to gently convex; pleonite 6 two-thirds width of pleonite 3; telson slightly wider than long, not embedded into pleonite 6. Pereopod 5 dactylus longer than upper margin of propodus Maximum cl. 14 mm. Intertidal, supralittoral, estuarine. Central Indo-West Pacific. 2 species (Schubart and Ng 2020).

Mindanium Naruse & Ng, 2020 Diagnosis. Carapace as long as exorbital width, quadrate, lateral margin slightly sinuous; with 2 epibranchial teeth; front about half exorbital width, front deflexed more than 90°, distally sloping ventroposteriorly; bilobed, recurved, directed anteriorly; mesial and lateral pairs of postfrontal lobes aligned; reaching beyond frontal margin in dorsal view, overhanging front. Pereopods 5 dactylus about two-thirds as long as propodus. Maximum cl. 17 mm. Habitat not documented. Philippines. 1 species (Naruse and Ng 2020).

Muradium Serène & Soh, 1970 Diagnosis. Carapace quadrate, pubescent, especially anteriorly; front with strong median concavity; postfrontal lobes well developed. Male cheliped palm upper surface with pectinate crest limited to median part; dactylus with rim of well spaced elongate-conical tubercles. Pereopods 2–5 dactyli about as long as propodi. Maximum cl. 36 mm. Estuarine. Western Indo-Pacific (India, Sri Lanka). 1 species (Serène and Soh 1970; not recently illustrated).

820

Marine Decapod Crustacea

Namlacium Serène & Soh, 1970 Diagnosis. Carapace wider than long, lateral margins strongly convex, widest at midlength, strongly convergent posteriorly; front about half exorbital width, front nearly straight, without median concavity; with 1 pair of postfrontal lobes, far from frontal margin. Antennular basal article swollen, little wider than long. Pereopods 2–5 long, narrow; merus with minute upper and lower distal spine; dactyli one-third length of propodi. Maximum cl. 12 mm. Habitat not documented, probably tree-climbing. Central Indo-Pacific (E Papua New Guinea). 1 species (Calman 1925: rediagnosis, redescription of species; Naruse and Ng 2020). The only species has apparently not been recorded since its description.

Nanosesarma Tweedie, 1950 Diagnosis. Carapace lateral margins convergent posteriorly; with prominent epibranchial tooth. Male cheliped palm upper surface with line of microscopic granules extending on to dactylus. Pereopods 2–5 with lower distal margin of meri with dentate arc and larger spine. Maximum cl. 8 mm (Pl. 70i). Intertidal. Western and Central Indo-Pacific. 10 species (­Naderloo and Türkay 2009).

Neosarmatium Serène & Soh, 1970 Diagnosis. Carapace wider than long, quadrate, dorsally strongly convex; with weak epibranchial tooth; front wider than half exorbital width, front nearly straight, without median concavity. Male cheliped palm upper surface flat, usually with longitudinal ridge; outer surface smooth or punstate, not tuberculate; inner surface with prominent vertical row of granules. Maximum cl. 37 mm (Pl. 70j). Estuarine, intertidal. Indo-West Pacific, Temperate Southern Africa. 19 species (Rahayu and Davie 2006: key to 16 Indo-West Pacific species; Schubart and Ng 2002: key to 14 species).

Neosesarma Serène & Soh, 1970 Diagnosis. Carapace epibranchial tooth acute, separated by deep sulcus from exorbital angle, with small second tooth; front nearly straight, without median concavity; postfrontal lobes obsolete. Male cheliped palm with 1 pectinate crest clearly separate from upper inner margin; dactylus with row of conical tubercles. Pereopods 2–5 dactyli about two-thirds length of propodi. Maximum cl. 26 mm. Intertidal. Central Indo-Pacific. 3 species (Davie 2012: key to species).

Orisarma Schubart & Ng, 2020 Diagnosis. Carapace quadrate–trapezoid, lateral margins parallel or convergent posteriorly; front wider than half exorbital width; with or without epibranchial tooth. Male chelipeds asymmetrical; cheliped palm upper surface without pectinate ridge; upper and outer surfaces tuberculate. Male sternite 5 without excavation on

each wall of sternopleonal cavity. Male pleon lateral margins straight to gently concave; pleonite 6 0.6 times width of pleonite 3; telson not embedded into pleonite 6, at most shallowly recessed. Pereopod 5 dactylus longer than upper margin of propodus Maximum cl. 47 mm. Intertidal, supralittoral, estuarine into fresh water. Central Indo-West Pacific. 6 species (Schubart and Ng 2020).

Parasesarma De Man, 1895 Diagnosis. Carapace lateral margins parallel over anterior half, convergent posteriorly; without epibranchial tooth, or with weak epibranchial tooth, or with prominent epibranchial tooth. Male cheliped palm upper surface with 1–3 (usually 2) transverse pectinate crests of tall chitinous teeth flanked by large tubercle on inside; dactylus tuberculate. Maximum cl. 25 mm (Pl. 70k, l). Estuarine, intertidal. Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 54 species (Li et al. 2018; Rahayu and Ng 2010: revision of P. plicatum group; Shahdadi and Schubart 2018: diagnosis, list of species; Shahdadi et al. 2021: P. calypso group). The genus is monophyletic (Wang et al. 2018).

Perisesarma De Man, 1895 Diagnosis. Carapace lateral margins parallel over anterior half, convergent posteriorly; with weak epibranchial tooth. Male cheliped palm upper surface with 2 transverse pectinate crests of tall chitinous teeth flanked by 2 small tubercles on inside; dactylus tuberculate. Maximum cl. 24 mm. Estuarine, intertidal. Western and Central Indo-Pacific. 2 species (Davie 2003: lists of species all but 2 now in other genera; Rahayu and Davie 2002; Shahdadi et  al. 2017: second species; Shahdadi and Schubart 2015: species identification; Shahdadi and Schubart 2018: diagnosis).

Platychirarma Schubart & Ng, 2020 Diagnosis. Carapace trapezoidal, lateral margins convergent posteriorly; front wider than half exorbital width, gently deflexed, not obscuring antennae from anterior view, straight in dorsal view; postfrontal and epigastric lobes merged, forming near-continuous transverse crest; without epibranchial tooth. Male chelipeds asymmetrical; cheliped palm upper surface without longitudinal pectinated ridge; outer surface flat. Male pleon lateral margins straight to gently concave; pleonite 6 0.6 width of pleonite 3; telson margins divergent, slightly wider than long, not embedded into pleonite 6. Pereopod 5 dactylus as long as or longer than upper margin of propodus. Gonopod 1 chitinous tip elongated, more than twice as long as wide. Maximum cl. 12 mm. Intertidal, supralittoral, estuarine. Western Indo-West Pacific. 1 species (Schubart and Ng 2020).

Pseudosesarma Serène & Soh, 1970 Diagnosis. Carapace wider than long, quadrate, dorsally convex; with or without epibranchial tooth; front wider than half exorbital width, with weak median concavity; postfrontal lobes obsolete.

14 – Brachyura – crabs

821

Male cheliped palm upper margin without pectinate ridge; upper and outer surfaces tuberculate; inner surface with vertical row of granules reaching upper margin. Male sternite 5 usually with excavation on each wall of sternopleonal cavity with excavation for reception of swollen gonopods 1. Male pleonite 6 two-thirds width of pleonite 3. Maximum cl. 20 mm. Estuarine, intertidal, supralittoral (mangroves, freshwater streams, caves). Central Indo-Pacific. 6 species (Ng 2018: species list; Ng and Schubart 2017: rediagnosis)

tooth; front gently sinuous, with median concavity. Male cheliped palm upper surface with longitudinal granular ridge, inner surface granular in median part, without row of granules. Pereopods 2–5 dactyli as long as propodi (about two-thirds in some cave species). Maximum cl. 36 mm. Estuarine, intertidal, supralittoral (including mangroves, freshwater streams to 1500 m asl, caves, terrestrial). Tropical Atlantic, Temperate Northern Atlantic, Tropical Eastern Pacific. 18 species (Abele 1992: diagnosis, key; Schubart and Cuesta 1998: larvae, key).

Sarmatium Dana, 1851

Sesarmoides Serène & Soh, 1970

Diagnosis. Carapace quadrate, dorsally strongly convex; front nearly straight, without median concavity. Cornea narrower than swollen peduncle. Male cheliped palm upper surface with 4–7 wide transverse grooves separating wide ridges, some pectinate; dactylus in male with 1 or more large wide chitinous tubercle. Maximum cl. 12 mm (Pl. 70m). Intertidal. Western and Central Indo-Pacific. 5 species (Davie 1992: key to species; Nagai et al. 2011: figure).

Diagnosis. Carapace wider than long, trapezoidal, anterolateral margins divergent posteriorly; front about half exorbital width, with weak median concavity; exorbital tooth weakly developed, incurved; postfrontal lobes obsolete. Antennular basal article swollen, little wider than long. Male cheliped merus inner upper angle forming tooth, inner surface with raised ridge forming stridulating organ; palm almost smooth. Pereopods 2–5 long, narrow; dactyli about as long as propodi. Maximum cl. 16 mm (Pl. 70o). Estuarine, supralittoral (coastal caves). Central Indo-Pacific. 3 species (Davie and Ng 2007: rediagnosis, key to species).

Scandarma Schubart, Liu & Cuesta, 2003 Diagnosis. Carapace wider than long, quadrate, lateral margins parallel to weakly convex laterally, dorsally almost flat; without or with 1 or 2 small epibranchial teeth; front about half exorbital width, front deflexed, distally recurved, directed anteriorly; margin concave medially in dorsal view; mesial and lateral pairs of postfrontal lobes sometimes weakly separated; not reaching beyond frontal margin in dorsal view. Male cheliped palm upper surface with longitudinal line of regular granules, outer surface with sloping smooth area delimited by median protuberance and base of fingers; fixed finger lower margin with row of ventral spiny granules; dactylus with longitudinal row of horny tubercles. Pereopods 2–5 dactyli longer than half length of propodi. Maximum cl. 20 mm. Freshwater, in forest, tree-climbing, nocturnal. Western and Central Indo-Pacific. 7 species (Naruse and Ng 2020: rediagnosis, key to 6 species; Schubart et al. 2003).

Sesarmops Serène & Soh, 1970 Diagnosis. Carapace as wide as long, widest posteriorly, dorsally almost flat; with prominent epibranchial tooth; front about half exorbital width, front with strong median concavity; postfrontal lobes well developed. Male cheliped palm upper surface without pectinate crest; upper and outer surfaces tuberculate; inner surface with vertical row of granules reaching upper margin. Pereopods 2–5 margins slightly convex; dactyli about as long as propodi. Maximum cl. 38 mm. Terrestrial. Indian Ocean, SE Asia, N Australia, W Pacific islands. 6 species (McLay and Ryan 1990; Naruse and Ng 2020: figures).

Shinobium Naruse & Ng, 2020

Diagnosis. Carapace quadrate to longitudinally rectangular, lateral margins parallel to slightly divergent posteriorly, with or without epibranchial tooth; front with strong median concavity; postfrontal lobes well developed. Male cheliped palm upper surface with 1 longitudinal crest of numerous tubercles, partly pectinate; dactylus with c. 30 tubercles. Pereopods 2–5 dactyli about one-third length of propodi. Maximum cl. 36 mm. Supralittoral, tree-climbing. Western and Central IndoPacific. 2 species (Schubart et al. 2009: rediagnosis, species).

Diagnosis. Carapace longer than width, trapezoidal; with 1 epibranchial tooth; front exceeding half exorbital width, front deflexed more than 90°, distally sloping ventroposteriorly; bilobed, recurved, directed anteriorly; lateral pair of postfrontal lobes slightly exceeding mesial pair; both reaching beyond frontal margin in dorsal view, overhanging front. Antennular basal article swollen, little wider than long. Male cheliped palm upper surface with irregular lines of granules, outer surface without prominent sloping smooth area; dactylus with row tubercles. Pereopods 2–5 dactyli shorter than half length of propodi. Maximum cl. 35 mm. Estuarine, freshwater, on vertical riverine cliffs. Indo-West Pacific. 1 species (Jeng et al. 2003: ecology; Naruse and Ng 2020).

Sesarma Say, 1817

Sinosesarma Ng, Shih & Canniccci, 2019

Diagnosis. Carapace slightly wider than long, margins subparallel, appearing square; dorsally almost flat; with or without epibranchial

Diagnosis. Carapace trapezoidal, lateral margins convergent posteriorly; front wider than half exorbital width, downcurved,

Selatium Serène & Soh, 1970

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obscuring antennae from anterior view, straight in dorsal view; without epibranchial tooth. Male cheliped merus inner margin weakly lamelliform; palm upper surface without longitudinal pectinate ridge; palm inner surface with low vertical granular row almost reaching upper margin; finger tips occlusal surface pectinate, upper margin with proximal row of minute corneous granules preceding row or widely spaced tubercles on central portion of margin. Maximum cl. 15 mm. Intertidal, supralittoral. Central Indo-West Pacific. 1 species (Ng et al. 2019).

Stelgistra Ng & Liu, 1999 Diagnosis. Carapace lateral margins convergent posteriorly; without epibranchial tooth. Male cheliped merus inner margin widely lamelliform; palm upper surface with scattered granules; finger tips hollowed, hoof-shaped occlusal surface smooth, not pectinate. Maximum cl. 17 mm. Supralittoral. Central Indo-Pacific. 1 species (Ng and Liu 1999).

Tiomanium Serène & Soh, 1970 Diagnosis. Carapace quadrate, dorsally strongly convex; front wider than half exorbital width, front nearly straight, without median concavity. Male cheliped palm upper surface sharp, ending in distal tooth; inner surface with denticles; carpus with long, acute inner angle. Maximum cl. 34 mm. Supralittoral. Western and Central Indo-Pacific. 1 species (Li 2014: redescription, figure).

Trapezarma Schubart & Ng, 2020 Diagnosis. Carapace trapezoidal, lateral margins convergent posteriorly; front wider than half exorbital width, gently deflexed, not obscuring antennae from anterior view, straight in dorsal view; postfrontal and epigastric lobes merged, forming near-continuous transverse crest; without epibranchial tooth. Male chelipeds asymmetrical; cheliped palm upper surface without longitudinal pectinated ridge; outer surface flat. Male pleon lateral margins straight to gently concave; pleonite 6 two-thirds width of pleonite 3; telson margins parallel, slightly longer than wide, not embedded into pleonite 6. Pereopod 5 dactylus as long as or longer than upper margin of propodus. Gonopod 1 chitinous tip short, about as long wide. Maximum cl. 34 mm. Intertidal, supralittoral, estuarine into freshwater. Tropical E Atlantic. 1 species (Schubart and Ng 2020). References Abele LG (1992) A review of the grapsid crab genus Sesarma (Crustacea: Decapoda: Grapsidae) in America, with the description of a new genus. Smithsonian Contributions to Zoology 527, 1–60. doi:10.5479/ si.00810282.527 Boon PY, Yeo DCJ, Todd PA (2009) Sound production and reception in mangrove crabs Perisesarma spp. (Brachyura: Sesarmidae). Aquatic Biology 5, 107–116. doi:10.3354/ab00136

Brösing A, Spiridonov VA, Al-Aidaroos AM, Türkay M (2014) Description of a new genus and new species of Sesarmidae (Decapoda: Brachyura) from the Farasan Islands, Saudi Arabia, Red Sea. Journal of Crustacean Biology 34, 273–282. doi:10.1163/1937240X-00002223 Calman WT (1925) A new crab of the genus Sesarma from New Guinea. Annals and Magazine of Natural History (ser. 9) 15, 454–456. Cannicci S, Ng PKL (2017) A new species of micro-mangrove crab of the genus Haberma Ng & Schubart, 2002 (Crustacea, Brachyura, Sesarmidae) from Hong Kong. ZooKeys 662, 67–78. doi:10.3897/ zookeys.662.11908 Cuesta JA, García-Guerrero MU, Hendrickx M, Rodríguez A (2006a) Larval morphology of the sesarmid crab, Aratus pisonii (H. Milne Edwards, 1837) (Decapoda, Brachyura, Grapsoidea) from laboratory-reared material. Crustaceana 79, 175–196. doi:10.1163/156854006776952838 Cuesta JA, Gueroa G, Liu H-C, Schubart CD (2006b) Morphology of the first zoeal stages of eleven Sesarmidae (Crustacea, Brachyura, Thoracotremata) from the Indo-West Pacific, with a summary of familial larval characters. Invertebrate Reproduction & Development 49, 151–173. doi:10.1080/07924259.2006.9652206 Davie PJF (1992) Revision of Sarmatium Dana (Crustacea: Brachyura: Sesarminae) with descriptions of three new species. Memoirs of the Queensland Museum 32, 79–97. Davie PJF (1994) Variations in diversity of mangrove crabs in tropical Australia. Memoirs of the Queensland Museum 36, 55–58. Davie PJF (2003) A new species of Perisesarma (Crustacea: Brachyura: Sesarmidae) from the Bay of Bengal. Raffles Bulletin of Zoology 51, 387–391. Davie PJF (2010) A new species of Perisesarma (Brachyura, Sesarmidae) from Western Australia. In: Fransen C, De Grave S, Ng P (Eds) Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume. Crustaceana Monographs 14, 195–207. doi:10.1163/9789047427759_012 Davie PJF (2012) A revision of Neosesarma (Crustacea: Brachyura: Sesarmidae) with the description of a new species. Memoirs of the Queensland Museum - Nature 56, 221–233. Davie PJF, Ng PKL (2007) A new genus for cave-dwelling crabs previously assigned to Sesarmoides (Crustacea: Decapoda: Brachyura: Sesarmidae). Raffles Bulletin of Zoology 16, 227–231. Davie PJF, Ng PKL (2013) A review of Chiromantes obtusifrons (Dana, 1851) (Decapoda: Brachyura: Sesarmidae), with descriptions of four new sibling-species from Christmas Island (Indian Ocean), Guam and Taiwan. Zootaxa 3609, 1–25. doi:10.11646/zootaxa.3609.1.1 Dole-Olivier M-J, Galassi DMP, Marmonier P, Creuzé Des Châtelliers M (2000) The biology and ecology of lotic microcrustaceans. Freshwater Biology 44, 63–91. doi:10.1046/j.1365-2427.2000.00590.x Fratini S, Vannini M, Cannicci S, Schubart CD (2005) Tree-climbing mangrove crabs: a case of convergent evolution. Evolutionary Ecology Research 7, 219–233. Goh SLR, Vishnu H, Ng NK (2019) The sounds of fighting: contests between violet vinegar crabs, Episesarma versicolor (Tweedie, 1940) (Decapoda: Brachyura: Sesarmidae), are resolved through acoustic communication. Journal of Crustacean Biology 39, 331–341. doi:10.1093/​ jcbiol/ruz023 Jeng M-S, Liu H-C, Tzeng C-S, Ng PKL (2003) On the taxonomy and ecology of Labuanium trapezoideum (Decapoda, Brachyura, Sesarmidae), a crab living on riverine cliffs in Taiwan. Crustaceana 76, 227–240. doi:10.1163/156854003321824576 Kawaida S, Nanjo K, Ohtsuchi N, Kohno H, Sano M (2019) Cellulose digestion abilities determine the food utilization of mangrove estuarine crabs. Estuarine, Coastal and Shelf Science 222, 43–52. doi:10.1016/j.ecss.2019.04.004 Lee BY, Ng NK, Ng PKL (2013) On the identity of Clistocoeloma balansae A. Milne-Edwards, 1873, and C. tectum (Rathbun, 1914), with description

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of a new species from the West Pacific (Crustacea: Decapoda: Sesarmidae). Zootaxa 3641, 420–432. doi:10.11646/zootaxa.3641.4.8 Lee BY, Ng NK, Ng PKL (2015) The taxonomy of five species of Episesarma De Man, 1895, in Singapore (Crustacea: Decapoda: Brachyura: Sesarmidae). Raffles Bulletin of Zoology Supplement 31, 199–215. Li J-J (2014) Redescription of two poorly known sesarmid crabs from Taiwan. Platax 2014, 83–93. Li JJ, Rahayu DL, Ng PKL (2018) Identity of the tree-spider crab, Parasesarma leptosoma (Hilgendorf, 1869 (Decapoda: Brachyura: Sesarmidae), with descriptions of seven new species from the Western Pacific. Zootaxa 4482, 451–490. doi:10.11646/zootaxa.4482.3.2 Li J-J, Shih H-T, Ng PKL (2020) The Taiwanese and Philippine species of the terrestrial crabs Bresedium Serène and Soh, 1970 and Sesarmops Serène and Soh, 1970 (Crustacea: Decapoda: Brachyura), with descriptions of two new species. Zoological Studies 59, e16. Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 McLay CL, Ryan PA (1990) The terrestrial crabs Sesarma (Sesarmops) impressum and Geograpsus crinipes (Brachyura, Grapsidae, Sesarminae) recorded from the Fiji Is. Journal of the Royal Society of New Zealand 20, 107–118. doi:10.1080/03036758.1990.10426735 Miranda JF, Mendoza-Carranza M, Sánchez AJ, Barba E (2017) Selective foraging of Aratus pisonii (Arthropoda: Sesarmidae) on mangrove leaves in laboratory experiments. Journal of Experimental Marine Biology and Ecology 488, 38–43. doi:10.1016/j.jembe.2016.12.012 Naderloo R, Türkay M (2009) A new species of the genus Nanosesarma (Crustacea: Decapoda: Brachyura: Sesarmidae), and redescription of Nanosesarma jousseaumei (Nobili, 1906) including new records from the Persian Gulf. Journal of Natural History 43, 2911–2923. doi:10.1080/00222930903219996 Nagai T, Naruse T, Maenosono T, Fujita Y, Komai T (2011) Taxonomy and distribution of the three sesarmid genera, Neosarmatium, Sarmatium and Tiomanium (Crustacea: Decapoda: Brachyura) in the Ryukyu Archipelago. Biological Magazine Okinawa 49, 15–36. Naruse T, Ng NK (2012) Establishment of a new genus for Cyclograpsus lophopus Nobili, 1905, within Sesarmidae Dana, 1851 (Crustacea: Decapoda: Brachyura). Zootaxa 3572, 63–68. doi:10.11646/zootaxa.​ 3572.1.8 Naruse T, Ng PKL (2020) Revision of the sesarmid crab genera Labuanium Serène and Soh, 1970, Scandarma Schubart, Liu and Cuesta, 2003 and Namlacium Serène and Soh, 1970 (Crustacea: Decapoda: Brachyura: Sesarmidae), with descriptions of four new genera and two new species. Journal of Natural History 54, 445–532. doi:10.1080/00222933. 2020.1763491 Ng PKL (2018) On two new species of potamid and sesarmid crabs (Decapoda, Brachyura) from caves in Myanmar. Crustaceana 91, 185–197. doi:10.1163/15685403-00003735 Ng PKL, Davie PJF (1995) The terrestrial sesarmine crabs of the genera Metasesarma and Geosesarma (Crustacea: Decapoda: Brachyura: Grapsidae) of Ujung Kulon, West Jawa, Indonesia. Tropical Biodiversity 3, 29–43. Ng PKL, Davie PJF (2011) Labuanium vitatum (Crustacea: Decapoda: Brachyura: Sesarmidae), a new Indo-West Pacific species of arboreal crab. Zootaxa 2889, 35–48. doi:10.11646/zootaxa.2889.1.2 Ng PKL, Liu H-C (1999) The taxonomy of Sesarma tangi Rathbun, 1931, and S. stormi De Man, 1895 (Crustacea: Decapoda: Brachyura: Grapsidae: Sesarminae), with establishment of a new genus for S. stormi. Zoological Studies 38, 228–237. Ng PKL, Schubart CD (2002) Haberma nanum, a new genus and new species of mangrove crab (Crustacea: Decapoda: Brachyura: Sesarmidae) from Singapore. Raffles Bulletin of Zoology 50, 437–442.

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Ng PKL, Schubart CD (2017) On the taxonomy of Pseudosesarma edwardsii (De Man, 1887) and P. crassimanum (De Man, 1887) (Crustacea: Decapoda: Brachyura: Sesarmidae), with description of a new species from Sri Lanka. Raffles Bulletin of Zoology 65, 655–669. Ng PKL, Wowor D (2019) The vampire crabs of Java, with descriptions of five new species from Mount Halimun Salak National Park, West Java, Indonesia (Crustacea: Brachyura: Sesarmidae: Geosesarma). Raffles Bulletin of Zoology 67, 217. Ng PKL, Lee BY, Tan HH (2015) Notes on the taxonomy and ecology of Labuanium politum (De Man, 1887) (Crustacea: Decapoda: Sesarmidae), an obligate arboreal crab on the nipah palm, Nypa fruticans (Arecales: Arecaceae). Raffles Bulletin of Zoology Supplement 31, 216–225. Ng PKL, Shih HT, Cannicci S (2019) A new genus for Sesarma (Holometopus) tangi Rathbun, 1931 (Decapoda: Brachyura: Sesarmidae) from mangrove forests, with notes on its ecology and conservation. Journal of Crustacean Biology 40, 89–96. doi:10.1093/jcbiol/ruz078 Ngo-Massou VM, Din N, Kenne M, Dongmo AB (2018) Brachyuran crab diversity and abundance patterns in the mangroves of Cameroon. Regional Studies in Marine Science 24, 324–335. doi:10.1016/j.rsma.​ 2018.09.010 Niem VH (1996) Phylogenetic relationships among American species of Sesarma (subgenus Armases) (Brachyura, Grapsidae). Crustaceana 69, 330–348. doi:10.1163/156854096X00934 Poupin J, Crestey N, Guelte JL (2018) Cave-dwelling crabs of the genus Karstarma from lava tubes of the volcano ‘Piton de la Fournaise’, in Reunion Island, with description of a new species and redescription of Karstarma jacksoni (Balss, 1934) from Christmas Island (Decapoda, Brachyura, Sesarmidae). Zootaxa 4497, 381–397. doi:10.11646/zootaxa.​ 4497.3.3 Ragionieri L, Fratini S, Vannini M, Schubart CD (2009) Phylogenetic and morphometric differentiation reveal geographical radiation and pseudo-cryptic speciation in a mangrove crab from the Indo-West Pacific. Molecular Phylogenetics and Evolution 52, 825–834. doi:10.1016/​ j.ympev.2009.04.008 Rahayu DL, Davie PJF (2002) Two new species and a record of Perisesarma (Decapoda, Brachyura, Grapsidae, Sesarminae) from Indonesia. Crustaceana 75, 597–607. doi:10.1163/156854002760095624 Rahayu DL, Davie PJF (2006) Two new species of mangrove crabs of the genus Neosarmatium Serène and Soh, 1970 (Decapoda, Brachyura, Sesarmidae) from Papua, Indonesia. Zoosystema 28, 573–584. Rahayu DL, Ng PKL (2010) Revision of the Parasesarma plicatum (Latreille, 1803) species-group (Crustacea: Decapoda: Brachyura: Sesarmidae). Zootaxa 2327, 1–22. doi:10.11646/zootaxa.2327.1.1 Rathbun MJ (1897) Description of a new genus and four new species of crabs from the West Indies. Proceedings of the United States National Museum 19, 141–144. doi:10.5479/si.00963801.1104.141 Rathbun MJ (1918) The grapsoid crabs of America. Bulletin of the United States National Museum 97, 1–461, pls 1–161. Schubart CD (2011) Reconstruction of phylogenetic relationships within Grapsidae (Crustacea: Brachyura) and comparison of trans-isthmian versus amphi-atlantic gene flow based on mtDNA. Zoologischer Anzeiger 250, 472–478. doi:10.1016/j.jcz.2011.06.003 Schubart CD, Cuesta JA (1998) First zoeal stages of four Sesarma species from Panama, with identification keys and remarks on the American Sesarminae (Crustacea: Brachyura: Grapsidae). Journal of Plankton Research 20, 61–84. doi:10.1093/plankt/20.1.61 Schubart C, Ng PKL (2002) The sesarmid genus Neosarmatium (Decapoda: Brachyura): new distribution records and a new species from Sulawesi. Crustacean Research 31, 28–38. doi:10.18353/crustacea.31.0_28

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Schubart CD, Ng PKL (2020) Revision of the intertidal and semiterrestrial crab genera Chiromantes Gistel, 1848, and Pseudosesarma Serène & Soh, 1970 (Crustacea: Brachyura: Sesarmidae), using morphology and molecular phylogenetics, with the establishment of nine new genera and two new species. Raffles Bulletin of Zoology 68, 891–994. Schubart CD, Liu HC, Cuesta JA (2003) A new genus and species of treeclimbing crab (Crustacea: Brachyura: Sesarmidae) from Taiwan with notes on its ecology and larval morphology. Raffles Bulletin of Zoology 51, 49–60. Schubart CD, Cannicci S, Vannini M, Fratini S (2006) Molecular phylogeny of grapsoid crabs (Decapoda, Brachyura) and allies based on two mitochondrial genes and a proposal for refraining from current superfamily classification. Journal of Zoological Systematics and Evolutionary Research 44, 193–199. doi:10.1111/j.1439-0469.2006.​ 00354.x Schubart CD, Liu H-C, Ng PKL (2009) Revision of Selatium Serène & Soh, (Crustacea: Brachyura: Sesarmidae), with description of a new genus and two new species. Zootaxa 2154, 1–29. doi:10.11646/zootaxa.​ 2154.1.1 Serène R, Soh CL (1970) New Indo-Pacfic genera allied to Sesarma Say 1817 (Brachyura, Decapoda, Crustacea). Treubia 27, 387–431. Shahdadi A, Schubart CD (2015) Evaluating the consistency and taxonomic importance of cheliped and other morphological characters that potentially allow identification of species of the genus Perisesarma De Man, 1895 (Brachyura, Sesarmidae). Crustaceana 88, 1079–1095. doi:10.1163/15685403-00003473 Shahdadi A, Schubart CD (2018) Taxonomic review of Perisesarma (Decapoda: Brachyura: Sesarmidae) and closely related genera based on morphology and molecular phylogenetics: new classification, two new genera and the questionable phylogenetic value of the epibranchial tooth. Zoological Journal of the Linnean Society 182, 517–548. doi:10.1093/zoolinnean/zlx032 Shahdadi A, Davie PJF, Schubart CD (2017) Perisesarma tuerkayi, a new species of mangrove crab from Vietnam (Decapoda, Brachyura, Sesarmidae), with an assessment of its phylogenetic relationships. Crustaceana 90, 1155–1175. doi:10.1163/15685403-00003663 Shahdadi A, Schubart CD, Mendoza JCE (2021) Conspicuous genetic similarity within a widely distributed and newly described species of Parasesarma De Man, 1895 from Western pacific oceanic islands, with notes on the allied P. calypso group (Crustacea: Brachyura: Sesarmidae). Invertebrate Systematics 35, 542–569. Shanij K, Praveen VP, Suresh S, Oommen MM, Nayar TS (2016) Tree climbing and temporal niche shifting: an anti-predatory strategy in the mangrove crab Parasesarma plicatum (Latreille, 1803). Current Science 111, 1201–1207. doi:10.18520/cs/v111/i7/1201-1207 Thiercelin N, Schubart CD (2014) Transisthmian differentiation in the tree-climbing mangrove crab Aratus H. Milne Edwards, 1853 (Crustacea, Brachyura, Sesarmidae), with description of a new species from the tropical eastern Pacific. Zootaxa 3793, 545–560. doi:10.11646/ zootaxa.3793.5.3 Wang Z, Wang Z, Shi X, Wu Q, Tao Y, et al. (2018) Complete mitochondrial genome of Parasesarma affine (Brachyura: Sesarmidae): gene rearrangements in Sesarmidae and phylogenetic analysis of the Brachyura. International Journal of Biological Macromolecules 118, 31–40. doi:10.1016/j.ijbiomac.2018.06.056 Wowor D, Ng PKL (2018) A new sesarmid crab of the genus Karstarma (Crustacea: Decapoda: Brachyura) associated with limestone formations in East Java, Indonesia. Zootaxa 4482, 355–366. doi:10.11646/ zootaxa.4482.2.7

Varunidae H. Milne Edwards, 1853 Figures 14.182–14.189, Plate 71a–g Varunids are typical crabs of muddy shores. Most scavenge on detritus, hide under rocks or live in burrows. Very few are known to live exclusively subtidally – in contrast, many live high on the shore or inhabit estuarine environments. Most are small and cryptic, but the mitten crabs (Eriocheir), the largest of the varunids, are highly prized as culinary delicacies in East and South-east Asia, as well as being major invasive pests in Europe (Naser et al. 2012). Varunidae, long considered a subfamily of Grapsidae, was settled as a full family after long debate (Davie et al. 2015a, 2015b). Their separation from Sesarmidae, in spite of similar stridulatory systems in some genera, has become more settled and subfamilies have emerged (Davie and Ng 2007; Davie and Nguyen 2003). The most thorough treatment of Varunidae is a PhD thesis by Ng Ngan Kee (Ng 2006) in which new genera and species were introduced. Some of the names proposed have been formally published subsequently. The work has proved a valuable resource for the keys to subfamilies and genera in this work. More recently, the Cyclograpsus-like Pinnotherelia laevigata (Kinahan, 1857) (Pinnotherelinae), long considered to be a pinnotheroid (though originally described as a species of Cyclograpsus), was recently shown to be varunid (Palacios Theil et al. 2016). Pinnotherelia requires detailed revision but we herein formally include Pinnotherelinae as a varunid subfamily. The key does not include Grapsodius Holmes, 1900, a poorly described and problematic genus (Ng 2006; Schubart et al. 2000). Diagnosis. Carapace subquadrate-polygonal, smooth; front entire; orbit with lateral opening; suborbital crest granulate or dentate, stridulatory; pterygostomial region sparingly setose, without pattern of reticulated setae, or covered in reticulated network of short, hooked, setae, without groove subparallel to lateral margin of buccal cavity. Maxilliped 3 merus and ischium with or without distinct oblique setose ridge; distinct rhomboidal gap between closed maxillipeds 3 exposing mandibles or very small rhomboidal gap concealing mandibles. Male chelipeds equal; dactylus straight, usually with large truncate tooth at or proximal to midlength of occlusal margin. Male pleonites 1–6 and telson free, or pleonites 3–6 fused. Implicit subfamily attributes. Carapace quadrate, lateral margins almost parallel, or ovate, lateral margins convex. Frontal margin sinuous. Pterygostome without complex reticulation of setae, without vertical groove parallel to buccal cavity. Maxilliped 3 ischium-merus with longitudinal sulcus; palp not elongate, without setal brush, without sternal groove. Ambulatory legs propodi without distal spine on flexor margin.

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825

Key to subfamilies of Varunidae 1. – 2. – 3. – 4. – 5. –

Carapace pterygostome with complex reticulation of setae, with deep vertical groove parallel to buccal cavity. Maxilliped 3 with oblique setose crest across merus and distolateral corner of ischium (Fig. 14.183i, j)���������������������������������������������������������������������������������������������������������������������������������������������������� Cyclograpsinae Carapace pterygostome without complex reticulation of setae, without vertical groove parallel to buccal cavity. Maxilliped 3 without oblique setose crest across merus (usually with longitudinal sulcus)������������������������������2 Maxillipeds 3 palp with dense brush of long setae resting on sternal plastron (Figs 14.184j–n, 14.185b). Thoracic sternal plastron usually with medial sulcus or shallow groove (accommodating maxillipedal setae) (Fig. 14.184l)����������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Maxillipeds 3 palp without dense brush of long setae. Male pleonites 3–6 usually free (Fig. 14.182b, 14.188u) or 5–6 fused (Fig. 14.186b)�����������������������������������������������������������������������������������������������������������������������������������������4 Male pleonites 3–6 fused (Fig. 14.184h, o)�������������������������������������������������������������������������������������������������������������������Gaeticinae Male pleonites 5–6 fused (Fig. 14.185c)������������������������������������������������������������������������������������������������������������� Pinnotherelinae Carapace trapezoidal, greatest width posterior to midlength (Fig. 14.182a)�������������������������������������������Asthenognathinae Carapace subquadrate or oval, greatest width usually at or close to exorbital width, never twice exorbital width�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������5 Lateral edge of carapace front with short oblique sulcus separating frontal margin from inner orbital angle (Fig. 14.186a). Maxillipeds 3 medial margins contiguous when closed. Pereopods 2–5 propodi with distal spine on flexor margin (Fig. 14.186d)�����������������������������������������Thalassograpsinae Carapace frontal margin sinuous or straight. Maxillipeds 3 moderately gaping when closed (Fig. 14.188t). Pereopods 2–5 propodi without distal spine on flexor margin��������������������������������������������������������������������������Varuninae

Subfamily Asthenognathinae Stimpson, 1858 Asthenognathidae and its only genus were once considered as members of Pinnotheroidea based on their reduced body ornamentation and commensal associations with infaunal invertebrates and holothurian. Their transfer to Varunidae as a subfamily by (Ng et al. 2008) was justified based on comparative morphology of sternal, mouthpart and reproductive structures and corroborated by molecular phylogenetic analysis (Palacios Theil et al. 2016). The only genus is immediately distinguished from other varunids by its wide trapezoidal carapace shape. Diagnosis. Carapace trapezoid, greatest width twice extraorbital width. Male pleonites 1–6 movable. Maxillipeds 3 broadly gaping. Asthenognathus Stimpson, 1858

(Fig.  14.183i, j). These structures are significant because several species spend considerable time out of the water (Marsden and Dewa 1994). In this they resemble Sesarmidae, but sesarmids lack the vertical groove running alongside the buccal cavity (Davie 2002). The distinction between these family groups has been confirmed by molecular analysis (Schubart et  al. 2006) indicating that these structures have arisen more than once in grapsoid crabs. Not all species are as round and smooth as the subfamily name suggests. Diagnosis. Carapace pterygostome with complex reticulation of setae, with deep vertical groove parallel to buccal cavity (Fig. 14.183i, j). Male pleonites 1–6 movable. Maxillipeds 3 separated by narrow gape when closed; maxilliped 3 with oblique setose crest across merus and distolateral corner of ischium (Fig. 14.183i, j).

Maximum cl. 9 mm. Intertidal–shelf (0–210 m; associated with holothurians and burrowing worms). Temperate Northern E Atlantic, Central Indo-Pacific. 3 species. Yang and Tang’s (2008) key to species included A. gallardoi Serène & Soh, 1976, a species removed to Gopkittisak, a genus of Gaeticinae, by Naruse and Clark (2009).

Subfamily Cyclograpsinae H. Milne Edwards, 1853 Cyclograpsines are recognised by the water-channelling setose ridges on maxilliped 3 and the setose pterygostome

Fig. 14.182.  Varunidae. Asthenognathinae. Asthenognathus: a, carapace; b, male pleon, telson; c, maxilliped 3.

826

Marine Decapod Crustacea

Key to genera of Cyclograpsinae 1. – 2. – 3. – 4. – 5. – 6. – 7. –

Carapace subquadrilateral to rounded, broader than long, regions not clearly defined; anterolateral margin with wide exorbital tooth plus 1 or 2 lobes or small teeth (Fig. 14.183c)�������������������������������������������������������Cyclograpsus Carapace subquadrilateral, regions well defined, anterolateral margin with distinct tooth or teeth���������������������������������2 Carapace anterolateral margin with 1 sharp tooth behind exorbital tooth (Fig. 14.183d)������������������������������� Helograpsus Carapace anterolateral margin with 2 teeth behind exorbital tooth����������������������������������������������������������������������������������������3 Carapace frontal margin projecting forward over antennules, slightly concave (Fig. 14.183f)����������������������� Paragrapsus Carapace frontal margin not projecting, weakly bilobed or almost straight��������������������������������������������������������������������������4 Eyestalk about as long as front, each occupying third of distance between exorbital angles (Fig. 14.183e)������������������������������������������������������������������������������������������������������������������������������������������������������������������Metaplax Eyestalk half or less width of front, each occupying much less than third of distance between exorbital angles���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������5 Maxilliped 3 merus about half as wide distally as long����������������������������������������������������������������������������������� Chasmagnathus Maxilliped 3 merus about as wide distally as long����������������������������������������������������������������������������������������������������������������������6 Male suborbital ridge with granules of similar sizes (Fig. 14.183k)������������������������������������������������������������������������������������������7 Male suborbital ridge with granules of different sizes (Fig. 14.183l–n)�����������������������������������������������������������������������������������8 Male suborbital ridge of 2 rows of granules (Fig. 14.183j)�������������������������������������������������������������������������������������������Neohelice Male suborbital ridge of 1 row of granules (Fig. 14.183k)��������������������������������������������������������������������������������������Austrohelice

Fig. 14.183.  Varunidae. Cyclograpsinae. a, Austrohelice; b, Chasmagnathus; c, Cyclograpsus granulosus H. Milne Edwards, 1853; d, Helograpsus haswellianus (Whitelegge, 1889); e, Metaplax; f, Paragrapsus gaimardii (H. Milne Edwards, 1837); g, Parahelice; h, Pseudohelice subquadrata (Dana, 1851). Buccal cavity, maxillipeds 3, pterygostomial region (p groove and oblique setose crest arrowed): i, Helice; j, Neohelice. Suborbital ridge, male: k, Austrohelice; l, Helicana; m, Parahelice; n, Pseudohelice. Gonopod 1: o, Helicana; p, Helice; q, Parahelice; r, Pseudohelice.

14 – Brachyura – crabs

8. – 9. – 10. –

827

Male suborbital ridge lateral part with 5 or fewer large tubercles (Fig. 14.183m, n)��������������������������������������������������������������9 Male suborbital ridge lateral part with more than 5 large tubercules (Fig. 14.183i, l)���������������������������������������������������������10 Gonopod 1 with sperm channel suture straight; horny tip elongate, clearly flattened (Fig. 14.183r)������������ Pseudohelice Gonopod 1 with sperm channel suture twisted; horny tip scarcely flattened, thick to tubular (Fig. 14.183q)�Parahelice Gonopod 1 distal part broad, tip thickened, triangular (Fig. 14.183p)���������������������������������������������������������������������������Helice Gonopod 1 distal part slender, tip small (Fig. 14.183o)������������������������������������������������������������������������������������������������Helicana Austrohelice Sakai, Türkay & Yang, 2006

Diagnosis. Cyclograpsinae. Carapace quadrilateral, wider than long; frontal margin not projecting, almost straight; anterolateral margin with 2 teeth behind exorbital tooth; male suborbital ridge with granules of similar sizes in 1 row. Maximum cl. 10 mm. Intertidal. Temperate Australasia (New Zealand only). 1 species (McLay 1988: description, biology as Helice crassa Dana, 1851; Sakai et al. 2006).

Chasmagnathus De Haan, 1833 Diagnosis. Cyclograpsinae. Carapace quadrilateral, wider than long; frontal margin sinuous; anterolateral margin with 3 low teeth behind exorbital tooth; male suborbital ridge bearing strong crests in median region. Maxilliped 3 merus about half as wide distally as long. Maximum cl. 30 mm. Estuarine, intertidal (nocturnal). Central Indo-Pacific. 1 species (Sakai et al. 2006: rediagnosis, details of stridulating apparatus, key to 6 similar genera).

Cyclograpsus H. Milne Edwards, 1837 Diagnosis. Cyclograpsinae. Carapace subquadrilateral to rounded, broader than long, regions not clearly defined; frontal margin smoothly deflexed, straight; anterolateral margin with wide exorbital tooth plus 1 or 2 lobes or microscopically notched; male suborbital ridge with granules of similar sizes in 1 row. Maximum cl. 40 mm (Pl. 71b). Estuarine, marine, intertidal (muddy shores, rocky reefs, under rocks). Cosmopolitan on temperate and tropical shores. 20 species (Campbell and Griffin 1966: diagnosis, key to 13 IndoWest Pacific species; Hangai et al. 2009: cryptic Japanese species).

Helice De Haan, 1833 Diagnosis. Cyclograpsinae. Carapace quadrilateral, wider than long; frontal margin not projecting, almost straight; anterolateral margin with 3 teeth behind exorbital tooth, plus more posterior faint notch; male suborbital ridge with granules of different sizes, lateral part with more than 5 large tubercles. Maximum cl. 26 mm. Estuarine, intertidal. Temperate Northern Pacific, Central IndoPacific. 5 species (Ng et al. 2018: fifth species; Sakai et al. 2006: key to 4 species; Yin et al. 2009: phylogenetic network of 4 species).

Helicana Sakai & Yatsuzuka, 1980 Diagnosis. Cyclograpsinae. Carapace quadrilateral, wider than long; frontal margin sinuous; anterolateral margin with 3 teeth

behind exorbital tooth, plus more posterior faint notch; male suborbital ridge with granules of different sizes, lateral part with more than 5 large tubercles. Maximum cl. 24 mm. Estuarine, intertidal. Temperate Northern W Pacific, Central Indo-Pacific. 3 species (Sakai et al. 2006: rediagnosis, key to species; Sakai and Yatsuzuka 1980).

Helograpsus Campbell & Griffin, 1966 Diagnosis. Cyclograpsinae. Carapace quadrilateral, wider than long; frontal margin not projecting, almost straight; anterolateral margin with 1 sharp tooth behind exorbital tooth; male suborbital ridge with granules of similar sizes in 1 row. Maximum cl. 20 mm (Pl. 71c). Estuarine, intertidal. Central Indo-Pacific, Temperate E and SE Australia. 1 species (Campbell and Griffin 1966; Poore 2004).

Metaplax H. Milne Edwards, 1852 Diagnosis. Cyclograpsinae. Carapace hexagonal, lateral margin convex, broader than long; widest at hepatic region; frontal margin short, about as long as each eyestalk; anterolateral margin with 2–4 teeth behind exorbital tooth. Maximum cl. 18 mm (Pl. 71e). Estuarine, intertidal. Western and Central Indo-Pacific. 11 species (Davie and Nguyen 2003: discussion of species).

Neohelice Sakai, Türkay & Yang, 2006 Diagnosis. Cyclograpsinae. Carapace quadrilateral, wider than long; frontal margin sinuous; anterolateral margin with 2 teeth behind exorbital tooth; male suborbital ridge with granules of similar sizes in 2 rows. Maximum cl. 25 mm. Intertidal. Tropical Atlantic, Temperate South America (Atlantic coast). 1 species (Ituarte et  al. 2012: population structure; Sakai et al. 2006).

Paragrapsus H. Milne Edwards, 1853 Diagnosis. Cyclograpsinae. Carapace lateral margin convex, broader than long; frontal margin projecting forward over antennules, slightly concave; anterolateral margin with 1 or 2 teeth behind exorbital tooth; male suborbital ridge absent. Maximum cl. 45 mm (Pl. 71f). Intertidal, rocky shores. Temperate SE Australia. 3 species (Poore 2004: key to species).

828

Marine Decapod Crustacea

Parahelice Sakai, Türkay & Yang, 2006 Diagnosis. Cyclograpsinae. Carapace quadrilateral, wider than long; frontal margin not projecting, weakly bilobed; anterolateral margin with 2 teeth behind exorbital tooth; male suborbital ridge with granules of different sizes, lateral part with 5 or fewer large tubercles. Gonopod 1 with sperm channel suture twisted; horny tip scarcely flattened, thick to tubular. Maximum cl. 22 mm. Estuarine, intertidal, mangroves. Western and Central IndoPacific. 5 species (Sakai et al. 2006: diagnosis as subgenus, key to species of Pseudohelice and Parahelice).

Pseudohelice Sakai, Türkay & Yang, 2006 Diagnosis. Cyclograpsinae. Carapace quadrilateral, wider than long; frontal margin not projecting, weakly bilobed; anterolateral margin with 3 teeth behind exorbital tooth, plus more posterior faint notch; male suborbital ridge 2–4 striated crests in lateral half, 4–6 low closely-set tubercles, 7–11 tubercles at mesial end. Gonopod 1 with sperm channel suture straight, not twisted; horny tip elongate, clearly flattened. Maximum cl. 24 mm.

Intertidal, mudflats. Temperate Northern Pacific, Indo-West Pacific, Temperate SE Australia. 2 species (Poore 2004: discussed as Helice leachii Hess, 1865; Hsu et al. 2022a: genus revised).

Subfamily Gaeticinae Davie & Ng, 2007 Gaeticines differ from other varunids in usually inhabiting burrows excavated by other invertebrates, notably upogebiid shrimps or echiuroid worms. Some may make their own burrows. The long setae at the end of maxillipeds 3 are used for filter-feeding to supplement their usual scavenging diet (Depledge 1989; Wahyudi et  al. 2015). Two genera were included in the family when it was distinguished from other varunids by Davie and Ng (2007). Naruse (2015) reviewed and tabulated differences between six genera and offered a key to five genera. Diagnosis. Male pleonites 3–6 fused (Fig.  14.184o), sutures sometimes visible (Fig. 14.184h). Maxillipeds 3 separated by narrow gape when closed; maxilliped 3 palp elongate, with long setal brush usually lying at rest in sternal groove or concavity (Fig. 14.184j–n).

Key to genera of Gaeticinae 1. – 2.

Maxilliped 3 ischium-merus suture oblique (Fig. 14.184k, l)����������������������������������������������������������������������������������������������������2 Maxilliped 3 ischium-merus suture transverse (Fig. 14.184j, m, n)�����������������������������������������������������������������������������������������4 Carapace quadrate, slightly wider than long (Fig. 14.184c)��������������������������������������������������������������������������������������������Gaetice

Fig. 14.184.  Varunidae, Gaeticinae. Carapace: a, Acmaeopleura; b, Brankocleistostoma; c, Gaetice; d, Gopkittisak; e, Proexotelson; f, Pseudopinnixa; g, Sestrostoma. Anterior thoracic sternites: h, Proexotelson (with male pleon, telson) i, Sestrostoma. Maxilliped 3: j, Acmaeopleura; k, Gaetice; l, Gopkittisak; m, Pseudopinnixa; n, Sestrostoma. Male pleon, telson: o, Gopkittisak.

14 – Brachyura – crabs

– 3. – 4. – 5. – 6. –

829

Carapace elliptical-rectangular, distinctly wider than long (Fig. 14.184b, d)�������������������������������������������������������������������������3 Carapace anterolateral margin straight, merging with posterolateral margin (Fig. 14.184d)�������������������������� Gopkittisak Carapace anterolateral margin undulate, forming angle with posterolateral margin (Fig. 14.184b)� Brankocleistostoma Male sternopleonal cavity, telson, gonopod 1 entering buccal cavity (Fig. 14.184h)���������������������������������������� Proexotelson Male sternopleonal cavity, telson, gonopod 1 not reaching thoracic suture 2/3 (Fig. 14.184i)���������������������������������������������5 Maxilliped 3 palp when folded, reaching near to proximal margin of ischium (Fig. 14.184m)������������������Pseudopinnixa Maxilliped 3 palp when folded reaching ischium-merus margin at most (Fig. 14.184j, n)��������������������������������������������������6 Maxilliped 3 dactylus ovate, not reaching ischium-merus margin (Fig. 14.184j)������������������������������������������ Acmaeopleura Maxilliped 3 dactylus elongate, reaching ischium-merus margin (Fig. 14.184n)�����������������������������������������������Sestrostoma Acmaeopleura Stimpson, 1858

Diagnosis. Gaeticinae. Carapace anterolateral margin entire. Maxilliped 3 ischium-merus suture transverse; dactylus elongate, reaching ischium-merus margin. Maximum cl. 10 mm. Intertidal–shelf (0–37 m; in burrows of upogebiid shrimps and echiuroids). Temperate Northern W Pacific, Central Indo-Pacific. 2 species (Naruse 2015).

Brankocleistostoma Števčić, 2011 Diagnosis. Gaeticinae. Carapace elliptical, distinctly wider than long; anterolateral margin undulate, forming angle with posterolateral margin. Maxilliped 3 ischium-merus suture oblique. Maximum cl. 3.3 mm. Intertidal. Western Indo-Pacific (Mozambique only). 1 species (Barnard 1955: description of only species as Paracleistostoma fossula Barnard, 1955; Števčić 2011).

Gaetice Gistel, 1848 Diagnosis. Gaeticinae. Carapace quadrate, slightly wider than long; anterolateral margin with 2 teeth behind exorbital tooth. Maxilliped 3 ischium-merus suture oblique; dactylus elongate, reaching ischium-merus margin when folded. Maximum cl. 21 mm. Estuarine, intertidal. Temperate Northern W Pacific, Central Indo-Pacific. 2 species (Davie and Ng 2007: rediagnosis, key to species).

Subtidal (1–8 m; in burrows). Central Indo-Pacific (Ryukyus, Japan only). 1 species (Naruse 2015).

Pseudopinnixa Ortmann, 1894 Diagnosis. Gaeticinae. Carapace anterolateral margin entire. Maxilliped 3 ischium-merus suture transverse; palp reaching near to proximal margin of ischium when folded. Maximum cl. 13 mm. Intertidal (burrowing). Temperate Northern Pacific (Japan). 1 species (Komai and Konishi 2012).

Sestrostoma Davie & Ng, 2007 Diagnosis. Gaeticinae. Carapace anterolateral margin entire. Maxilliped 3 ischium-merus suture transverse; dactylus ovate, not reaching ischium-merus margin. Maximum cl. 14 mm. Intertidal (in burrows of upogebiid shrimps and echiuroids). Temperate Northern Pacific, Central Indo-Pacific. 3 species (Davie and Ng 2007; Marin et  al. 2011: redescription of species; Naruse 2015).

Subfamily Pinnotherelinae Alcock, 1900 Despite being free-living and its varunid, especially Cyclograpsus-like, features, Pinnotherelia has long been

Gopkittisak Naruse & Clark, 2009 Diagnosis. Gaeticinae. Carapace elliptical, distinctly wider than long; anterolateral margin straight, merging with posterolateral margin. Maxilliped 3 ischium-merus suture oblique. Maximum cl. 5 mm. Intertidal, subtidal (0–1 m; in burrows). Western and Central Indo-Pacific. 2 species (Komai 2011; Naruse and Clark 2009).

Proexotelson Naruse, 2015 Diagnosis. Gaeticinae. Carapace anterolateral margin entire. Male sternopleonal cavity, telson, gonopod 1 entering buccal cavity. Maxilliped 3 ischium-merus suture transverse. Maximum cl. 3.5 mm.

Fig. 14.185.  Varunidae, Pinnotherelinae, Pinnotherelia laevigata H. Milne Edwards & Lucas, 1843. a, habitus; b, carapace front, maxillipeds 3; c, male pleon.

830

Marine Decapod Crustacea

placed in Pinnotheroidea (Ng et  al. 2008). Palacios Theil et  al. (2016), however, showed Pinnotherelia to be a varunid. Diagnosis. Male pleonites 5, 6 fused. Maxillipeds 3 separated by narrow gape when closed; maxilliped 3 palp elongate, with long, dense setal brush, dactylus reaching ischium-merus articulation. Pinnotherelia H. Milne Edwards & Lucas, 1843 Maximum cl. 11 mm. Intertidal. Eastern Indo-Pacific (Marquesas), Tropical Eastern Pacific, Temperate South America (Chile, Peru). 1 species (Rathbun 1918: redescription, figures).

Subfamily Thalassograpsinae Davie & Ng, 2007 The subfamily recognises the unique form of the front, maxilliped 3, male pleon, walking legs, and thoracic sternite of a single species (Davie and Ng 2007). Thalassograpsus harpax (Hilgendorf, 1882) is widespread under corals and rocks on sheltered rocky shores in the Indo-West Pacific. Diagnosis. Lateral margins of carapace front with short oblique sulcus separating frontal margin from inner orbital angle (Fig. 14.186a). Male pleonites 5, 6 fused (Fig. 14.186b). Maxillipeds 3 mesial margins contiguous when closed; ischium-merus without longitudinal sulcus. Ambulatory legs propodi with distal spine on flexor margin (Fig. 14.186d).

Fig. 14.186.  Varunidae, Thalassograpsinae. Thalassograpsus harpax (Hilgendorf, 1882). a, carapace; b, male pleon, telson; c, maxilliped 3; d, Pereopod 4.

Thalassograpsus Tweedie, 1950 Maximum cl. 11.3 mm. Intertidal. Western and Central IndoPacific. 1 species (Davie and Ng 2007).

Subfamily Varuninae H. Milne Edwards, 1853 Varunine crabs lack the specialised pterygostome reticulation or maxillipeds 3 seen in the other subfamilies (Davie and Ng 2007). Most inhabit intertidal mudflats shores where they shelter under rocks or burrow into the sediment. Many occur around river mouths or among mangroves. Few reach fresh water streams. Diagnosis. Male pleonites 1–6 movable (Fig.  14.188u). Maxillipeds 3 moderately gaping when closed (Fig. 14.188t).

Key to genera of Varuninae 1. – 2. – 3. – 4. – 5. – 6. –

Carapace subhexagonal, with prominent rounded tooth between postero- and anterolateral margins (Fig. 14.188i)��������������������������������������������������������������������������������������������������������������������������������������������������������������Otognathon Carapace quadrate, subrectangular, trapezoid or with convex lateral margins���������������������������������������������������������������������2 Carapace anterolateral margin with 1 or 2 small lobes or teeth behind exorbital lobe���������������������������������������������������������3 Carapace anterolateral margin with 2 or more teeth behind exorbital tooth�������������������������������������������������������������������������6 Carapace about as long as wide������������������������������������������������������������������������������������������������������������������������������������������������������4 Carapace wider than long���������������������������������������������������������������������������������������������������������������������������������������������������������������5 Maxilliped 3 exopod as wide as ischium (Fig. 14.189h). Carapace anterolateral margin with 1 or 2 indistinct lobes or small teeth behind exorbital tooth (Fig. 14.188n)���������������������������������������������������������Ptychognathus Maxilliped 3 exopod narrower than ischium (Fig. 14.189i). Carapace anterolateral margin with 2 indistinct lobes behind exorbital tooth (Fig. 14.188p)����������������������������������������������������������������������������������������������������������� Scutumara Carapace ~1.4 times as wide as long, widest anteriorly; anterolateral margin with wide exorbital lobe plus 1 anterolateral lobe (Fig. 14.188h)���������������������������������������������������������������������������������������������������������������������������������� Orcovita Carapace ~1.2 times as wide as long, laterally evenly convex; anterolateral margin with 2 lobes behind exorbital tooth (Fig. 14.188o)�����������������������������������������������������������������������������������������������������������������������������Pyxidognathus Carapace anterolateral margin with 3 or more teeth behind exorbital tooth�������������������������������������������������������������������������7 Carapace anterolateral margin with 2 teeth behind exorbital tooth��������������������������������������������������������������������������������������11

14 – Brachyura – crabs

831

7.

Carapace anterolateral margin with 3 sharp teeth behind exorbital tooth, last largest and widest (Fig. 14.188c)�����������������������������������������������������������������������������������������������������������������������������������������������������������Cyrtograpsus – Carapace anterolateral margin with 2 or 3 teeth behind exorbital tooth, last tooth smallest����������������������������������������������8 8. Carapace widest at midlength��������������������������������������������������������������������������������������������������������������������������������������������������������9 – Carapace widest posteriorly����������������������������������������������������������������������������������������������������������������������������������������������������������10 9. Carapace anterolateral margin with 3 sharp teeth behind sharp exorbital tooth; frontal margin with 4 lobes or teeth (Figs 14.187, 14.188d). Adult male chela densely covered with plumose setae almost completely obscuring all surfaces���������������������������������������������������������������������������������������������������������������������������������������������������� Eriocheir – Carapace anterolateral margin with 3 small teeth behind blunt exorbital tooth; frontal margin weakly bilobed (Fig. 14.188b). Adult male chela with setal mat partially covering surfaces of fingers (Fig. 14.189l)������������������������������������������������������������������������������������������������������������������������������ ‘Brachynotus’ spinosus 10. Cheliped carpus and chela inner surface densely setose (Fig. 14.189n)�������������������������������������������������������������� Neoeriocheir – Cheliped palm outer surface densely setose (Fig. 14.189o)��������������������������������������������������������������������������������� Platyeriocheir 11. Carapace surface granulated������������������������������������������������������������������������������������������������������������������������������������������������������� 12 – Carapace surface glabrous or setose��������������������������������������������������������������������������������������������������������������������������������������������13 12. Carapace with granulated ridges demarcating regions (Fig. 14.188g)�������������������������������������������������������������� Noarograpsus – Carapace with wide V-shaped ridge on gastric region, horizontal setose granulated ridge on branchial region (Fig. 14.188r)������������������������������������������������������������������������������������������������������������������������������������������������������������ Utica 13. Carapace anterolateral teeth broad, blunt (Fig. 14.188m). Maxilliped 3 merus expanded laterally; exopod half as wide as ischium, with sharp distomesial tooth (Fig. 14.189g)������������������������������������������� Pseudograpsus – Carapace anterolateral and exorbital teeth acute. Maxilliped 3 merus not expanded or expanded distolaterally; exopod narrower or wider than ischium, without distomesial tooth (with lobe in Varuna)���������������� 14 14. Maxilliped 3 exopod almost as wide or wider than ischium (Fig. 14.189d, k)����������������������������������������������������������������������15 – Maxilliped 3 exopod markedly narrower than ischium (Fig. 14.189a, f, i)���������������������������������������������������������������������������16 15. Carapace exorbital and anterolateral teeth separated by V-shaped notch, lateral margin weakly convex (Fig. 14.188j). Maxilliped 3 merus larger than narrow ischium, with strong distolateral lobe; exopod much wider than ischium (Fig. 14.189d)�����������������������������������������������������Parapyxidognathus – Carapace exorbital and anterolateral teeth broad, adjacent, lateral margin strongly convex (Fig. 14.188s). Maxilliped 3 merus with prominent distolateral lobe; exopod almost as wide as ischium, with distomesial lobe (Fig. 14.189k)������������������������������������������������������������������������������������������������������������� Varuna 16. Carapace exorbital and first anterolateral teeth lobate, second anterolateral tooth sharp, all well spaced. Cheliped with pulvinus (Fig. 14.188a)�������������������������������������������������������������������������������������������Brachynotus – Carapace exorbital and anterolateral teeth similar, second anterolateral tooth less prominent, all separated by narrow V-shaped gap. Cheliped without pulvinus����������������������������������������������������������������������������������17 17. Carapace exorbital and 2 anterolateral teeth diminishing in size, aligned (Fig. 14.188q). Maxilliped 3 merus wider distally, longer than ischium (Fig. 14.189j)���������������������������������������������������������� Tetragrapsus – Carapace anterolateral teeth set lateral to exorbital tooth, arranged in distinct arc (Fig. 14.188e, l). Maxilliped 3 merus about as long as ischium (Fig. 14.189a, f)������������������������������������������������������������������������������������������18 18. Maxilliped 3 palp reaching distal end of ischium when folded (Fig. 14.189f). Male cheliped dactylus with broad proximal tooth (Fig. 14.189m)�������������������������������������������������������������� Pseudogaetice – Maxilliped 3 palp not reaching distal end of ischium when folded (Fig. 14.189a). Male cheliped dactylus without tooth����������������������������������������������������������������������������������������������������������������Hemigrapsus Brachynotus De Haan, 1833 Diagnosis. Varuninae. Carapace anterolateral margin with exorbital and first anterolateral teeth lobate, second anterolateral tooth sharp, all well spaced. Maxilliped 3 merus expanded laterally. Maximum cl. 15 mm (Pl. 71a).

Estuarine, intertidal, subtidal (0–21 m). Temperate Northern Atlantic. 4 species. The genus includes three species in the Mediterranean, one in NE Atlantic (Schubart et  al. 2001). ‘Brachynotus’ spinosus (H. Milne Edwards, 1853), a common species in SE A ­ ustralia (Figs 14.188b, 14.189l), is very different

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Marine Decapod Crustacea

from the northern species (Poore 2004) and deserves a separate genus closer to Eriocheir. It appears alone in the key above.

Cyrtograpsus Dana, 1851 Diagnosis. Varuninae. Carapace subcircular, slightly wider than long; anterolateral margin 3 sharp teeth behind exorbital tooth, third largest and widest. Maximum cl. 42 mm. Estuarine, intertidal, subtidal, lagoons. Tropical Atlantic, Temperate South America (Pacific and Atlantic coasts). 4 species (Lezcano et al. 2012: discussion of all species).

Eriocheir De Haan, 1835 mitten crabs Diagnosis. Varuninae. Carapace wider than long, widest at midlength; frontal margin with 4 teeth or lobes; anterolateral margin with 3 teeth behind exorbital tooth. Cheliped fingers with tufts of long setae (Fig. 14.187). Maxilliped 3 merus about as wide distally as long; exopod narrower than ischium. Maximum cl. 72 mm.

Fig. 14.187.  Varunidae, Varuninae. Eriocheir japonica (De Haan, 1835)

Freshwater, estuarine, intertidal, subtidal. Temperate Northern W Pacific (native range), Temperate Northern Atlantic, Western and Central Indo-Pacific (introduced to Europe, UK, NE and NW America, Persian Gulf). 4 species. Mitten crabs are a valuable

Fig. 14.188.  Varunidae, Varuninae. Carapace: a, Brachynotus (with left cheliped); b, ‘Brachynotus’ spinosa (H. Milne Edwards, 1853); c, Cyrtograpsus; d, Eriocheir; e, Hemigrapsus; f, Neoeriocheir; g, Noarograpsus; h, Orcovita; i, Otognathon; j, Parapyxidognathus; k, Platyeriocheir; l, Pseudogaetice; m, Pseudograpsus; n, Ptychognathus; o, Pyxidognathus; p, Scutumara; q, Tetragrapsus; r, Utica; s, Varuna. Buccal cavity, maxillipeds 3: t, Eriocheir. Male pleon, telson: u, Parapyxidognathus.

14 – Brachyura – crabs

food resource in China and SE Asia where they are cultured in hatcheries and rice fields (Cheng et  al. 2008; Naser et  al. 2012; Wang et  al. 2018), They are invasive pests elsewhere (Panning 1939), ‘arguably the most notorious brachyuran species on the planet’ (Naser et al. 2012). Naser et al. (2012) reviewed this conflict, the history of invasion and provided diagnostic characters for three species. They discussed divergent opinions on the number of species resulting from molecular analyses (Chu et al. 2003; Tang et al. 2003) and provided new diagnostic characters. Sakai (2013) used comparative morphology to recognise four species in two genera, a conclusion not followed here.

833

Neoeriocheir Sakai, 1983 Diagnosis. Varuninae. Carapace wider than long, widest posteriorly; anterolateral margin with 3 teeth behind exorbital tooth. Cheliped carpus and chela inner surface densely setose in male. Maxilliped 3 merus longer than wide; exopod narrower than ischium. Maximum cl. 20 mm. Estuarine, intertidal. Central Indo-Pacific. 1 species (Ng et al. 1999: rediagnosis, species description; Sakai 2013: rediagnosis, species description).

Noarograpsus Ng, Manuel & Ng, 2006 Hemigrapsus Dana, 1851 Diagnosis. Varuninae. Carapace anterolateral margin with 2 anterolateral teeth set lateral to exorbital tooth, arranged in distinct arc. Maxilliped 3 merus about as wide distally as long, about as long as ischium; exopod narrower than ischium. Maximum cl. 47 mm (Pl. 71d). Intertidal (mudflats, rocky reefs). Tropical Atlantic, Temperate Northern W Pacific, Central Indo-Pacific, Tropical Eastern Pacific, Temperate South America, Temperate Australasia. 14 species (Asakura and Watanabe 2005: new cryptic species; Dai and Yang 1991: key to 4 species). Two species have been introduced to Europe and the USA (Breton et  al. 2002; O’Connor 2001; Soors et al. 2010).

Diagnosis. Varuninae. Carapace about as wide as long, widest at first anterolateral tooth; with granulated ridges demarking regions; anterolateral margin with 2 teeth behind exorbital tooth. Maxilliped 3 merus expanded laterally; exopod narrower than ischium. Maximum cl. 8 mm. Intertidal. Central Indo-Pacific (Philippines). 1 species (Ng et al. 2006).

Otognathon Ng & Števčić, 1993 Diagnosis. Varuninae. Carapace subhexagonal, with prominent rounded tooth between postero- and anterolateral margins.

Fig. 14.189.  Varunidae, Varuninae. Maxilliped 3: a, Hemigrapsus; b, Neoeriocheir; c, Orcovita; d, Parapyxidognathus; e, Platyeriocheir; f, Pseudogaetice; g, Pseudograpsus; h, Ptychognathus; i, Scutumara; j, Tetragrapsus; k, Varuna. Chela, outer face: l, ‘Brachynotus’ spinosa; m, Pseudogaetice. Right cheliped, upper view: n, Neoeriocheir; o, Platyeriocheir.

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Maxilliped 3 merus broader than long, with prominent distolateral lobe; exopod half as wide as ischium. Maximum cl. 4 mm. Intertidal. Temperate Northern W Pacific, Central IndoPacific. 2 species (Ng et al. 2009: rediagnosis, second species; Ng and Števčić 1993: systematic position).

laterally; exopod half as wide as ischium, with sharp distomesial tooth. Maximum cl. 40 mm. Freshwater, estuarine, intertidal (migrating in streams to 800 m asl.) Indo-West Pacific. 6 species (Ng et al. 2002: taxonomy, ecology; Ng and Nakasone 1993: notes on a diagnosis).

Orcovita Ng & Tomascik, 1994

Ptychognathus Stimpson, 1858

Diagnosis. Varuninae. Carapace ~1.4 times as wide as long, widest anteriorly; anterolateral margin with wide exorbital lobe plus 1 anterolateral lobe. Maxilliped 3 merus broader than long, with prominent distolateral lobe; exopod as wide or wider than ischium. Maximum cl. 15 mm. Marine anchialine caves. Western and Central Indo-Pacific. 11 species (Davie and Ng 2012: key to 10 species; Ng and Tomascik 1994).

Diagnosis. Varuninae. Carapace as wide as long, with convex lateral margins. Anterolateral margin with 1 or 2 indistinct lobes or small teeth behind exorbital tooth. Maxilliped 3 merus broader than long, with prominent distolateral lobe; exopod as wide or wider than ischium. Maximum cl. 34 mm. Freshwater, estuarine, intertidal. Temperate Northern Pacific, Indo-West Pacific. 28 species. Most species of this large varunid genus have not been reviewed since Tesch’s (1918) key to 15 species. See Hsu et al. (2022b) and references therein for recent species descriptions.

Parapyxidognathus Ward, 1941 Diagnosis. Varuninae. Carapace quadrilateral, wider than long; anterolateral margin with 1 blunt, 1 sharp teeth behind sharp exorbital tooth. Maxilliped 3 merus larger than narrow ischium, with strong distolateral lobe; exopod much wider than ischium. Maximum cl. 12 mm. Freshwater, estuarine, intertidal (mudflats). Western and Central Indo-Pacific. 1 species. The type species, P. deianira (de Man, 1888), is from mangrove swamps in Myanmar and has been recorded in Indonesia and the Philippines (Ward 1941); Ng (2006) included a second species from rivers in India.

Platyeriocheir Ng, Guo & Ng, 1999 Diagnosis. Varuninae. Carapace wider than long, widest posteriorly; anterolateral margin with 2 teeth behind exorbital tooth. Cheliped palm outer surface densely setose in male. Maxilliped 3 merus as wide as long; exopod narrower than ischium. Maximum cl. 60 mm. Estuarine. Central Indo-Pacific (Taiwan). 1 species (Ng et al. 1999: diagnosis, species description).

Pseudogaetice Davie & Ng, 2007 Diagnosis. Varuninae. Carapace anterolateral margin with 2 teeth behind exorbital tooth, set lateral to exorbital tooth, arranged in distinct arc. Cheliped dactylus of male with broad proximal tooth. Maxilliped 3 merus about as long as ischium; palp reaching distal end of ischium when folded; exopod narrower than ischium. Maximum cl. 17 mm. Estuarine. Tropical Eastern Pacific (Mexico). 1 species (Davie and Ng 2007).

Pseudograpsus H. Milne Edwards, 1837 Diagnosis. Varuninae. Carapace as wide as long, with convex lateral margins; setose; anterolateral margin with 2 blunt teeth behind blunt exorbital tooth. Maxilliped 3 merus expanded

Pyxidognathus A. Milne-Edwards, 1879 Diagnosis. Varuninae. Carapace ~1.2 times as wide as long, with convex lateral margins; anterolateral margin with 2 lobes behind exorbital tooth. Maximum cl. 17 mm. Freshwater, estuarine. Western and Central Indo-Pacific. 3 species (Tesch 1918: key to 4 species, all described, one since synonymised).

Scutumara Ng & Nakasone, 1993 Diagnosis. Varuninae. Carapace as wide as long or slightly wider, with convex lateral margins, widest anteriorly; anterolateral margin with 2 lobes behind exorbital tooth. Maxilliped 3 merus broader than long, with prominent distolateral lobe; exopod narrower than ischium. Maximum cl. 8 mm. Intertidal (coral beaches). Western and Central Indo-Pacific. 3 species (Ng and Nakasone 1993: 3 species compared).

Tetragrapsus Rathbun, 1916 Diagnosis. Varuninae. Carapace trapezoid; anterolateral margin with exorbital and 2 anterolateral teeth diminishing in size, aligned. Maxilliped 3 merus wider distally, longer than ischium; exopod narrower than ischium. Maximum cl. 10 mm. Intertidal (burrows in saltmarsh). Tropical Eastern Pacific, Temperate South America. 1 species (Campos and Campos 1998: description, figure).

Utica White, 1847 Diagnosis. Varuninae. Carapace about as wide as long, widest at second anterolateral tooth; with wide V-shaped ridge on gastric region, horizontal setose granulated ridge on branchial region; cnterolateral margin with 2 teeth behind exorbital tooth. Maxilliped 3 merus wider than long; exopod half as wide as ischium. Maximum cl. 26 mm.

14 – Brachyura – crabs

Freshwater, estuarine, intertidal. Central Indo-Pacific, Temperate SE Australia. 7 species (Tesch 1918: key to 5 species). See Naruse et al. (2004) for recent species description.

Varuna H. Milne Edwards, 1853 river swimming crab Diagnosis. Varuninae. Carapace anterolateral margin with exorbital and anterolateral teeth broad, sharp, adjacent. Maxilliped 3 merus broader than long, with prominent distolateral lobe; exopod almost as wide as ischium, with distomesial lobe. Maximum cl. 51 mm (Pl. 71g). Freshwater, estuarine, intertidal, subtidal. Indo-West Pacific, Temperate Southern Africa, Temperate Australasia. 2 species (Hwang and Takeda 1986: second species; Mos et al. 2017: range extension into SE Australia; Naruse et  al. 2004: description of more common V. litterata (Fabricius, 1798)). References Asakura A, Watanabe S (2005) Hemigrapsus takanoi, new species, a sibling species of the common Japanese intertidal crab H. penicillatus (Decapoda: Brachyura: Grapsoidea). Journal of Crustacean Biology 25, 279–292. doi:10.1651/C-2514 Barnard KH (1955) Additions to the fauna-list of South African Crustacea and Pycnogonida. Annals of the South African Museum 43, 1–107. Breton G, Faasse M, Noel P, Vincent T (2002) A new alien crab in Europe: Hemigrapsus sanguineus (Decapoda: Brachyura: Grapsidae). Journal of Crustacean Biology 22, 184–189. doi:10.1163/20021975-99990221 Campbell BM, Griffin DJG (1966) The Australian Sesarminae (Crustacea: Brachyura): genera Helice, Helograpsus nov., Cyclograpsus, and Paragrapsus. Memoirs of the Queensland Museum 14, 127–174, pls 20–23. Campos E, Campos ARd (1998) Distribution and taxonomic remarks for five crab species of the family Grapsidae (Crustacea: Sesarminae and Varuninae) of the Mexican Pacific. Bulletin of the Southern California Academy of Sciences 97, 96–103. Cheng Y, Wu X, Yang X, Hines AH (2008) Current trends in hatchery techniques and stock enhancement for Chinese Mitten Crab, Eriocheir japonica sinensis. Reviews in Fisheries Science 16, 377–384. doi:10.1080/​ 10641260701681698 Chu KH, Ho HY, Li CP, Chan T-Y (2003) Molecular phylogenetics of the mitten crab species in Eriocheir, sensu lato (Brachyura: Grapsidae). Journal of Crustacean Biology 23, 738–746. doi:10.1651/C-2347 Dai A-Y, Yang S-L (1991) Crabs of the China Seas. English edn. China Ocean Press and Springer-Verlag, Beijing and Berlin. Davie PJF (2002) Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells A, Houston WWK, Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing, Melbourne. Davie PJF, Ng PKL (2007) Two new subfamilies of Varunidae (Crustacea: Brachyura), with descriptions of two new genera. Raffles Bulletin of Zoology Supplement 16, 257–272. Davie PJF, Ng PKL (2012) Two new species of Orcovita (Crustacea: Decapoda: Brachyura: Varunidae) from anchialine caves on Christmas Island, eastern Indian Ocean. Raffles Bulletin of Zoology 60, 57–70. Davie PJF, Nguyen VX (2003) A new species of Metaplax (Crustacea: Brachyura: Varunidae: Cyclograpsinae) from Vietnam. Raffles Bulletin of Zoology 51, 379–386.

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Davie PJF, Guinot D, Ng PKL (2015a) Chapter 71–16. Phylogeny of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 921–979. Brill, Leiden. Davie PJF, Guinot D, Ng PKL (2015b) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. Depledge MH (1989) Observations on the feeding behaviour of Gaetice depressus (Grapsidae: Varuninae) with special reference to suspension feeding. Marine Biology 100, 253–259. doi:10.1007/BF00391966 Hangai R, Kitaura J, Wada K, Fukui Y (2009) A new species of Cyclograpsus (Brachyura: Varunidae) from Japan, co-occurring with C. intermedius Ortmann, 1894. Crustacean Research 38, 21–27. doi:10.18353/ crustacea.38.0_21 Hsu J-W, Shih H-T, Innocenti G (2022a) A review of the mud crab genus Pseudohelice Sakai, Türkay & Yang, 2006 (Crustacea: Brachyura: Varunidae), with redescription of Cyclograpsus latreillii H. Milne Edwards, 1837, from the western Indian Ocean. Raffles Bulletin of Zoology 70, 94–107. doi:10.26107/RBZ-2022-0007 Hsu J-W, Shih H-T, Li J-J (2022b) Description of a new species of brackishwater crab of the genus Ptychognathus Stimpson, 1858 (Crustacea: Brachyura: Varunidae) from southern Taiwan. Nauplius 30, e2022002. doi:10.1590/2358-2936e2022002 Hwang JJ, Takeda M (1986) A new freshwater crab of the family Graspidae from Taiwan. Proceedings of the Japanese Society of Systematic Zoology 33, 11–18. Ituarte RB, D’Anatro A, Luppi TA, Ribeiro PD, Spivak ED, et al. (2012) Population structure of the SW Atlantic estuarine crab Neohelice granulata throughout its range: a genetic and morphometric study. Estuaries and Coasts 35, 1249–1260. doi:10.1007/s12237-012-9516-9 Komai T (2011) A new species of the varunid crab genus Gopkittisak (Crustacea: Decapoda: Brachyura: Grapsoidea) from the Ryukyu Islands. Species Diversity 16, 103–111. doi:10.12782/specdiv.16.103 Komai T, Konishi K (2012) Reappraisal of the systematic position of the supposed pinnotherid crab Pseudopinnixa carinata (Crustacea: Decapoda: Brachyura). Species Diversity 17, 29–37. doi:10.12782/ sd.17.1.029 Lezcano AH, González-José R, Spivak ED, Dellatorre FG (2012) Geographic differences in the carapace shape of the crab Cyrtograpsus affinis (Decapoda: Varunidae) and its taxonomic implications. Scientia Marina 76, 329–337. doi:10.3989/scimar.03391.16A Marin I, Korn OM, Kornienko ES (2011) Symbiotic crab Sestrostoma balssi (Shen, 1932) (Varunidae: Gaeticinae) from Vostok Bay, Sea of Japan: a new species for the fauna of Russia. Russian Journal of Marine Biology 37, 509–511. doi:10.1134/S1063074011060113 Marsden ID, Dewa RS (1994) Diel and tidal activity patterns of the smooth shore crab Cyclograpsus lavauxi (Milne Edwards 1853). Journal of the Royal Society of New Zealand 24, 429–438. doi:10.1080/03014223.1994.​ 9517478 McLay CL (1988) Brachyura and crab-like Anomura of New Zealand [Crabs of New Zealand]. Leigh Laboratory Bulletin 22, 1–463. Mos B, Ahyong ST, Burnes CN, Davie PJF, McCormack RB (2017) Range extension of a euryhaline crab, Varuna litterata (Fabricius, 1798) (Brachyura: Varunidae), in a climate change hot-spot. Journal of Crustacean Biology 37, 258–262. doi:10.1093/jcbiol/rux030 Naruse T (2015) Description of a new genus and a new species of gaeticine crab (Crustacea: Brachyura: Varunidae) from the Ryukyu

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Islands, and a review of Acmaeopleura Stimpson, 1858, and Sestrostoma Davie & N.K. Ng, 2007. Zootaxa 3925, 211–228. doi:10.11646/ zootaxa.​3925.2.4 Naruse T, Clark PF (2009) Establishment of a new genus for Asthenognathus gallardoi Serène & Soh, 1976 within Gaeticinae Davie & N. K. Ng, 2007 (Crustacea: Decapoda: Brachyura: Varunidae). Zootaxa 1987, 61–68. doi:10.11646/zootaxa.1987.1.3 Naruse T, Shih H-T, Ng NK, Hsu H-L (2004) On two new records of varunid crabs (Crustacea: Brachyura: Varunidae) from southern Taiwan. Collection and Research 18, 69–79. Naser MD, Page TJ, Ng NK, Apel M, Yasser AG, et  al. (2012) Invasive records of Eriocheir hepuensis Dai, 1991 (Crustacea: Brachyura: Grapsoidea: Varunidae): Implications and taxonomic considerations. BioInvasions Records 1, 71–86. doi:10.3391/bir.2012.1.1.15 Ng NK (2006) The systematics of the crabs of the family Varunidae (Brachyura, Decapoda). PhD thesis. National University of Singapore, Singapore. Ng PKL, Nakasone Y (1993) Scutumara enodis, a new genus and species of grapsid crab (Decapoda: Grapsidae) from Okinawa, Ryukyus, Japan. Crustacean Research 22, 1–6. doi:10.18353/crustacea.22.0_1 Ng PKL, Števčić Z (1993) The identity of Dentoxanthus komodoensis Serène, 1971 (Crustacea: Decapoda: Barchyura: Pilumnidae) with description of a new genus. Tropical Biodiversity 1, 101–106. Ng PKL, Tomascik T (1994) Orcovita saltatrix, a new genus and new species of anchialine varunine crab (Crustacea: Decapoda: Brachyura: Grapsidae) from Kakaban Island, Indonesia. Raffles Bulletin of Zoology 42, 937–948. Ng NK, Junyao G, Ng PKL (1999) Generic affinities of Eriocheir leptognathus and E. formosa with description of a new genus (Brachyura: Grapsidae: Varuninae). Journal of Crustacean Biology 19, 154–170. doi:10.2307/​1549557 Ng NK, Jeng M-S, Ng PKL (2002) On the taxonomy of Pseudograpsus setosus (Fabricius, 1798) (Decapoda, Brachyura, Grapsidae). Crustaceana 75, 759–775. doi:10.1163/156854002760289746 Ng NK, Manuel MR, Ng PKL, Boyko CB (2006) A new genus of varunid crab from the Philippines (Crustacea: Decapoda: Brachyura: Varunidae). Proceedings of the Biological Society of Washington 119, 49–57. doi:10.2988/0006-324X(2006)119[49:ANGOVC]2.0.CO;2 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Ng NK, Komai T, Ng PKL (2009) Otognathon uru, a new species of crabs (Brachyura, Varunidae) from coral reefs in Okinawa Island, The Ryukyus, Japan. Bulletin of the National Museum of Nature and Science. Series A, Zoology 3(Supplement), 205–214. Ng NK, Naruse T, Shih HT (2018) Helice epicure, a new species of varunid mud crab (Brachyura, Decapoda, Grapsoidea) from the Ryukyus, Japan. Zoological Studies 15, e15. doi:10.6620/ZS.​2018.57-15. O’Connor NJ (2001) The Asian shore crab Hemigrapsus sanguineus in New England: changes in resident crab populations. Proceedings of the Second International Conference on Marine Bioinvasions, New Orleans, La., April 9–11, 2001 105–106. Palacios Theil E, Cuesta JA, Felder DL (2016) Molecular evidence for nonmonophyly of the pinnotheroid crabs (Crustacea : Brachyura : Pinnotheroidea), warranting taxonomic reappraisal. Invertebrate Systematics 30, 1–27. doi:10.1071/IS15023 Panning A (1939) The Chinese mitten crab. Report of the Board of Regents of the Smithsonian Institution 3508, 361–375. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Rathbun MJ (1918) The grapsoid crabs of America. Bulletin of the United States National Museum 97, 1–461, pls 1–161.

Sakai K (2013) A review of the genus Eriocheir De Haan, 1835 and related genera, with the description of a new genus and a new species (Brachyura, Grapsoidea, Varunidae). Crustaceana 86, 1103–1138. doi:10.1163/​ 15685403-00003185 Sakai K, Yatsuzuka K (1980) Notes on some Japanese and Chinese Helice with Helice (Helicana) n. subgen., including Helice (Helicana) japonica n. sp. (Crustacea: Decapoda). Senckenbergiana Biologica 60, 393–411. Sakai K, Türkay M, Yang SL (2006) Revision of the Helice/Chasmagnathus-complex (Crustacea: Decapoda: Brachyura). Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 565, 1–76. Schubart CD, Cuesta JA, Diesel R, Felder D (2000) Molecular phylogeny, taxonomy, and evolution of nonmarine lineages within the American grapsoid crabs (Crustacea: Brachyura). Molecular Phylogenetics and Evolution 15, 179–190. doi:10.1006/mpev.1999.0754 Schubart CD, Cuesta JA, Rodríguez A (2001) Molecular phylogeny of the crab genus Brachynotus (Brachyura: Varuninae) based on the 16S rRNA gene. Hydrobiologia 449, 41–46. doi:10.1023/A:1017564229866 Schubart CD, Cannicci S, Vannini M, Fratini S (2006) Molecular phylogeny of grapsoid crabs (Decapoda, Brachyura) and allies based on two mitochondrial genes and a proposal for refraining from current superfamily classification. Journal of Zoological Systematics and Evolutionary Research 44, 193–199. doi:10.1111/j.1439-0469.2006.00354.x Soors J, Faasse M, Stevens M, Verbessem I, De Regge N, et al. (2010) New crustacean invaders in the Schelde estuary (Belgium). Belgian Journal of Zoology 140, 3–10. Stebbing TRR (1893) A History of Crustacea. Recent Malacostraca. Kegan Paul, Trench, Trübner and C., London. Števčić Z (2011) Addition to the reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Part I. New taxa. Natura Croatica 20, 125–139. Tang B, Zhou K, Song D, Yang G, Dai A (2003) Molecular systematics of the Asian mitten crabs, genus Eriocheir (Crustacea: Brachyura). Molecular Phylogenetics and Evolution 29, 309–316. doi:10.1016/ S1055-7903(03)00112-X Tesch JJ (1918) The Decapoda Brachyura of the Siboga Expedition. I Hymenosomatidae, Retropilumidae, Ocypodidae, Grapsidae and Gecarcinidae. Siboga-Expéditie 39c, 1–148, pls 1–6. Wahyudi AJ, Wada S, Aoki M, Hama T (2015) Gaetice depressus (Crustacea, Varunidae): Species profile and its role in organic carbon and nitrogen flow. Ocean Science Journal 50, 389–401. doi:10.1007/s12601-015-0036-y Wang J, Xu P, Zhou G, Li X, Lu Q, et al. (2018) Genetic improvement and breeding practices for Chinese mitten crab, Eriocheir sinensis. Journal of the World Aquaculture Society 49, 292–301. doi:10.1111/jwas.12500 Ward M (1941) New Brachyura from the Gulf of Davao, Mindanao, Philippine Islands. American Museum Novitates 1104, 1–15. Yang S-L, Tang B-P (2008) Description of the male of Asthenognathus hexagonum Rathbun, 1909 (Decapoda, Pinnotheridae). Crustaceana 81, 595–600. doi:10.1163/156854008784092201 Yin W, Fu C, Guo L, He Q, Li J, et al. (2009) Species delimitation and historical biogeography in the genus Helice (Brachyura: Varunidae) in the Northwestern Pacific. Zoological Science 26, 467–475. doi:10.2108/zsj.26.467

Xenograpsidae Ng, Davie, Schubart & Ng, 2007 Figure 14.190 All members of the small genus, Xenograpsus Takeda & Kurata, 1977, are confined to shallow hydrothermal vent environments where they can be common (Ng et al. 2014). Around shallow-water vents, Xenograpsus have been

14 – Brachyura – crabs

Fig. 14.190.  Xenograpsidae, Xenograpsus. a, carapace; b, front, orbit, antennule, antenna; c, maxilliped 3.

observed to feed on marine snow comprising zooplankton killed by the toxic vent plumes. The crabs themselves avoid the vent plumes by emerging from deep rock crevices only at slack water when the plumes are directed vertically (Jeng et al. 2004). A family separate from other grapsoids was justified for Xenograpsus on larval and adult morphology, and on DNA evidence (Ng et  al. 2007). The basal article of the antenna is immobile, locked laterally against the inner orbital tooth – a unique apomorphy among grapsoids. Maxillipeds 3 meet mesially without any gap, a condition not seen in other grapsoids. Diagnosis. Carapace subquadrate, smooth; front entire; orbit totally closed; suborbital crest with small granules; pterygostomial region sparingly setose, without pattern of reticulated setae, or covered in reticulated network of short, hooked, setae, with distinct groove subparallel to lateral margin of buccal cavity. Maxilliped 3 merus and ischium together subrectangular; maxillipeds 3 with mesial margins meeting when closed. Chelipeds robust, swollen, end of finger with subapical brush of setae; carpus unarmed, inner margin rounded. Male pleonites 1–6 and telson free. Xenograpsus Takeda & Kurata, 1977 vent crabs Maximum cl. 26 mm. Subtidal, shelf (5–270 m; hydrothermal vents). Temperate Northern Pacific, Central Indo-Pacific, Temperate ­Australasia (New Zealand). 3 species (Ng et  al. 2007: family, genus diagnosed, key to species; Ng et al. 2000: 3 species compared, ecology). References Jeng M-S, Ng NK, Ng PKL (2004) Hydrothermal vent crabs feast on sea ‘snow’. Nature 432, 969. doi:10.1038/432969a

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Ng PKL, Huang JF, Ho P-O (2000) Description of a new species of hydrothermal crab, Xenograpsus testudinatus (Crustacea: Decapoda: Brachyura: Grapsidae) from Taiwan. In Proceedings of the International Symposium on Marine Biology in Taiwan – Crustacea and zooplankton taxonomy, ecology and living resources, 26–27 May, 1998, Taiwan. (Eds Hwang J-S, Wang C-H, Chan T-Y) pp. 191–199. National Taiwan Museum, Taipei. Ng NK, Davie PJF, Schubart C, Ng PKL (2007) Xenograpsidae, a new family of grapsoid crabs (Crustacea: Brachyura) associated with shallow water hydrothermal vents. Raffles Bulletin of Zoology Supplement 16, 233–256. Ng NK, Suzuki H, Shih H-T, Dewa S-i, Ng PKL (2014) The hydrothermal crab, Xenograpsus testudinatus Ng, Huang & Ho, 2000 (Crustacea: Decapoda: Brachyura: Grapsidae) in southern Japan. Proceedings of the Biological Society of Washington 127, 391–399. doi:10.2988/​ 0006-324X-127.2.391

Ocypodoidea Rafineque, 1815 The superfamily Ocypodoidea comprises seven families of intertidal and shallow-water crabs from muddy shores and sandy beaches. The stalk-eyed crabs seen next to their burrows at low tide on intertidal mud-and sand flats (and which quickly hide when approached) are ocypodoids. The most familiar are variously known as fiddler crabs, sentinel crabs or semaphore crabs, others are semiterrestrial ghost crabs. Some are exceptionally common in tropical and subtropical environments (Sharifian et al. 2020), especially so because they are active out of the water when the tide is out. Many species build permanent burrows and are important bioengineers. Ocypodoidea and Grapsoidea are the two largest thoracotreme superfamilies and despite substantial taxonomic refinement in past decades, neither is monophyletic (Schubart et al. 2006; Tsang et al. 2018; Tsang et al. 2014). Such a conclusion might be inferred from the difficulty in clearly separating the two superfamilies. Although a core of graspoid families clearly differs from a core of ocypodoids, many are less easily assigned and will no doubt be reclassified when comprehensive phylogenetic data become available. The superfamily is currently characterised by the front being narrow relative to the anterior width, usually narrower than the orbit, and correspondingly long stalked eyes. Diagnosis. Thoracotremata. Carapace well calcified, hard in both sexes, usually squarish or transversely ovate; front narrow, usually narrower than or as wide as orbit (wider in Shenius). Maxilliped 3 ischium and merus free, or ischium and merus fused (rare, some Camptandriidae); very small rhomboidal gap between closed maxillipeds 3. Mandibles not visible when mouthparts closed. Pereopods 2–5 slender, with pointed dactyli.

838

Marine Decapod Crustacea

Key to families of Ocypodoidea 1. – 2. – 3. – 4. – 5. – 6. –

Orbit placed dorsally, forming longitudinal slits perpendicular to carapace front (Fig. 14.201)�����������Xenophthalmidae Orbit transverse or sloping posteriorly, placed along anterior carapace margin�������������������������������������������������������������������2 Carapace high, subglobular, longer than wide. Eyestalk as long as or shorter than cornea. Pleon similar in both sexes, broad, nearly covering most of thoracic sternum (Fig. 14.199)������������������������Mictyridae Carapace subglobular to subquadrate, usually wider than long. Eyestalk longer than cornea, sometimes markedly so. Pleon usually distinctly sexually dimorphic, male pleon never substantially covering thoracic sternum�������������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Gonopod 1 strongly bent, U-shaped (Fig. 14.194)������������������������������������������������������������������������������������������� Camptandriidae Gonopod 1 straight or sinuous, not strongly bent����������������������������������������������������������������������������������������������������������������������4 Male chelipeds unequal (Fig. 14.200)������������������������������������������������������������������������������������������������������������������������Ocypodidae Male chelipeds equal������������������������������������������������������������������������������������������������������������������������������������������������������������������������5 Carapace transversely subovate, regions weakly indicated; lateral margins unarmed behind exorbital angle. Cheliped dactylus curved, twisted, without truncate tooth on occlusal margin (Fig. 14.196)�����������Heloeciidae Carapace quadrate or polygonal, regions usually well indicated; lateral margins with teeth or lobes behind exorbital angle. Cheliped dactylus straight, not twisted, usually with large truncate tooth at or proximal to midlength of occlusal margin�����������������������������������������������������������������������������������������������������������������������������������������������6 Male pleon with constriction near pleonite 5 (pleonite 5 narrower than 6) or pleonite 4 with transverse setose brush partially covering pleonite 5. Carapace lateral margins usually without teeth or lobes behind exorbital angle. Merus of one or more walking legs usually with ovate typanum (Fig. 14.195)���������� Dotillidae Male pleon without constriction near pleonite 5, without transverse setose brush on pleonite 4. Carapace lateral margins with teeth or lobes behind exorbital angle. Merus of walking legs without typana (Figs 14.197, 14.198)�������������������������������������������������������������������������������������������������������� Macrophthalmidae

References Schubart CD, Cannicci S, Vannini M, Fratini S (2006) Molecular phylogeny of grapsoid crabs (Decapoda, Brachyura) and allies based on two mitochondrial genes and a proposal for refraining from current superfamily classification. Journal of Zoological Systematics and Evolutionary Research 44, 193–199. doi:10.1111/j.1439-0469.2006.00354.x Sharifian S, Kamrani E, Saeedi H (2020) Global biodiversity and biogeography of mangrove crabs: Temperature, the key driver of latitudinal gradients of species richness. Journal of Thermal Biology 92, 102692. doi:10.1016/j.jtherbio.2020.102692 Tsang LM, Schubart CD, Ahyong ST, Lai JCY, Au EYC, et al. (2014) Evolutionary history of true crabs (Crustacea: Decapoda: Brachyura) and the origin of freshwater crabs. Molecular Biology and Evolution 31, 1173–1187. doi:10.1093/molbev/msu068 Tsang LM, Ahyong ST, Shih H-T, Ng PKL (2018) Further polyphyly of pinnotheroid crabs: the molecular phylogenetic position of the polychaete-associated Aphanodactylidae. Invertebrate Systematics 32, 92–99. doi:10.1071/IS17038

Camptandriidae Stimpson, 1858 Figures 14.191–14.194, Plate 71h–l Camptandriid crabs inhabit intertidal and shallow muddy environments including in estuaries. The family is confined to the tropics of the Western and Central IndoPacific and extends around southern Africa into the eastern Atlantic. Camptandriids are surface feeders using

the spoon-shaped fingers (Fig. 14.193c) on their chelae to gather mud for sorting (Jones and Clayton 1983). They retreat to burrows at high tide, sometimes protecting these burrows by building towers or shelters over the opening (Kim et al. 2011). Camptandriids differ from other ocypodoids in possessing a U-shaped gonopod 1 (Fig.  14.194). The taxonomy of the group, then a subfamily of 12 genera, was reviewed by Manning and Holthuis (1981). Their key, later updated for 15 genera (Al-Khayat and Jones 1996), relied on features including the visibility of the gonopods when the pleon is closed (Fig. 14.193l, m) and the orbit (Fig. 14.192v–x). These can be difficult to interpret and are not included in this key to 24 genera. Diagnosis. Carapace transversely to longitudinally subovate or subquadrate, regions weakly indicated. Chelipeds usually subequal, chelae usually larger in males; dactylus of males usually with subproximal tooth; chelae of female weak, slender, with spatulate tips. Gonopod 1 strongly bent, U-shaped. Implicit generic attributes. Suborbital ridge not visible in dorsal view. Chelipeds sexually dimorphic, one cheliped of male larger. Pereopods 2–5 meri extensor margin without subdistal tooth. Male pleonite 1 about as wide as pleonite 2, not reaching coxae of pereopods 5.

14 – Brachyura – crabs

839

Key to genera of Camptandriidae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. – 9. – 10. –

Carapace ~1.3 times as long as wide (Figs 14.191a, 14.192l)�������������������������������������������������������������������������������������������������������2 Carapace about as long as wide or wider than long��������������������������������������������������������������������������������������������������������������������3 Carapace front one-half exorbital width, anterolateral margin with blunt teeth (Fig. 14.192l). Male pleonites 3–5 fused, with lateral concavity (Fig. 14.193n)������������������������������������������������������������������������������� Moguai Carapace front one-third exorbital width, anterolateral margin convex, without teeth (Fig. 14.191a). Male pleonites 2–5 fused, with angular shoulder (Fig. 14.193q)��������������������������������������������������������������������� Paramoguai Gonopod 1 with 2 long appendages (Fig. 14.194b, g, t)��������������������������������������������������������������������������������������������������������������4 Gonopod 1 without appendages or with short unequal appendages���������������������������������������������������������������������������������������6 Front not produced, orbits not visible dorsally, anterolateral and posterolateral margins equal, meeting at rounded angle (Fig. 14.192h). Male pleonites free������������������������������������������������������������������������������������������������ Exagorium Front produced, orbits visible dorsally, anterolateral and posterolateral margins equal, separated by tooth. Male pleonites 2–5 fused������������������������������������������������������������������������������������������������������������������������������������������������������������5 Carapace wider than long, anterolateral margin oblique, with low lobe (Fig. 14.192c). Gonopod 1 with 1 simple and 1 bifid appendage (Fig. 14.194b)���������������������������������������������������������������� Camptandrium Carapace little wider than long, anterolateral and posterolateral margins separated by tooth, with small tooth behind postorbital angle (Fig. 14.192t). Gonopod 1 with 2 simple appendages (Fig. 14.194t)����������������������������������������������������������������������������������������������������������������������������������������������������������������Takedellus Male pleonite 1 much wider than more posterior pleonites, reaching coxae of pereopods 5 (Fig. 14.193g). Suborbital ridge visible in dorsal view (Fig. 14.192d)�����������������������������������������������������������������������������������7 Male pleonite 1 about as wide as pleonite 2, not reaching coxae of pereopods 5 (Fig. 14.193h, i). Suborbital ridge usually not visible in dorsal view����������������������������������������������������������������������������������������������������������������8 Gonopod 1 with acute apex, not swollen. Chelipeds strongly sexually dimorphic������������������������������������������ Cleistostoma Gonopod 1 with blunt swollen apex (Fig. 14.193h). Chelipeds not sexually dimorphic���������������������������������������Ilyogynnis Pereopods 2–5 meri dilated, with teeth on flexor margin arranged in U-shape (Fig. 14.193d). Carapace anterolateral margin slightly shorter than posterolateral margin, with or without crenulate lobes (Fig. 14.192a)�������������������������������������������������������������������������������������������������������������������������� Baruna Pereopods 2–5 meri flexor margin smooth or with denticles at most�������������������������������������������������������������������������������������9 Male pleonite sutures all visible (articulating or 2–4 effectively immobile) (Fig. 14.193m, s)�������������������������������������������10 Male pleonites 2–4, 2–5 or 3–5 fused (Fig. 14.193j–l, n–p, r)���������������������������������������������������������������������������������������������������13 Carapace slightly wider than long, subhexagonal (Fig. 14.192g). Gonopod 1 with broad subapical lobe (Fig. 14.194f)��������������������������������������������������������������������������������������������������������������������������������������������������������������� Ecphantor Carapace transversely subovate, lateral margins convex. Gonopod 1 without broad subapical lobe�������������������������������11

Fig. 14.191.  Camptandriidae. a, Paramoguai kavieng Ahyong, 2014; b, Telmatothrix powelli Manning & Holthuis, 1981.

840

Marine Decapod Crustacea

11. Gonopod 1 with massive reflexed portion (Fig. 14.194l)������������������������������������������������������������������������������������Mortensenella – Gonopod 1 tapering to obliquely truncate apex����������������������������������������������������������������������������������������������������������������������� 12 12. Carapace anterolateral margin convex, with few obsolete lobes; posterolateral margin distinct; posterior margin straight (Fig. 14.192k). Chelipeds sexually dimorphic����������������������������������������������������������Manningis – Carapace anterolateral margin oblique, continuous with posterolateral margin, unarmed (Fig. 14.192p). Chelipeds not sexually dimorphic������������������������������������������������������������������������������������������������������� Opusia 13. Carapace anterolateral and posterolateral margins more or less continuous (Fig. 14.192e, f, i, o, q, r, s)������������������������14 – Carapace anterolateral and posterolateral margins separated by angle or tooth (Figs 14.191b, 14.192b, j, n, u)�������������������������������������������������������������������������������������������������������������������������������������������������� 20

Fig. 14.192.  Camptandriidae. Carapace: a, Baruna; b, Calabarium; c, Camptandrium; d, Cleistostoma; e, Danielella; f, Deiratonotus; g, Ecphantor; h, Exagorium; i, Leptochryseus; j, Lillyanella; k, Manningis; l, Moguai; m, Mortensenella; n, Nanusia; o, Nasima; p, Opusia; q, Paracleistostoma; r, Paratylodiplax; s, Serenella; t, Takedellus; u, Tylodiplax. Carapace front, orbit, epistome: v, Calabarium; w, Camptandrium; x, Nasima.

14 – Brachyura – crabs

841

Fig. 14.193.  Camptandriidae. Male major, minor chelipeds: a, b, Lillyanella. Female chela: c, Paracleistostoma. Pereopod 4: d, Baruna; e, Calabarium; f, Lillyanella. Pleonites 1, 2, pereopod 5 coxae (numbered): g, Cleistostoma; h, Exagorium; i, Leptochryseus. Male pleon: j, Camptandrium; k, Danielella; l, Deiratonotus (with thoracic sternites 5–8); m, Ecphantor (with thoracic sternites 4–8); n, Moguai; o, Nasima; p, Paracleistostoma; q, Paramoguai; r, Takedellus; s, Telmatothrix.

14. – 15. – 16. – 17. – 18. – 19. –

Carapace ~1.5 times as wide posteriorly as exorbital width, anterolateral margins convex, posterolateral margins diverging significantly, smooth or with blunt tooth (Fig. 14.192r)��������������������Paratylodiplax Carapace widest anteriorly or at midpoint or scarcely wider posteriorly������������������������������������������������������������������������������15 Carapace anterolateral margins diverging slightly posteriorly, with small anterolateral tooth (Fig. 14.192e). Male pleonites 3–5 fused (Fig. 14.193k). Gonopod 1 truncate, with 2 distal appendages, one small (Fig. 14.194d)�����������������������������������������������������������������������������������������������������������������������Danielella Carapace lateral margins diverging slightly posteriorly, with small anterolateral tooth. Male pleonites 2–4 or 2–5 fused. Gonopod 1 acute or swollen������������������������������������������������������������������������������������������16 Gonopod 1 swollen distally, with 1 or more short distal appendages (Fig. 14.194p)���������������������������������Paracleistostoma Gonopod 1 truncate or acute, without distal appendages�������������������������������������������������������������������������������������������������������17 Carapace distinctly wider than long, transversely quadrate, with 3-part transverse hairy ridge (Fig. 14.192f)�����������������������������������������������������������������������������������������������������������������������������������������������������������Deiratonotus Carapace about as wide as long, dorsally smooth���������������������������������������������������������������������������������������������������������������������18 Carapace lateral margins almost parallel, straight, with obsolete anterolateral lobe (Fig. 14.192o). Chelipeds not sexually dimorphic. Gonopod 1 with recurved portion broadened to flat lobe halfway along (Fig. 14.194n)������������������������������������������������������������������������������������������������������������������������������������������Nasima Carapace lateral margins convex. Chelipeds sexually dimorphic. Gonopod 1 without broadened section��������������������19 Carapace widest at anterior one-third (Fig. 14.192i). Gonopod with oblique apical margin bearing toadstool-shaped spines (Fig. 14.194i). Male pleonites 2–5 fused����������������������������������������������������������������� Leptochryseus Carapace widest at midlength (Fig. 14.192s). Gonopod 1 with narrow acute apex (Fig. 14.194s). Male pleonites 2–4 fused�������������������������������������������������������������������������������������������������������������������������������������������������������Serenella

842

Marine Decapod Crustacea

Fig. 14.194.  Camptandriidae. Gonopod 1: a, Calabarium; b, Camptandrium; c, Cleistostoma; d, Danielella; e, Deiratonotus; f, Ecphantor; g, Exagorium; h, Ilyogynnis; i, Leptochryseus; j, Manningis; k, Moguai; l, Mortensenella; m, Nanusia; n, Nasima; o, Opusia; p, Paracleistostoma; q, Paramoguai; r, Paratylodiplax; s, Serenella; t, Takedellus; u, Telmatothrix; v, Tylodiplax.

20. Carapace anterolateral margin with few sharp teeth (Fig. 14.192b, j). Pereopods 2–4 meri extensor margin with subdistal tooth (Fig. 14.193e, f). Gonopod 1 tapering to slender apex��������������������������������������������������������������������21 – Carapace anterolateral margin without or with blunt tooth (Figs 14.191b, 14.192n, u). Pereopods 2–4 meri extensor margin without subdistal tooth. Gonopod 1 wider distally or tapering to bifid apex����������������� 22 21. Carapace with prominent front, medially excavate (Fig. 14.192b). Pereopods 2–5 meri flexor margin with 1 subdistal tooth (Fig. 14.193e)��������������������������������������������������������������������������������������������������������������������� Calabarium – Carapace with wide front, medially almost straight (Fig. 14.192j). Pereopods 2–5 meri extensor margins with 1 distal and 1 subdistal tooth (Fig. 14.193f)���������������������������������������������������������������������������������� Lillyanella 22. Carapace widest posteriorly, anterolateral margin short, oblique, with obtuse tooth separating diverging posterolateral margin (Fig. 14.192u). Gonopod 1 wider distally, with 2 distal lobes (Fig. 14.194v)��������������������������������������������������������������������������������������������������������������������������������������������������������������� Tylodiplax – Carapace anterolateral margin with 2 teeth, posterior one larger, at juncture of posterolateral margin������������������������ 23 23. Carapace wider than long, front one-third exorbital width (Fig. 14.191b). Gonopod 1 with bifid apex (Fig. 14.194u). Chelipeds strongly sexually dimorphic������������������������������������������������������������������������������������Telmatothrix – Carapace as wide as long, front one-half exorbital width (Fig. 14.192n). Gonopod with bulbous apex, with 1 bifid appendage (Fig. 14.194m). Chelipeds not sexually dimorphic����������������������������������������������������������Nanusia Baruna Stebbing, 1904 Diagnosis. Carapace about as long as wide, dorsal surface granular; front almost one-half exorbital width; anterolateral margin slightly shorter than posterolateral margin, with or without crenulate lobes. Pereopods 2–5 meri dilated, with teeth on flexor margin arranged in U-shape. Male pleonites free. Gonopod 1 distal portion with spines, subapical process. Maximum cl. 6 mm (Pl. 71h).

Estuarine, intertidal (on wood and shells, mangroves, rocks). Western and Central Indo-Pacific. 4 species (Harminto and Ng 1991: rediagnosis, 3 species).

Calabarium Manning & Holthuis, 1981 Diagnosis. Carapace as wide as long, subhexagonal, dorsal surface uneven; front one-third exorbital width, protruding, medially

14 – Brachyura – crabs

843

excavate; anterolateral margin with few sharp teeth. Chelipeds not sexually dimorphic. Pereopods 2–5 meri extensor margin with subdistal tooth. Male pleonites 2–4 fused. Gonopod 1 tapering to slender apex. Maximum cl. 7.8 mm. Estuarine (on aquatic lily, Crinum natans). Tropical E Atlantic. 1 species (Manning and Holthuis 1981).

and posterolateral margins diverging posteriorly, separated by obtuse angle. Pereopods 2–5 meri extensor margin without subdistal tooth. Male pleonites free. Gonopod 1 with broad subapical lobe. Maximum cl. 4.5 mm. Estuarine. Tropical E Atlantic. 1 species (Manning and Holthuis 1981).

Camptandrium Stimpson, 1858

Exagorium Naruse, Chung & Tangah, 2015

Diagnosis. Carapace wider than long, subhexagonal, dorsal surface with few interrupted crests, with pubescence extending on to pereopods; front one-third exorbital width; suborbital ridge weak; anterolateral and posterolateral margins, separated by rounded blunt too, obtuse lobe more anteriorly; posterolateral angles angular. Male pleonites 2–5 fused. Gonopod 1 with 1 blade-like appendage, 1 bifid appendage. Maximum cl. 10 mm (Pl. 71k). Intertidal, subtidal (muddy sediments). Western and Central Indo-Pacific. 1 species (Tan and Ng 1999: rediagnosis).

Diagnosis. Carapace wider than long, hexagonal, dorsal surface epigastric regions with pair of convexities; front not protruding; anterolateral and posterolateral margins equal, separated by rounded angle. Male pleonites free; pleonite 1 much wider than pleonite 2 and others, reaching coxae of pereopods 5. Gonopod 1 with 2 long appendages. Maximum cl. 7.6 mm. Estuarine, intertidal, subtidal (0–0.3 m; in burrows). Central Indo-Pacific. 1 species (Naruse et al. 2015).

Cleistostoma De Haan, 1833 Diagnosis. Carapace wider than long, transversely subovate, dorsal surface mostly smooth; front less than one-third exorbital width; suborbital ridge visible in dorsal view; anterolateral and posterolateral margins evenly convex. Male pleonites 2–4 fused; pleonite 1 much wider than pleonite 2 and others, reaching coxae of pereopods 5. Gonopod 1 acute, with toadstool-shaped, subterminal spines. Maximum cl. 19 mm. Intertidal (mudflats). Central Indo-Pacific. 2 species (Ng et al. 2009: figures, discussion; Shen 1932: description, figures).

Danielella Ng & Clark, 2012 Diagnosis. Carapace wider than long, dorsal surface mostly smooth; front one-third exorbital width; lateral margins diverging slightly posteriorly, with small anterolateral tooth. Male pleonites 3–5 fused. Gonopod 1 truncate, with 2 distal appendages, one small. Maximum cl. 8 mm. Estuarine, intertidal. Temperate Southern Africa. 1 species (Ng and Clark 2012: replacement name; Ng et al. 2010: diagnosed as Danielita).

Deiratonotus Manning & Holthuis, 1981 Diagnosis. Carapace distinctly wider than long, transversely quadrate, dorsal surface with 3-part hairy transverse ridge; front one-third exorbital width; anterolateral and posterolateral margins diverging posteriorly, separated by obtuse angle. Male pleonites 2–5 fused. Gonopod 1 regularly narrowing to acute tip, without subapical lobe. Maximum cl. 10.5 mm. Intertidal. Temperate Northern W Pacific, Central IndoPacific. 3 species (Kawane et al. 2005: figures, synonymy of species; Manning and Holthuis 1981).

Ilyogynnis Manning & Holthuis, 1981 Diagnosis. Carapace about as long as wide, dorsal surface regions marked, pubescent; front one-third exorbital width; suborbital ridge visible in dorsal view; anterolateral margin with 2 obsolete lobes. Male pleonites 2–5 fused; pleonite 1 much wider than pleonite 2 and others, reaching coxae of pereopods 5. Gonopod 1 with blunt swollen apex. Maximum cl. 6 mm (Pl. 71j). Estuarine. Central Indo-Pacific. 1 species (Manning and Holthuis 1981; Tweedie 1937: description, figures).

Leptochryseus Al Khayat & Jones, 1996 Diagnosis. Carapace distinctly wider than long, transversely quadrate, dorsal surface granular; front about one-quarter exorbital width; suborbital ridge visible in dorsal view; anterolateral margin with broad rounded lobe, posterolateral margins converging posteriorly. Male pleonites 2–5 fused. Gonopod 1 acute, with toadstool-shaped, subterminal spines. Maximum cl. 21 mm. Estuarine, intertidal. Central Indo-Pacific. 1 species (Ng et al. 2009: rediagnosis, discussed possible synonymy with Nasima).

Lillyanella Manning & Holthuis, 1981 Diagnosis. Carapace wider than long, subhexagonal, dorsal surface mostly smooth; front almost one-half exorbital width; anterolateral margin oblique, with 2 large and few small teeth, posterolateral margin converging posteriorly. Pereopods 2–5 meri extensor margin with 1 distal and 1 subdistal tooth. Male pleonites 2–4 fused. Gonopod 1 slightly swollen apically, with recurved spines. Maximum cl. 4.3 mm. Estuarine, intertidal. Tropical E Atlantic. 1 species (Manning and Holthuis 1981).

Ecphantor Manning & Holthuis, 1981

Manningis Al Khayat & Jones, 1996

Diagnosis. Carapace slightly wider than long, subhexagonal, dorsal surface uneven; front one-half exorbital width; anterolateral

Diagnosis. Carapace wider than long, transversely subovate, dorsal surface granular; front one-third exorbital width; anterolateral

844

Marine Decapod Crustacea

margin convex, with few obsolete lobes; posterolateral margin distinct; posterior margin straight. Male pleonites visible, 2–4 fused. Gonopod 1 abruptly recurved, recurved portion with strong distal spines. Maximum cl. 5.4 mm. Intertidal (muddy sediments). Western Indo-Pacific. 1 species (Al-Khayat and Jones 1996; Ng et al. 2009: rediagnosis).

anterolateral margin oblique, continuous with posterolateral margin, unarmed. Chelipeds not sexually dimorphic. Male pleonites free. Gonopod 1 gently bent, apex conical with numerous strong subapical spines. Maximum cl. 7 mm. Estuarine, intertidal. Western Indo-Pacific. 1 species (Ng et al. 2009).

Moguai Tan & Ng, 1999

Paracleistostoma De Man, 1895

Diagnosis. Carapace longer than wide, dorsal surface granular; front one-half exorbital width; anterolateral and posterolateral margins separated by tooth, with small tooth behind postorbital angle; posterolateral angles projecting, lobe-like. Chelipeds not sexually dimorphic. Male pleonites 3–5 fused. Gonopod 1 with 2 long appendages. Maximum cl. 7 mm. Intertidal, subtidal (muddy sediments, mangroves). Central Indo-Pacific. 2 species (Tan and Ng 1999).

Diagnosis. Carapace wider than long, dorsal surface mostly smooth; front one-third exorbital width; suborbital ridge visible in dorsal view; anterolateral margin convex, with or without obsolete lobes. Male pleonites 2–5 fused. Gonopod 1 swollen distally, with 1 or more short distal appendages. Maximum cl. 8 mm. Estuarine, intertidal, subtidal (mangroves, muddy sediments). Central Indo-Pacific, Temperate Australasia (Qld, Australia). 9 species (Manning and Holthuis 1981: rediagnosis; Rahayu and Ng 2003: 3 species described).

Mortensenella Rathbun, 1909 Diagnosis. Carapace wider than long, dorsal surface mostly smooth; front almost one-half exorbital width; anterolateral and posterolateral margins evenly convex. Chelipeds not sexually dimorphic. Male pleonites free. Gonopod 1 with massive reflexed portion. Maximum cl. 5 mm (Pl. 71i). Subtidal (in burrows of echiuroids and sipunculans). Central Indo-Pacific. 1 species (Dai and Yang 1991: figures, descriptions; Harminto and Ng 1991: transferred species to Camptandriinae; Rathbun 1909: described species as pinnotherid).

Paramoguai Ahyong, 2014 Diagnosis. Carapace longer than wide, vase-shaped, dorsal surface with few protuberances; front one-third exorbital width; anterolateral margin constricted behind postorbital cup, unarmed; posterolateral angles projecting, lobe-like. Chelipeds not sexually dimorphic. Male pleonites 2–5 fused. Gonopod 1 with 2 long appendages. Maximum cl. 5.4 mm. Estuarine, intertidal, subtidal (sand-mud sediments). Central Indo-Pacific (Papua New Guinea, Japan). 2 species (Ahyong 2014; Naruse 2005).

Nanusia Tan & Ng, 1999

Paratylodiplax Serène, 1974

Diagnosis. Carapace about as long as wide, dorsal surface uneven; front one-half exorbital width; suborbital ridge weak; anterolateral and posterolateral margins separated by tooth, with small tooth behind postorbital angle; posterolateral angles round. Chelipeds not sexually dimorphic. Male pleonites 2–5 fused. Gonopod 1 with bulbous apex, with 1 bifid appendage. Maximum cl. 5 mm. Estuarine, intertidal. Central Indo-Pacific. 1 species (Tan and Ng 1999).

Diagnosis. Carapace wider than long, dorsal surface mostly smooth or setose; front about one-quarter exorbital width; widest posteriorly, anterolateral margins convex, posterolateral margins diverging significantly, smooth or with blunt tooth. Chelipeds not sexually dimorphic. Male pleonites 3–5 fused. Gonopod 1 truncate, with 2 distal appendages, one small. Maximum cl. 7 mm. Intertidal, subtidal. Western Indo-Pacific, Temperate Southern Africa. 3 species (Guinot and Crosnier 1963: one species described as Tylodiplax derijardi Guinot & Crosnier, 1963; Ng et  al. 2009: figures).

Nasima Manning, 1991 Diagnosis. Carapace distinctly wider than long, transversely quadrate, dorsal surface convex, regions marked; front about one-quarter exorbital width; lateral margins almost parallel, straight, with obsolete anterolateral lobe. Chelipeds not sexually dimorphic. Male pleonites 2–5 fused. Gonopod 1 acute, recurved portion broadened to flat lobe halfway along. Maximum cl. 12.3 mm (Pl. 71l). Intertidal. Western Indo-Pacific. 1 species (Ng et  al. 2009: rediagnosis)

Diagnosis. Carapace about as long as wide, dorsal surface mostly smooth; front almost one-half exorbital width; lateral margin convex, widest posterior to midlength, unarmed. Male pleonites 2–4 fused. Gonopod 1 with acute apex. Maximum cl. 6.6 mm. Intertidal (mangroves). Western Indo-Pacific. 1 species (Manning and Holthuis 1981).

Opusia Ng, Rahayu & Naser, 2009

Takedellus Tan & Ng, 1999

Diagnosis. Carapace wider than long, transversely subovate, dorsal surface regions marked; front less than one-third exorbital width;

Diagnosis. Carapace slightly wider than long, subhexagonal, dorsal surface granular; front one-third exorbital width; anterolateral

Serenella Manning & Holthuis, 1981

14 – Brachyura – crabs

and posterolateral margins separated by tooth, with small tooth behind postorbital angle; posterolateral angles round. Male pleonites 2–5 fused. Gonopod 1 with 2 long appendages. Maximum cl. 4.8 mm. Estuarine, intertidal. Central Indo-Pacific. 1 species (Tan and Ng 1999).

Telmatothrix Manning & Holthuis, 1981 Diagnosis. Carapace wider than long, subhexagonal, dorsal surface regions marked; front one-third exorbital width; anterolateral and posterolateral margins separated by tooth, second more anterior tooth. Male pleonites 2–4 fused. Gonopod 1 with bifid apex. Maximum cl. 10 mm. Estuarine, intertidal. Tropical E Atlantic. 1 species (Manning and Holthuis 1981).

Tylodiplax De Man, 1895 Diagnosis. Carapace wider than long; front less than one-third exorbital width; anterolateral and posterolateral margins diverging posteriorly, separated by blunt tooth. Chelipeds not sexually dimorphic. Male pleonites 2–4 fused. Gonopod 1 widened distally, with 2 distal lobes. Maximum cl. 9 mm. Intertidal. Central Indo-Pacific. 1 species (Manning and Holthuis 1981: rediagnosis; Ng et al. 2009: figures). References Ahyong ST (2014) Paramoguai kavieng, a new genus and species of camptandriid crab from Papua New Guinea (Crustacea: Brachyura). Zootaxa 3856, 578–584. doi:10.11646/zootaxa.3856.4.6 Al-Khayat JA, Jones DA (1996) Two new genera, Manningis and Leptochryseus (Decapoda: Camptandriidae), and descriptions of the first zoea of six brachyurans from the Arabian Gulf. Journal of Crustacean Biology 16, 797–813. doi:10.2307/1549197 Dai A-Y, Yang S-L (1991) Crabs of the China Seas. English edn. China Ocean Press and Springer-Verlag, Beijing and Berlin. Guinot D, Crosnier A (1963) Remarques sure les genres Cleistostoma, Paracleistostoma et Tylodiplax, et description de Tylodiplax derijardi sp. nov. (Crust. Decap. Brachyura). Bulletin du Muséum National d’Histoire Naturelle, ser. 2 35, 606–619. Harminto S, Ng PKL (1991) A revision of the camptandriine genus Baruna Stebbing, 1904 (Crustacea: Decapoda: Brachyura: Ocypodidae), with descriptions of two new species from the Indo-West Pacific. Raffles Bulletin of Zoology 39, 187–207. Jones DA, Clayton D (1983) The systematics and ecology of crabs belonging to the general Cleistostoma De Haan and Paracleistostoma De Man on Kuwait Mudflats. Crustaceana 45, 183–199. doi:10.1163/​ 156854083X00613 Kawane M, Wada K, Kitaura J, Watanabe K (2005) Taxonomic re‐examination of the two camptandriid crab species Deiratonotus japonicus (Sakai, 1934) and D. tondensis Sakai, 1983, and genetic differentiation among their local populations. Journal of Natural History 39, 3903– 3918. doi:10.1080/00222930500445085 Kim TW, Ryu HJ, Choi JB, Choe JC (2011) Tower construction by the manicure crab Cleistostoma dilatatum during dry periods on an intertidal mudflat. Journal of Ethology 29, 459–465. doi:10.1007/s10164-011-0280-2 Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306

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Naruse T (2005) Species of Moguai Tan and Ng, 1999 (Decapoda: Brachyura: Camptandriidae) from brackish waters in the Ryukyu Islands, Japan, with the description of a new species. Zootaxa 1044, 57–64. doi:10.11646/zootaxa.1044.1.6 Naruse T, Chung AY, Tangah J (2015) Description of a new genus and a new species of the family Camptandriidae Stimpson, 1858 (Crustacea: Decapoda: Brachyura) from Lower Kinabatangan-Segama Wetlands, Sabah, Malaysia. Raffles Bulletin of Zoology 63, 327–333. Ng PKL, Clark PF (2012) Danielita Ng, Clark & Tan, 2010 (Crustacea: Decapoda: Brachyura: Ocypodoidea: Camptandriidae), preoccupied by Danielita Kiriakoff, 1970 (Lepidoptera: Glossata: Heteroneura: Noctuoidea: Notodontidae). Zootaxa 3490, 48. doi:10.11646/zootaxa.​ 3490.1.3 Ng PKL, Rahayu DL, Naser MD (2009) The Camptandriidae of Iraq, with description of a new genus and notes on Leptochryseus Al-Khayat & Jones, 1996 (Crustacea: Decapoda: Brachyura). Zootaxa 2312, 1–26. doi:10.11646/zootaxa.2312.1.1 Ng PKL, Clark PF, Tan CGS (2010) A new genus for Cleistostoma edwardsii Macleay, 1838 (Ocypodoidea, Camptandriidae) from South Africa. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 251–260. Rahayu DL, Ng PKL (2003) The Camptandriidae of Irian Jaya, Indonesia, including the descriptions of two new species (Decapoda: Brachyura). Journal of Crustacean Biology 23, 951–962. doi:10.1651/C-2381 Rathbun MJ (1909) New crabs from the Gulf of Siam. Proceedings of the Biological Society of Washington 22, 107–114. Shen CJ (1932) The brachyuran Crustacea of North China. Zoologia Sinica. Series A. Invertebrates of China 9, 1–320, pls 1–10. Tan CGS, Ng PKL (1999) A revision of the genus Camptandrium Stimpson, 1858 (Crustacea: Decapoda: Brachyura: Camptandriidae). Raffles Bulletin of Zoology 47, 193–219. Tweedie MWF (1937) On the crabs of the family Ocypodidae in the collection of the Raffles Museum. Bulletin of the Raffles Museum 13, 140–170.

Dotillidae Stimpson, 1858 sand bubbler crabs Figure 14.195, Plate 71m, n Sand bubbler crabs are small crabs of intertidal and shallow muddy environments in tropical and subtropical regions of the Eastern Pacific and Indian Ocean. Some species are very common (e.g. Ramesh et  al. 2019). All are burrowers and some build earthen barriers around their homes (Wada 1994). Their impact on sediment structure is well documented (Sassa and Yang 2019). Dotillids differ from other ocypodoids in being more globose and having the antennules folded lengthwise and parallel. The mouthfield is broad, covered by wide maxillipeds 3 that meet in the middle. Some genera have thinly chitinised tympana, possibly used for respiration, on the meri of pereopods 2–5. The foundations of the taxonomy of the family, then treated as subfamily Scopimerinae of the Ocypodidae, were established by Alcock (1900) but only half the genera were then recognised. One species, Camptandrium anomalum Shen, 1935 has had a convoluted taxonomic history – it was included in its own genus

846

Marine Decapod Crustacea

Shenius Serène, 1974 and is now included in its own subfamily, Sheniinae, of Dotillidae (Ng et al. 2010). Diagnosis. Carapace anterolateral margin with or without small tooth behind exorbital angle. Male chelipeds equal; dactylus straight, usually with large truncate tooth at or proximal to midlength of occlusal margin. Pereopods with merus of one or more walking legs usually with ovate

typanum. Male pleon with constriction near pleonite 5 (pleonite 5 narrower than 6) or pleonite 4 with transverse setose brush partially covering pleonite 5. Implicit generic attributes. Eyestalk not prolonged beyond cornea. Maxilliped 2 propodus not expanded, dactylus terminal. Male chelipeds subequal. Chelipeds palm and fingers stout, longer than pereopods 2–4 in male.

Key to genera of Dotillidae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8. – 9. –

Carapace with prominent postorbital tooth, 2 anterolateral teeth. Pereopods 2–5 meri flexor margins with teeth (Fig. 14.195i). Male telson trilobed (Fig. 14.195w)�������������������������������������������������������������������������������������������� Shenius Carapace with small postorbital tooth, with or without 1 anterolateral tooth. Pereopods 2–5 meri flexor margins smooth. Male telson rounded������������������������������������������������������������������������������������������������������������������������������������2 Eyestalk prolonged beyond cornea as styliform process (Fig. 14.195j)�����������������������������������������������������������Tmethypocoelis Eyestalk not prolonged beyond cornea�����������������������������������������������������������������������������������������������������������������������������������������3 Carapace subovate, lateral margins convex. Pereopods 2–5 meri without tympana (Fig. 14.195c)�������������� Potamocypoda Carapace subquadrate, lateral margins straight or angular. Pereopods 2–5 meri usually with tympana (Fig. 14.195d), sometimes absent (Fig. 14.195b) or ill-defined (Fig. 14.195a)���������������������������������������������������������������������4 Maxilliped 2 propodus greatly expanded, dactylus applied as narrow strip (Fig. 14.195l). Pereopods 2–5 meri with large tympana (Fig. 14.195d)��������������������������������������������������������������������������������������������������������5 Maxilliped 2 propodus not expanded, dactylus terminal (Fig. 14.195k). Pereopods 2–5 meri without or with or without ill-defined tympana (fully developed only in Pseudogelasimus)�����������������������������������������6 Carapace subquadrate, as long as wide or slightly wider, lateral walls and usually dorsal surface deeply sculptured (Fig. 14.195e). Male pleonite 4 overlapping pleonite 5, setose over anterior half; pleonite 5 not constricted (Fig. 14.195n)������������������������������������������������������������������������������������������������������������������������������������������������ Dotilla Carapace pentagonal, much wider than long, lateral walls and sometimes dorsal surface granular, rarely sculptured (Fig. 14.195d). Male pleonite 4 not overlapping pleonite 5, not setose; pleonite 5 constricted (Fig. 14.195u, v)�������������������������������������������������������������������������������������������������������������������������������������� Scopimera Pereopods 2–5 meri with large tympana. Male chelipeds unequal�������������������������������������������������������������� Pseudogelasimus Pereopods 2–5 meri without or with ill-defined tympana. Male chelipeds equal�����������������������������������������������������������������7 Carapace lateral walls deeply sculptured, dorsal surface sculptured or not (Fig. 14.195a, f)�����������������������������������������������8 Carapace lateral walls rarely sculptured, dorsal surface smooth (Fig. 14.195b, g)�����������������������������������������������������������������9 Carapace dorsal surface smooth, lateral walls deeply sculptured (Fig. 14.195f). Male pleonite 4 twice as wide as pleonite 5 anterior margin (Fig. 14.195o)������������������������������������������������������������������������������Dotilloplax Carapace dorsal surface and lateral walls deeply sculptured (Fig. 14.195a). Male pleonite 4 laterally expanded, 3 times as wide as pleonite 5 anterior margin (Fig. 14.195p)�������������������������������������������� Dotillopsis Carapace anterolateral margin usually with small tooth just behind postorbital tooth (Fig. 14.195b)������������������������������������������������������������������������������������������������������������������������������������������������������������������� Ilyoplax Carapace lateral margin with prominent tooth at midpoint (Fig. 14.195g)��������������������������������������������Lazarocleistostoma Dotilla Stimpson, 1858

Diagnosis. Carapace subquadrate, as long as wide or slightly wider; lateral walls and usually dorsum deeply sculptured. Maxilliped 2 propodus greatly expanded, dactylus applied as narrow strip. Chelipeds palm and fingers slender, tapering, about as long as pereopods 2–4 in male. Pereopods 2–5 meri with large tympana. Male pleonite 4 overlapping pleonite 5, setose over anterior half, pleonite 5 not constricted. Maximum cl. 10 mm (Pl. 71n).

Intertidal. Western and Central Indo-Pacific. 9 species (Kemp 1919: key to 8 species; Mitra et al. 2020: review of species).

Dotilloplax Tweedie, 1950 Diagnosis. Carapace subquadrate, wider than long; dorsal surface smooth, lateral walls deeply sculptured. Pereopods 2–5 meri with ill-defined tympana. Male pleonite 4 twice as wide as pleonite 5, pleonite 5 not constricted. Maximum cl. 4.2 mm.

14 – Brachyura – crabs

847

Fig. 14.195.  Dotillidae. a, Dotillopsis brevitarsis (De Man, 1888) (male); b, Ilyoplax gangetica (Kemp, 1919) (male); c, Pomatocypoda pugil Tweedie, 1938 (male); d, Scopimera pilula Kemp, 1919 (male). Carapace: e, Dotilla; f, Dotilloplax; g, Lazarocleistostoma; h, Pseudogelasimus; i, Shenius; j, Tmethypocoelis. Maxilliped 2: k, Ilyoplax; l, Scopimera. Pereopod 4: m, Shenius. Male pleon: n, Dotilla; o, Dotilloplax; p, Dotillopsis; q, r, Ilyoplax; s, Potamocypoda; t, Pseudogelasimus; u, v, Scopimera; w, Shenius.

Estuarine, intertidal. Central Indo-Pacific. 1 species (Tweedie 1950).

Dotillopsis Kemp, 1919 Diagnosis. Carapace subquadrate, wider than long; dorsal surface and lateral walls deeply sculptured. Pereopods 2–5 meri with illdefined tympana. Male pleonite 4 laterally expanded, 3 times as wide as pleonite 5, pleonite 5 not constricted. Maximum cl. 8 mm. Intertidal. Western and Central Indo-Pacific. 2 species (Kemp 1919).

Ilyoplax Stimpson, 1858 Diagnosis. Carapace subquadrate, wider than long; lateral walls granular, usually not sculptured, usually with small anterolateral tooth. Pereopods 2–5 meri with tympana ill-defined or absent. Male pleonite 5 constricted, sometimes linear. Maximum cl. 7 mm (Pl. 71m). Estuarine, intertidal, subtidal. Western and Central IndoPacific. 28 species (Davie and Naruse 2010: review of species; Kitaura and Wada 2006a: evolution of waving display, distribution and ecology; Kitaura and Wada 2006b: recognition of 3 species groups; Serène and Lundoer 1974: key to 23 species).

848

Marine Decapod Crustacea

Lazarocleistostoma Števčić, 2011 Diagnosis. Carapace subquadrate, wider than long; wider than long, with 1 prominent anterolateral and 1 triangular lateral tooth. Maximum cl. 3.3 mm. Shelf. Central Indo-Pacific. 1 species (Ng et  al. 2008: noted only species close to Ilyoplax, male unknown; Števčić 2011; Tesch 1918: figure as Paracleistostoma dentatum Tesch, 1918).

Potamocypoda Tweedie, 1938 Diagnosis. Carapace subovate, wider than long; dorsal surface smooth. Male chelipeds subequal, or unequal. Pereopods 2–5 meri without tympana. Male pleonite 5 constricted. Maximum cl. 8 mm. Freshwater. Central Indo-Pacific. 2 species (Tai and Manning 1984; Tweedie 1938).

Pseudogelasimus Tweedie, 1937 Diagnosis. Carapace widest at exorbital margin, converging posteriorly or quadrate; dorsal surface with pair of prominences. Male chelipeds unequal. Pereopods 2–5 meri with large tympana. Male pleonite 5 constricted. Maximum cl. 7 mm. Estuarine. Central Indo-Pacific. 2 species (Serène 1982; Tweedie 1937).

Scopimera De Haan, 1833 Diagnosis. Carapace pentagonal, much wider than long; lateral walls granular, usually not sculptured, usually with small anterolateral tooth. Maxilliped 2 propodus greatly expanded, dactylus applied as narrow strip. Chelipeds palm and fingers slender, tapering, about as long as pereopods 2–4 in male. Pereopods 2–5 meri with large tympana. Male pleonite 5 constricted, sometimes linear. Maximum cl. 8 mm. Intertidal. Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Australasia (SE Australia). 16 species (Kemp 1919: key to 8 species; Wong et al. 2011: key to 8 species of ‘normal’ group).

Shenius Serène, 1971 Diagnosis. Carapace wider than long, with prominent postorbital tooth, 2 anterolateral teeth; dorsal surface with symmetrically arranged tubercles. Chelipeds palm and fingers slender, tapering, about as long as pereopods 2–4 in male. Pereopods 2–5 meri without tympana, with spines along extensor margin. Male pleonite 5 constricted, telson trilobed. Maximum cl. 5.5 mm. Intertidal, subtidal. Central Indo-Pacific. 1 species (Ng et  al. 2010: diagnosis of genus and subfamily, figures).

Tmethypocoelis Koelbel, 1897 Diagnosis. Carapace widest at exorbital margin, converging posteriorly; dorsal surface, lateral walls not sculptured. Eyestalk prolonged beyond cornea as styliform process. Pereopods 2–5 meri with large tympana. Male pleonite 5 not constricted. Maximum cl. 4.8 mm.

Estuarine. Central Indo-Pacific. 4 species (Davie and Kosuge 1995: new species, male cheliped waving behaviour; Davie 1990: rediagnosis). References Alcock A (1900) Materials for a carcinological fauna of India. No. 6. The Brachyura Catometopa or Grapsoidea. Journal of the Asiatic Society of Bengal 69, 279–456. Davie PJF (1990) New and rare crabs of the subfamily Dotillinae (Crustacea: Ocypodidae) from northern Australia and Papua New Guinea. Memoirs of the Queensland Museum 28, 463–473. Davie PJ, Kosuge T (1995) A new species of Tmethypocoelis (Crustacea: Brachyura: Ocypodidae) from Japan. Raffles Bulletin of Zoology 43, 207–216. Davie PJF, Naruse T (2010) A new species of Ilyoplax (Decapoda, Brachyura, Dotillidae) from Panglao, Philippines. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 75–82. Kemp BA (1919) XXII. Notes on the Crustacea Decapoda in the Indian Museum. Records of the Indian Museum 16, 305–348, pls 12–14. Kitaura J, Wada K (2006a) Evolution of waving display in brachyuran crabs of the genus Ilyoplax. Journal of Crustacean Biology 26, 455–462. doi:10.1651/S-2678.1 Kitaura J, Wada K (2006b) New species of Ilyoplax (Brachyura: Ocypodidae: Dotillinae) from the Philippines and Indonesia: behavioural, molecular, and morphological evidence. Raffles Bulletin of Zoology 54, 373–379. Mitra S, Trivedi JN, Mendoza JCE (2020) On the two ‘forms’ of Dotilla intermedia De Man, 1888 (Crustacea: Brachyura: Dotillidae) from the Bay of Bengal, India, with description of a new species. Zootaxa 4809, 521–534. doi:10.11646/zootaxa.4809.3.6 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Ng PKL, Clark PF, Cuesta JA (2010) Establishment of a new subfamily for Shenius anomalus (Shen, 1935) (Crustacea: Decapoda: Brachyura: Dotillidae. Journal of Natural History 44, 1531–1553. doi:10.1080/​ 00222931003616362 Ramesh CH, Koushik S, Shunmugaraj T, Murthy MVR (2019) Occurrence of unusual swarm of soldier crab, Dotilla myctiroides (H. Milne Edwards, 1852) in Gulf of Mannar, southeast coast of India. Emergent Life Sciences Research 5, 63–66. doi:10.31783/elsr.2019.516366 Sassa S, Yang S (2019) Suction-induced habitat selection in sand bubbler crabs. Royal Society Open Science 6, 190088. doi:10.1098/rsos.190088 Serène R (1982) Trois nouvelles espèces de Brachyoures (Crustacea, Decapoda) provenant de la baie de Nhatrang (Vietnam). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série 3, 1127–1134, pls 1–4. Serène R, Lundoer S (1974) Observations on the male pleopod of the species of Ilyoplax Stimpson with a key to the identification of the species. Phuket Marine Biological Center Research Bulletin 3, 1–10. Števčić Z (2011) Addition to the reclassification of brachyuran crabs (Crustacea: Decapoda: Brachyura). Part I. New taxa. Natura Croatica 20, 125–139. Tai A-y, Manning RB (1984) A new species of Potamocypoda (Crustacea: Brachyura: Ocypodidae) from Malaysia. Proceedings of the Biological Society of Washington 3, 615–617. Tesch JJ (1918) The Decapoda Brachyura of the Siboga Expedition. II Goneplacidae and Pinnotheridae. Siboga-Expéditie 39c1, 149–296, pls 7–18. Tweedie MWF (1937) On the crabs of the family Ocypodidae in the collection of the Raffles Museum. Bulletin of the Raffles Museum 13, 140–170.

14 – Brachyura – crabs

Tweedie MWF (1938) A new scopimerine crab from the Malay Peninsula. Bulletin of the Raffles Museum 14, 198–202. Tweedie MWF (1950) Grapsoid crabs from Labuan and Sarawak. Sarawak Museum Journal 5, 338–369. Wada K (1994) Earthen structures built by Ilyoplax dentimerosa (Crustacea, Brachyura, Ocypodidae). Ethology 96, 270–282. doi:10.1111/j.1439-0310.​ 1994.tb01015.x Wong KJ, Shih HT, Chan BK (2011) Two new species of sand-bubbler crabs, Scopimera, from North China and the Philippines (Crustacea: Decapoda: Dotillidae). Zootaxa 2962, 21–35. doi:10.11646/zootaxa.​ 2962.1.2

Heloeciidae H. Milne Edwards, 1852 semaphore crab Figure 14.196, Plate 71o The only species Heloecius cordiformis (H. Milne Edwards, 1837) is endemic to south-eastern Australia. It lives intertidally near mangroves and on seagrass sandflats where it builds permanent burrows. The species is recognised in Australia by the twisted fingers on the chelipeds. Males are territorial using their chela in a waving display (Detto et al. 2004; Griffin 1968). The species’ systematics was reviewed by Türkay (1983) who confirmed its subfamily, now family, placement. His conclusion was supported by zoeal morphology (Greenwood and Fielder 1985). Diagnosis. Carapace transversely subovate, regions weakly indicated; anterolateral margin unarmed. Chelipeds equal, larger in male; dactylus curved, twisted, without truncate tooth on occlusal margin. Gonopod 1 straight. Heloecius Dana, 1851 Maximum cl. 25 mm (Pl. 71o). Estuarine, marine, intertidal. Temperate Australasia (SE Australia). 1 species (Poore 2004: description, figure).

Fig. 14.196.  Heloeciidae, Heloecius cordiformis (H. Milne Edwards, 1837).

849

References Detto T, Zeil J, Magrath RD, Hunt S (2004) Sex, size and colour in a semiterrestrial crab, Heloecius cordiformis (H. Milne Edwards, 1837). Journal of Experimental Marine Biology and Ecology 302, 1–15. doi:10.1016/j.jembe.2003.09.023 Greenwood JG, Fielder DF (1985) The systematic position of Heloecius cordiformis (H. Milne Edwards, 1837) (Decapoda, Ocypodidae) as revealed by larval morphology. Crustaceana 48, 244–248. doi:10.1163/​ 156854085X00954 Griffin DJG (1968) Social maintenance behaviour in two Australian ocypodid crabs (Crustacea: Brachyura). Journal of Zoology 156, 291–305. doi:10.1111/j.1469-7998.1968.tb04353.x Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Türkay M (1983) The systematic position of an Australian mangrove crab Heloecius cordiformis (Crustacea: Decapoda: Brachyura). Australian Museum Memoir 18, 107–111. doi:10.3853/j.0067-1967.18.1984.376

Macrophthalmidae Dana, 1851 sentinel crabs Figures 14.197, 14.198, Plate 72a–d Sentinel crabs are common on tropical to temperate intertidal mudflats and reef flats of the West Pacific and Indian Ocean. They get their common name from the courtship cheliped-waving behaviour of the males (Kitaura and Wada 2004; McLay et  al. 2010). Sentinel crabs build permanent burrows that they inhabit when the tide is in or when disturbed while feeding while the tide is out. Most species were described as members of the oldest genus, Macrophthalmus. Subgenera were recognised (Barnes 1967, 2010) but some of these have now been elevated to generic rank. Three subfamilies are currently recognised (Ng et  al. 2008) but the recent discovery of new genera has made these difficult to define morphologically (Komai and Fujita 2018; Komai and Wada 2008). While Tritodynamia (Tritodynamiinae) is distinct from other genera, the genera presently assigned to Ilyograpsinae appear embedded in Macrophthalminae as reflected in this key and as supported by zoeal features (Flores et al. 2003). The only available molecular phylogeny does not address this issue (McLay et al. 2010). Earlier keys to genera did not include all genera now recognised (Barnes 1967, 2010; Davie 2012; Mendoza and Ng 2007). Diagnosis. Carapace anterolateral margin with 1 or more spines, or unarmed. Male chelipeds equal; dactylus straight, usually with large truncate tooth at or proximal to midlength of occlusal margin. Without brush of long setae edging pouch at base or pereopods. Male pleon without constriction near pleonite 5, without transverse setose brush on pleonite 4. Merus of walking legs without typana.

850

Marine Decapod Crustacea

Implicit generic attributes. Postfrontal ridges absent; front not constricted at base of eyestalks; suborbital margin smooth or with small granules. Eyestalk stout, cornea well developed. Maxillipeds 3 mostly closing mouthfield; maxilliped 3 dactylus small, cylindrical, articulating at end of

propodus. Cheliped fingers with acute apex; male cheliped fingers cutting margins not obscured externally by dense setae; male cheliped dactylus with dominant tooth on cutting margin.

Key to genera of Macrophthalmidae 1. – 2. – 3. – 4. – 5. – 6. –

Carapace transversely oval or hexagonal, widest at posterior margin, anterolateral margins converging anteriorly, without teeth (Fig. 14.198m). Maxilliped 3 dactylus large, flat, articulating at base of propodus (Fig. 14.198z)��������������������������������������������������������������������������������������������������������������������������������������� Tritodynamia Carapace quadrate-trapezoid, widest anterior to posterior margin, anterolateral margins not usually converging anteriorly, with teeth. Maxilliped 3 dactylus small, cylindrical, articulating at end of propodus���������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������2 Carapace longer than wide, anterolateral margin with 2 obsolete or broad teeth behind exorbital tooth (Fig. 14.198a, k)����������������������������������������������������������������������������������������������������������������������������������������������������������������������������3 Carapace wider than long, anterolateral margin with 2 or 3 teeth blunt teeth behind exorbital tooth�����������������������������4 Carapace postfrontal ridges longitudinal, with long setae; anterolateral margin oblique, second tooth prominent (Fig. 14.198a)����������������������������������������������������������������������������������������������������������������������������������������� Apograpsus Carapace postfrontal ridges absent; anterolateral margin almost parallel, second tooth obsolete (Fig. 14.198k)������������������������������������������������������������������������������������������������������������������������������������������������������������Mirograpsus Male cheliped fingers cutting margins obscured externally by dense setae; dactylus without tooth on cutting margin (Fig. 14.198o)�������������������������������������������������������������������������������������������������������������������������������Australoplax Male cheliped fingers cutting margins not obscured by dense setae; dactylus with tooth on cutting margin (Fig. 14.198p–u)���������������������������������������������������������������������������������������������������������������������������������������������������������������������������5 Front more than 0.4 exorbital width, not constricted at base of eyestalks. Carapace 1.1–1.2 times as wide as long (Fig. 14.198d, g)�������������������������������������������������������������������������������������������������������������������������������������������������������������������6 Front less than 0.3 exorbital width, constricted or not at base of eyestalks. Carapace greater than 1.2 times as wide as long�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������7 Carapace subovate, posterolateral margin convex; anterolateral margin with 3 teeth behind exorbital angle, last minute (Fig. 14.198g). Cheliped fingers spoon-shaped (Fig. 14.198s)��������������������������������������������Ilyograpsus Carapace subquadrate, posterolateral margin with rounded corner; anterolateral margin with 2 teeth behind exorbital angle. Cheliped fingers acute (Fig. 14.198q)������������������������������������������������������������������������� Enigmaplax

Fig. 14.197.  Macrophthalmidae. a, Chaenostoma punctulatum (Miers, 1884); b, Tasmanoplax latifrons (Haswell, 1881).

14 – Brachyura – crabs

7. – 8. – 9. –

851

Maxillipeds 3 gaping, not covering mouthfield (Fig. 14.198w, x). Carapace widest posteriorly. Cornea reduced, not wider than eyestalk (Fig. 14.198e, h)�����������������������������������������������������������������������������������������������������������������8 Maxillipeds 3 mostly covering mouthfield (Fig. 14.198y). Carapace usually widest anteriorly. Cornea fully pigmented, wider than eyestalk������������������������������������������������������������������������������������������������������������������������������������������������9 Maxilliped 3 ischium wider and longer than subrectangular merus (Fig. 14.198w)�������������������������������������������������� Euplax Maxilliped 3 ischium narrower and about as long as ovate merus (Fig. 14.198x)�����������������������������������������������Lutogemma Carapace front ~0.15 exorbital width, constricted at base of eyestalks (Fig. 14.198i, j, n)��������������������������������������������������10 Carapace front ~0.2 or more exorbital width, not constricted at base of eyestalks (Figs 14.197a, 14.198c, f, l)��������������������������������������������������������������������������������������������������������������������������������������������������������11

Fig. 14.198.  Macrophthalmidae. Carapace: a, Apograpsus; b, Australoplax; c, Chaenostoma; d, Enigmaplax; e, Euplax; f, Hemiplax; g, Ilyograpsus; h, Lutogemma; i, j, Macrophthalmus; k, Mirograpsus; l, Tasmanoplax; m, Tritodynamia; n, Venitus. Cheliped, outer view: o, Australoplax; p, Chaenostoma; q, Enigmaplax; r, Hemiplax; s, Ilyograpsus; t, u, Macrophthalmus. Maxilliped 3, mouthfield: v, Chaenostoma; w, Euplax; x, Lutogemma; y, Macrophthalmus; z, Tritodynamia.

852

Marine Decapod Crustacea

10.

Carapace branchial region typically with 3 clumps of granules on each posterior half; anterolateral margin with 2 or 3 teeth behind exorbital tooth; suborbital margin granular. Eyestalks more than half carapace length, extending to or beyond exorbital angle (Fig. 14.198i, j)���������������������������������������������������������������Macrophthalmus – Carapace branchial region granular; anterolateral margin with 2 broad, sharp teeth behind prominent sharp exorbital tooth; suborbital margin with large, spinose granules or tubercles. Eyestalks less than half carapace length, not reaching exorbital angle (Fig. 14.198n)��������������������������������������������������������������������������� Venitus 11. Carapace subquadrate, 1.3–1.4 times as wide as long; branchial ridges absent; anterolateral margin with 3 teeth behind exorbital tooth, last minute (Figs 14.197a, 14.198c). Cheliped fingers spoon-shaped, dactylus with prominent tooth���������������������������������������������������������������������������������������������������������������������������Chaenostoma – Carapace subquadrate, 1.5 times as wide as long; branchial ridges present; anterolateral margin with 2 teeth behind exorbital tooth (Fig. 14.198j, l). Cheliped fingers acute, dactylus cutting edge with long basal crenellate crest (Fig. 14.198r)���������������������������������������������������������������������������������������������������������������������������������������� 12 12. Carapace branchial region with 2 longitudinal ridge on each side; front 0.2 exorbital width (Fig. 14.197b, 14.198l)��������������������������������������������������������������������������������������������������������������������������������������������Tasmanoplax – Carapace branchial region with oblique and rows of granules; front 0.3 exorbital width (Fig. 14.198f)���������������Hemiplax Apograpsus Komai & Wada, 2008 Diagnosis. Carapace longer than wide; branchial region with ridge; anterolateral margins oblique, with 1 prominent tooth behind exorbital tooth; postfrontal ridges longitudinal, with long setae; front 0.4 exorbital width. Maxilliped 3 merus shorter than ischium in midline. Cheliped fingers with spoon-shaped apex. Maximum cl. 6.4 mm. Estuarine, intertidal (pebbly muddy sediments). Central IndoPacific (Ryukyu Is, Japan). 1 species (Komai and Wada 2008).

Australoplax Barnes, 1966 Diagnosis. Carapace subquadrate, 1.3 times as wide as long; branchial region without ridges; anterolateral margins with 2 blunt teeth behind exorbital tooth; front about half exorbital width. Maxilliped 3 merus shorter than ischium in midline. Male cheliped fingers cutting margins obscured externally by dense setae; dactylus without dominant tooth on cutting margin. Maximum cl. 15 mm. Estuarine, intertidal (mangroves). Central Indo-Pacific, Temperate E Australia. 1 species (Barnes 1966; Poore 2004: figure).

Chaenostoma Stimpson, 1858 Diagnosis. Carapace subquadrate, 1.3–1.4 times as wide as long; branchial region without ridges; anterolateral margins with 2 teeth behind exorbital angle, last minute; front 0.3 exorbital width. Maxilliped 3 merus shorter than ischium in midline. Cheliped fingers with spoon-shaped apex. Maximum cl. 15 mm. Intertidal. Western and Central Indo-Pacific. 7 species (Naderloo 2013: rediagnosis, key to species; Teng et al. 2016: review of species).

Enigmaplax Davie, 1993 Diagnosis. Carapace subquadrate, 1.1–1.2 times as wide as long; branchial region without ridges; anterolateral margins with 2

teeth behind exorbital angle; front one-third exorbital width. Maxilliped 3 merus shorter than ischium in midline. Maximum cl. 9 mm. Intertidal, subtidal (0–1 m; seagrass beds). Temperate E Australia. 1 species (Davie 1993).

Euplax H. Milne Edwards, 1852 Diagnosis. Carapace rounded-quadrate, widest posteriorly, 1.1–1.2 times as wide as long; branchial region granular; anterolateral margins with 2 teeth behind exorbital angle; front 0.2 exorbital width. Eyestalks thin; cornea reduced. Maxillipeds 3 gaping, not closing mouthfield; maxilliped 3 merus shorter than ischium in midline. Maximum cl. 15 mm. Estuarine, intertidal, subtidal (0–1 m; mangroves). Western and Central Indo-Pacific. 2 species (Mendoza and Ng 2007: rediagnosis, species compared).

Hemiplax Heller, 1865 Diagnosis. Carapace ~1.5 times as wide as long; branchial region with oblique rows of granules; anterolateral margins with 2 blunt teeth behind exorbital tooth; front 0.3 exorbital width. Eyestalks thin. Maxilliped 3 merus longer than ischium. Male cheliped dactylus with long basal crenellate tooth. Maximum cl. 20 mm. Intertidal–shelf (0–42 m; muddy sediments). Temperate Australasia (New Zealand endemic). 1 species (Barnes 1967: rediagnosis as subgenus; McLay et al. 2010: rediagnosis, systematic position).

Ilyograpsus Barnard, 1955 Diagnosis. Carapace subquadrate, 1.1–1.2 times as wide as long; branchial region usually with ridges; anterolateral margins with 3 teeth behind exorbital angle, last minute; front about half exorbital width. Maxilliped 3 merus shorter than ischium in midline. Cheliped fingers with spoon-shaped apex; male cheliped dactylus without dominant tooth on cutting margin, or with dominant tooth on cutting margin. Maximum cl. 10.4 mm (Pl. 72a).

14 – Brachyura – crabs

Estuarine, intertidal (mud, mangroves). Temperate Northern W Pacific, Western and Central Indo-Pacific. 5 species (Komai and Wada 2008: rediagnosis, key to species).

Lutogemma Davie, 2009 Diagnosis. Carapace subcircular, widest posteriorly, 1.3 times as wide as long; branchial region granular; anterolateral margins with 2 lobate teeth behind exorbital angle; front 0.25 exorbital width. Eyestalks thin, tapering; cornea reduced. Maxillipeds 3 gaping, not closing mouthfield; maxilliped 3 merus ovate, about as long as but wider than ischium. Maximum cl. 5.3 mm. Intertidal, subtidal (0–3 m; seagrass sediments). Central IndoPacific (NT, Australia). 1 species (Davie 2009).

Macrophthalmus Desmarest, 1823 Diagnosis. Carapace 1.5–2.0 times as wide as long; branchial region typically with 3 clumps of granules on each posterior half; front ~0.15 exorbital width, front constricted at base of eyestalks. Eyestalks thin, extending almost to and sometimes beyond exorbital angle. Maxilliped 3 merus shorter than ischium in midline. Maximum cl. 30 mm (Pl. 72b). Intertidal, subtidal (0–20 m; mud flats). Temperate Northern Pacific, Western and Central Indo-Pacific, Temperate SE Australia. 47 species (Barnes 1967: diagnosis as subgenus; Crosnier 1965: key to 6 species from Madagascar; Davie 2012: key to 12 Australian spp, some now in other genera; Naruse and Kosuge 2008: key to 8 Japanese species).

Mirograpsus Komai & Fujita, 2018 Diagnosis. Carapace longer than wide; branchial region without ridges; anterolateral margins almost parallel, with 2 obsolete behind exorbital tooth; front about half exorbital width. Maxilliped 3 merus shorter than ischium in midline. Male cheliped dactylus without dominant tooth on cutting margin. Maximum cl. 5.8 mm. Intertidal. Central Indo-Pacific (Ryukyu Is, Japan). 2 species (Komai and Fujita 2018).

Tasmanoplax Barnes, 1967 Diagnosis. Carapace ~1.5 times as wide as long; branchial region with 2 longitudinal ridges on each side; anterolateral margins with 2 blunt teeth behind exorbital tooth; front 0.2 exorbital width. Eyestalks thin. Maxilliped 3 merus longer than ischium. Male cheliped dactylus with long basal crenellate tooth. Maximum cl. 26 mm (Pl. 72c). Estuarine, intertidal (muddy seagrass beds). Temperate SE Australia. 1 species (Barnes 1967; McLay et al. 2010: rediagnosis, systematic position; Poore 2004: figure).

Tritodynamia Ortmann, 1894 Diagnosis. Carapace widest at posterior margin, transversely oval or hexagonal; branchial region without ridges; anterolateral margins

853

convex, beaded; front 0.4 exorbital width. Maxillipeds 3 gaping, not closing mouthfield; maxilliped 3 merus longer than ischium; maxilliped 3 dactylus large, flat, articulating at base of propodus (palp appears biramous). Maximum cl. 8 mm. Intertidal, subtidal (0– 14 m; muddy sediments, some species symbiotic with burrowing sedentary polychaetes and enteropneusts). Temperate Northern W Pacific, Central Indo-Pacific. 13 species (Anker and Ng 2014; Rahayu and Ng 2021; Yang and Tang 2005: key to 9 species as member of Pinnotheridae).

Venitus Barnes, 1967 Diagnosis. Carapace 1.5–2.0 times as wide as long; branchial region granular; anterolateral margins with 2 broad, sharp teeth behind sharp exorbital tooth; front ~0.15 exorbital width, front constricted at base of eyestalks; suborbital margin with large, spinose granules or tubercles. Eyestalks thin. Maxilliped 3 merus shorter than ischium in midline. Maximum cl. 60 mm (Pl. 72d). Intertidal (mangrove mudflats). Western and Central IndoPacific, Temperate Australia. 5 species (Barnes 1967: diagnosis as subgenus; Serène 1971: 2 species described). References Anker A, Ng PKL (2014) Tritodynamia serratipes sp. nov., a new marine crab from Singapore (Crustacea: Decapoda: Brachyura: Macrophthalmidae). Zootaxa 3826, 369–376. doi:10.11646/zootaxa.3826.2.6 Barnes RSK (1966) The status of the crab genus Euplax H. Milne Edwards, 1852; and a new genus Austroplax of the subfamily Macrophthalminae Dana, 1851 (Brachyura: Ocypodidae). Australian Zoologist 13, 370–376, pl. 1. Barnes RSK (1967) The Macrophthalminae of Australasia: with a review of the evolution and morphological diversity of the type genus Macrophthalmus (Crustacea: Brachyura). Transactions of the Zoological Society of London 31, 195–261. doi:10.1111/j.1096-3642.1967.tb00367.x Barnes RSK (2010) A review of the sentinel and allied crabs (Crustacea: Brachyura: Macrophthalmidae), with particular reference to the genus Macrophthalmus. Raffles Bulletin of Zoology 58, 31–49. Crosnier A (1965) Crustacés Décapodes Grapsidae et Ocypodidae. Faune de Madagascar 18, 1–143, pls 1–11. Davie PJF (1993) A new genus of macrophthalmine crab (Crustacea: Decapoda: Ocypodidae) from eastern Australia. Records of the Australian Museum 45, 5–9. doi:10.3853/j.0067-1975.45.1993.125 Davie PJF (2009) A new genus and species of Macrophthalmidae (Brachyura, Ocypodidae) from the Northern Territory, Australia. Crustaceana 82, 815–827. doi:10.1163/156854009X427360 Davie PJF (2012) A review of Macrophthalmus sensu lato (Crustacea: Decapoda: Macrophthalmidae) from Australia, including two new species and new records. Memoirs of the Queensland Museum – Nature 56, 149–219. Flores AAV, Paula J, Dray T (2003) First zoeal stages of grapsoid crabs (Crustacea: Brachyura) from the East African coast. Zoological Journal of the Linnean Society 137, 355–383. doi:10.1046/j.1096-3642.​ 2003.00054.x Kitaura J, Wada K (2004) Allocleaning, fighting, waving and mating behavior in sentinel crabs (Brachyura: Ocypodoidea: Macrophthalmus). Crustacean Research 33, 72–91. doi:10.18353/crustacea.33.0_72 Komai T, Fujita Y (2018) A new genus and new species assigned to Macrophthalmidae (Decapoda: Brachyura: Thoracotremata) from the Ryukyu Islands, Japan. Zootaxa 4531, 109–116. doi:10.11646/zootaxa.​ 4531.1.5

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Komai T, Wada K (2008) A revision of the estuarine crab genus Ilyograpsus Barnard, 1955 (Crustacea: Decapoda: Brachyura: Macrophthalmidae), with descriptions of a new genus and one new species. Raffles Bulletin of Zoology 52, 357–384. McLay CL, Kitaura J, Wada K (2010) Behavioural and molecular evidence for the systematic position of Macrophthalmus (Hemiplax) hirtipes Hombron & Jacquinot, 1846, with comments on macrophthalmine subgenera (Decapoda, Brachyura, Macrophthalmidae). In: Fransen C, De Grave S, Ng P (Eds) Studies on Malacostraca: Lipke Bijdeley Holthuis Memorial Volume. Crustaceana Monographs 14, 483–503. Mendoza JCE, Ng PKL (2007) Macrophthalmus (Euplax) H. Milne Edwards, 1852, a valid subgenus of ocypodoid crab (Decapoda: Brachyura: Macrophthalmidae), with description of a new species from the Philippines. Journal of Crustacean Biology 27, 670–680. doi:10.1651/S-2779.1 Naderloo R (2013) The sentinel crabs of the genus Chaenostoma (Stimpson, 1858) (Crustacea: Brachyura: Macrophthalmidae), with description of a new species and new records. Journal of Natural History 47, 2835– 2848. doi:10.1080/00222933.2013.807948 Naruse T, Kosuge T (2008) A new species of Macrophthalmus (Crustacea: Decapoda: Brachyura: Macrophthalmidae) from Iriomote Island, Ryukyu Islands, Japan. Species Diversity 13, 117–122. doi:10.12782/ specdiv.13.117 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Poore GCB (2004) Marine decapod Crustacea of southern Australia. A guide to identification. (Chapter 12. Ahyong, S, Stomatopoda – mantis shrimps). CSIRO Publishing, Melbourne. Rahayu DL, Ng PKL (2021) A new species of Tritodynamia (Brachyura, Macrophthalmidae) from Papua, Indonesia. Crustaceana 94, 1187–1200. Serène R (1971) Observations préliminaires sur des brachyoures nouveaux ou mal connus du Sud-Est Asiatique (Crustacea Decapoda). Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e série 42, 903–918. Teng S-J, Shih H-T, Naderloo R, Corbari L (2016) A review of the Chaenostoma boscii species-complex (Decapoda: Brachyura: Macrophthalmidae) from the Indo-West Pacific. Crustacean Research 45, 15–27. doi:10.18353/crustacea.45.0_15 Yang S, Tang B (2005) A new species of Tritodynamia (Decapoda, Brachyura, Pinnotheridae) from the Bohai Sea, China, with a key to the genus Tritodynamia Ortmann, 1894. Crustaceana 78, 781–786. doi:10.1163/​ 156854005774445528

Mictyridae Dana, 1851 soldier crabs Figure 14.199, Plates 1e, 72m Soldier crabs are conspicuous on many shores throughout tropical and temperate Australia, and the Western Pacific. They are ecologically important on intertidal flats where they occur in large numbers, often swarming in their thousands, foraging as the tide falls (Pl. 1e, Webb and Eyre 2004). Mictyris are among the few brachyurans to walk forwards, rather than sideways; they burrow by corkscrewing into the sand but do not make permanent burrows. They feed on meiofauna and detritus while tunnelling (Davie et al. 2013).

Fig. 14.199.  Mictyridae. Mictyris longicarpus Latreille, 1806.

Soldier crabs are taller and more globose than typical crabs, and often brightly coloured. The species have not been reviewed since McNeill’s (1926) review of three species and one variety. Eight species are now known (Davie et al. 2013; Davie et  al. 2010; Unno 2008; Unno and Semeniuk 2011). Unno and Semeniuk (2011) provided a key to seven species. Diagnosis. Carapace high, subglobular, longer than wide, cervical and cardiobranchial grooves well defined. Eyestalk long, slender, standing well beyond carapace margin. Chelipeds elongated, slender, subequal, similar in both sexes. Pleon similar in both sexes, broad, nearly covering most of thoracic sternum. Mictyris Latreille, 1806 Maximum cl. 25 mm (Pls 1e, 72m). Marine, intertidal. Western and Central Indo-Pacific, Temperate Australia. Maximum cl. 25 mm. 8 species. References Davie PJF, Shih H-T, Chan BKK (2010) A new species of Mictyris (Decapoda, Brachyura, Mictyridae) from the Ryukyu Islands, Japan. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 83–105. Davie PJ, Wisespongpand P, Shih HT (2013) A new species of Mictyris Latreille, 1806 (Crustacea: Decapoda: Brachyura: Mictyridae) from the Andaman coast of Thailand, with notes on its ecology and behavior. Zootaxa 3686, 65–76. doi:10.11646/zootaxa.3686.1.3 McNeill FA (1926) Studies in Australian carcinology. No. 2. Records of the Australian Museum 15, 100–131, pls 9, 10. Unno J (2008) A new species of soldier crab, Mictyris occidentalis (Crustacea: Decapoda: Brachyura: Mictyridae) from Western Australia, with congener comparisons. Journal of the Royal Society of Western Australia 91, 31–50. Unno J, Semeniuk V (2011) A new species of Mictyris (Crustacea: Decapoda: Mictyridae) recorded from northern Australia – Kimberley

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region to Cape York. Journal of the Royal Society of Western Australia 94, 45–54. Webb AP, Eyre BD (2004) The effect of natural populations of the burrowing and grazing soldier crab (Mictyris longicarpus) on sediment irrigation, benthic metabolism and nitrogen fluxes. Journal of Experimental Marine Biology and Ecology 309, 1–19. doi:10.1016/j.jembe.2004.05.003

Ocypodidae Rafinesque, 1815 fiddler crabs, ghost crabs Figure 14.200, Plate 72e–l Ocypodidae are a group of intertidal or semiterrestrial burrowing crabs found on tropical to warm temperate shores worldwide. The family can be divided into two main groups. Ghost crabs of the genus Ocypode are mainly inhabitants of tropical beaches and dunes exposed to the ocean. They run quickly over the beach, especially at night, where they scavenge and are able to forage almost a kilometre away from the sea. Ghost crabs are the fastest moving crustaceans on land but retreat to deep burrows above the high tide mark when not feeding. All have long eyes, held erect when the crab is active. Lucrezi and Schlacher (2014) reviewed the biology of ghost crabs. Species of Uca and related genera, fiddler crabs, have attracted considerable attention because of their bright colours, aggressive waving and fighting behaviour, and the extreme claw asymmetry found in males. The massively enlarged cheliped is used in mate attraction and competition; the small cheliped is used for feeding. Fiddler crabs inhabit intertidal sand and mudflats in estuaries and mangrove forests (Kristensen 2008; Rosenberg 2020; SayãoAguiar et  al. 2012). They too excavate burrows and may build complex structures around the opening that aid in attracting mates (Kim et al. 2004; Shih et al. 2005). The genus Uca has a complex and contested systematic history (Shih et al. 2010). Bott (1973) divided the genus into eight genera and subgenera. The genus was treated in detail by Crane (1975) who divided Uca into nine subgenera, recognising few of Bott’s taxa. The conflicts between these two schemes were addressed using cladistic methods by Beinlich

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and von Hagen (2006) who diagnosed eight subgenera. More recently a reappraisal of external and gastric mill morphology, and molecular studies (Shih et al. 2015; Shih et al. 2016b) concluded that three subfamilies and 13 genera, including Ocypode, can be supported within Ocypodidae. Most of Bott’s and Crane’s genera and subgenera, but not Beinlich and von Hagen’s, were synonymised. The genus v. subgenus status of some taxa was contested by Rosenberg (2019) who added more subgenera not included here. The distributions of all species are well documented (Rosenberg 2020). Here, we recognise four subfamilies and follow the generic rediagnoses of Shih et  al. (2016b). They used the morphology of the gastric mill in their key and diagnoses of gelasimine genera. These characters are included in our generic diagnoses but not in the key. Shih et al. (2016b) listed most of the species in each of the genera now recognised but few keys to species exist. Rathbun (1918) provided a key to 32 American species of ‘Uca’, many now Leptuca or Minuca. Regional keys to species of ‘Uca’ can be found in Crane’s (1975) substantial review but nomenclature has moved on since then and twice as many species are now known. A key to 19 Australian species of ‘Uca’ can be found in George and Jones (1982) and another to 18 East Asian species of ‘Uca’ and five subgenera (now genera) is in Shih et al. (2016a). Some of the controversy regarding subdivision of Uca depended on the front of the carapace being ‘broad’ or ‘narrow’. Neither condition is monophyletic. Nevertheless, the front is useful taxonomically and we use the descriptor ‘broad’ to indicate 0.2 maximum carapace width and ‘narrow’ to indicate ~0.1 maximum carapace width (Shih et  al. 2016b). Narrow fronts are sometimes waisted. All maximum sizes for this family are indicated by carapace width (cw.). Diagnosis. Carapace subquadrilateral to pentagonal, regions typically indistinct; orbit transverse or sloping posteriorly, placed along anterior carapace margin. Male chelipeds unequal.

Key to subfamilies and genera of Ocypodidae 1. – 2. –

Fronto-orbital distance 0.5–0.7 maximum carapace width. Front broad (Fig. 14.200i). Without brush of long setae between pereopods 3 and 4 bases�����������������������������������������������������������Ucidinae … Ucides Fronto-orbital distance 0.9 or more of maximum carapace width. Front narrow or broad. With brush of long setae between pereopods 3 and 4 bases�������������������������������������������������������������������������������������������������2 Orbital floor with tubercle at inner corner adjacent to antennule (Fig. 14.200j). Cornea of male of some species with distal ornament (horn, stylus or setal brush) (Fig. 14.200g)��������������������������������������������������������������������������3 Orbital floor without tubercle at inner corner adjacent to antennule (Fig. 14.200m) (small ridge in Petruca, Fig. 14.200k). Cornea of male without distal ornament (Fig. 14.200k–m)����������������������� Gelasiminae … 5

856

3. – 4. – 5. – 6. –

Marine Decapod Crustacea

Eyestalk thick; cornea large, ovate (Fig. 14.200j). Chelipeds unequal in both sexes; major cheliped not grossly enlarged (Fig. 14.200g)���������������������������������������������������������������������Ocypodinae … Ocypode Eyestalk slender, cornea round (Fig. 14.200e, h). Major cheliped of male grossly enlarged; minor cheliped of male and female small����������������������������������������������������������������������������������������������������������� Ucinae … 4 Carapace front narrow; not covered with tubercles (Fig. 14.200h)�������������������������������������������������������������������������������������Uca Carapace front wide; covered with prominent tubercles (Fig. 14.200e)������������������������������������������������������������������������Afruca Carapace front narrow (Fig. 14.200a, d, l, m)������������������������������������������������������������������������������������������������������������������������������6 Carapace front wide (Fig. 14.200b, c, f, k)������������������������������������������������������������������������������������������������������������������������������������8 Major cheliped carpus upper surface facing propodus not flat (Fig. 14.200a). Most males with major chela on right. Orbital floor without ornamentation. Carapace without striae�������������������������� Gelasimus Major cheliped carpus upper surface facing propodus flat, with sharp edge (Fig. 14.200d). Males with major chela equally on left and right. Orbital floor often with tubercles (Fig. 14.200l), ridge or mound. Carapace with posterolateral tuberculate stria, at least in female (Fig. 14.200o)��������������������������������������������������������������7

Fig. 14.200.  Ocypodidae. a, Gelasimus jocelynae (Shih, Naruse & Ng, 2010); b, Minuca burgersi (Holthuis, 1967); c, Petruca panamensis (Stimpson, 1859); d, Xeruca formosensis (Rathbun, 1921). Carapace: e, Afruca; f, Austruca; g, Ocypode; h, Uca; i, Ucides. Left orbit, anterior view: j, Ocypode; k, Petruca; l, Tubuca; m Xeruca. Carapace, right posterolateral corner with striae: n, Petruca; o Xeruca (female). Male major chela, outer view: p, Austruca; q, Cranuca; r, Gelasimus; s, Paraleptuca; t, Tubuca; u, Xeruca. Gonopod 1 apex: v, Leptuca; w, Minuca.

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857

7.

Major cheliped fingers forceps-shaped or with straight cutting margins not more than half finger length (Fig. 14.200t). Minor cheliped merus with short, stiff setae������������������������������������������������������Tubuca – Major cheliped fingers deep, cutting margins straight, more than half finger length (Fgi.14.200u). Minor cheliped merus with long, thin setae��������������������������������������������������������������������������������������� Xeruca 8. Dorsal edge of orbit broad (Fig. 14.200k)�������������������������������������������������������������������������������������������������������������������������������������9 – Dorsal edge of orbit narrow����������������������������������������������������������������������������������������������������������������������������������������������������������11 9. Major cheliped propodus expanded proximally on upper margin (Fig. 14.200c). Minor cheliped fingers with apical brush of long setae. Orbital floor with small ridge near inner corner������������������ Petruca – Major cheliped propodus not expanded proximally on upper margin (Fig. 14.200b). Minor cheliped fingers without apical brush of long setae. Orbital floor without ridge near inner corner���������������������������10 10. Carapace with 0–2 posterolateral striae (Fig. 14.200n). Gonopod 1 slightly curved distally, ‘thumb’ small or absent. Pleonites free, or pleonites 4–6 partly or completely fused���������������������������������������������������������Leptuca – Carapace with 2 posterolateral striae. Gonopod 1 with 2 apical flanges, ‘thumb’ subdistal (Fig. 14.200w). Pleonites free������������������������������������������������������������������������������������������������������������������������ Minuca 11. Major cheliped dactylus with subdistal tooth; propodus outer face tuberculate (Fig. 14.200q)�����������������������������Cranuca – Major cheliped dactylus without subdistal tooth; propodus outer face smooth (Fig. 14.200p, s)������������������������������������ 12 12. Major cheliped propodus with outer depression near base of fixed finger; carpal cavity not continued distally (Fig. 14.200s)����������������������������������������������������������������������������������������������������������������������������Paraleptuca – Major cheliped propodus without outer depression near base of fixed finger; carpal cavity with distal extension (Fig. 14.200p)������������������������������������������������������������������������������������������������������������������������������������ Austruca

Subfamily Gelasiminae Miers, 1886 fiddler crabs Most species formerly in the genus Uca are now in the nine gelasimine genera. They lack a tubercle at the inner corner of the orbital floor, substituted in the only species of Petruca with a low ridge. Distal ornaments on the eyestalk are never found in species of Gelasiminae. Diagnosis. Fronto-orbital distance 0.9 or more maximum carapace width; front broad, or narrow; orbital floor without or with vestigial tubercle at inner corner. With brush of long setae between pereopods 3 and 4 bases. Eyestalk slender, cornea round; cornea of male without distal ornament. Chelipeds strongly unequal in adult male; both chelipeds small in female. Implicit generic attributes. Anterolateral margin long, curving into upper-lateral surface. Males with major chela equally on left and right. Major cheliped carpus upper surface facing propodus not flat. Major cheliped propodus not expanded proximally on upper margin. Minor cheliped fingers without apical brush of long setae. Pleonal locking apparatus absent. Austruca Bott, 1973 Diagnosis. Gelasiminae. Carapace with 0–2 posterolateral striae; front broad; dorsal edge of orbit narrow. Major cheliped propodus outer face smooth, without small round outer depression near

base of fixed finger; carpal cavity with distal extension; fixed finger lower margin sometimes carinate; dactylus without subdistal tooth. Pleonal locking apparatus present. Gastric mill without large brownish setae at base of posterior tooth plate. Maximum cw. 15 mm (Pl. 72e). Estuarine, intertidal. Western and Central Indo-Pacific, Temperate SE Australia. 13 species (Naderloo et al. 2010: key to 6 species; Shih and Poupin 2020: new species, 3 species groups, synonymies of all species).

Cranuca Beinlich & von Hagen, 2006 Diagnosis. Gelasiminae. Carapace smooth; front broad; dorsal edge of orbit narrow. Major cheliped propodus outer face tuberculate; fixed finger lower margin not carinate; dactylus with large subdistal tooth. Gastric mill without large brownish setae at base of posterior tooth plate. Maximum cw. 20 mm. Estuarine, intertidal. Western Indo-Pacific. 1 species (Beinlich and von Hagen 2006).

Gelasimus Latreille, 1817 Diagnosis. Gelasiminae. Carapace smooth; front narrow; orbital floor without ornamentation. Most males with major chela on right. Major cheliped propodus outer face tuberculate; fixed finger lower margin not carinate. Gastric mill with setae on lateral margins of posterior stem region of urocardiac ossicles. Maximum cw. 30 mm (Pl. 72f). Estuarine, intertidal. Indo-West Pacific, Temperate SE Australia. 8 species (Shih et al. 2010: relationships, new species).

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Leptuca Bott, 1973 Diagnosis. Gelasiminae. Carapace with 0–2 posterolateral striae; front broad; anterolateral margin short; dorsal edge of orbit broad. Major cheliped propodus outer face smooth; fixed finger lower margin sometimes carinate. Gonopod 1 slightly curved distally, ‘thumb’ small or absent. Gastric mill with 2 large brown setae at base of posterior tooth plate. Maximum cw. 25 mm. Estuarine, intertidal. Tropical W Atlantic, Tropical Eastern Pacific. 29 species (Crane 1975: key to 7 species as Uca).

Minuca Bott, 1954 Diagnosis. Gelasiminae. Carapace with 2 posterolateral striae; front broad; dorsal edge of orbit broad. Major cheliped propodus outer face tuberculate, or outer face smooth; fixed finger lower margin not carinate. Gonopod 1 with 2 apical flanges, ‘thumb’ subdistal. Pleonal locking apparatus carina-like. Gastric mill with 2 large brown setae at base of posterior tooth plate. Maximum cw. 30 mm (Pl. 72g). Estuarine, intertidal. Tropical W Atlantic, Tropical Eastern Pacific. 18 species (Landstorfer and Schubart 2010: diagnoses of 8 Pacific species).

Paraleptuca Bott, 1973 Diagnosis. Gelasiminae. Carapace with 4 posterolateral striae; front broad; dorsal edge of orbit narrow. Major cheliped propodus outer face smooth, with small round outer depression near base of fixed finger; carpal cavity not continued distally; fixed finger lower margin not carinate; dactylus without subdistal tooth. Pleonal locking apparatus present. Gastric mill without large brownish setae at base of posterior tooth plate. Maximum cw. 20 mm (Pl. 72i). Estuarine, intertidal. Indo-West Pacific. 4 species (Shih et al. 2013: new species, relationships).

Petruca Shih, Ng & Christy, 2015 Diagnosis. Gelasiminae. Carapace with posterolateral striae; front broad; orbital floor with small ridge near inner corner; dorsal edge of orbit broad. Major cheliped propodus expanded proximally on upper margin; fixed finger lower margin not carinate. Minor cheliped fingers with apical brush of long setae. Gastric mill with 2 large brown setae at base of posterior tooth plate. Maximum cw. 15 mm. Intertidal (cobble beaches, not burrowing). Eastern IndoPacific. 1 species (Shih et al. 2015).

Tubuca Bott, 1973 Diagnosis. Gelasiminae. Carapace with posterolateral striae in female, absent in male; front narrow. Major cheliped carpus upper surface facing propodus flat, with sharp edge. Major cheliped propodus outer face tuberculate; fingers forceps-shaped or with straight cutting margins less than half finger length; fixed finger lower margin sometimes carinate. Minor cheliped merus with short, stiff setae. Gastric mill without setae on lateral

margins of posterior stem region of urocardiac ossicles. Maximum cw. 35 mm (Pl. 72j). Estuarine, intertidal. Western and Central Indo-Pacific. 22 species (Shih et al. 2018: new species).

Xeruca Shih, 2015 Diagnosis. Gelasiminae. Carapace smooth, female with margins beaded, with tuberculate swelling behind dorsolateral margin; front narrow. Major cheliped carpus upper surface facing propodus flat, with sharp edge. Major cheliped propodus outer face tuberculate; fingers deep, cutting margins straight, more than half finger length; fixed finger lower margin not carinate. Minor cheliped merus with long, thin setae. Gastric mill without setae on lateral margins of posterior stem region of urocardiac ossicles. Maximum cw. 35 mm. Estuarine, intertidal. Central Indo-Pacific (Taiwan). 1 species (Shih et al. 1999; Shih 2015).

Subfamily Ocypodinae Rafinesque, 1815 ghost crabs Ghost crabs inhabit sandy beaches, not muddy environments as do the other subfamilies. The eyestalk is thick with large ovate cornea and may possess a distal ornament. While the chelipeds are unequal in both sexes, males never have one grossly enlarged. Ocypode is the only genus. It was revised by Sakai and Türkay (2013) who introduced a new genus Hoplocypode Sakai & Türkay, 2013, but this was synonymised by Shih et al. (2016b). Diagnosis. Fronto-orbital distance 0.9 or more maximum carapace width; front narrow; orbital floor with tubercle at inner corner. With brush of long setae between pereopods 3 and 4 bases. Eyestalk thick; cornea large, ovate; cornea of male with or without distal ornament. Chelipeds unequal in both sexes; major cheliped not enlarged. Ocypode Weber, 1795 Maximum cw. 59 mm (Pl. 72h). Intertidal, burrowing well above the high tide mark. Cosmopolitan, tropical to warm temperate shores, except Eastern Indo-Pacific. 26 species (Sakai and Türkay 2013: rediagnosis, key to 22 species; Shih et al. 2016b: discussion of genus and species synonymies).

Subfamily Ucidinae Števčić, 2005 mangrove crabs The fronto-orbital width is much narrower than in the other subfamilies. Ucides is the only genus with one species each on the eastern and western coasts of the Americas. The only two species burrow in the high to mid-intertidal area

14 – Brachyura – crabs

and were long thought to belong to Gecarcinidae, a family of land crabs. They are clearly ocypodids (Chace and Hobbs 1969; Tavares 1990). Ucides cordatus (Linnaeus, 1763) is a large crab exploited for human consumption in Brazil (Côrtes et al. 2019). Diagnosis. Fronto-orbital distance 0.5–0.7 maximum carapace width; front broad; orbital floor with tubercle at inner corner. Without brush of long setae between pereopods 3 and 4 bases. Eyestalk short, cornea terminal; cornea of male without distal ornament. Chelipeds strongly unequal in adult male, less so in female. Ucides Rathbun, 1897 Maximum cw. 75 mm (Pl. 72l). Estuarine, intertidal (semiterrestrial mangrove forests). Tropical W Atlantic, Tropical Eastern Pacific. 2 species (Rathbun 1918: rediagnosis, key to species).

Subfamily Ucinae Dana, 1851 fiddler crabs The subfamily contains Uca, the generic name widely used for all fiddler crabs in the past. Uca is now confined to the Americas. The second genus includes only one species from the Eastern Atlantic. Diagnosis. Fronto-orbital distance 0.9 or more maximum carapace width; front broad, or narrow; orbital floor with tubercle at inner corner. With brush of long setae between pereopods 3 and 4 bases. Eyestalk slender, cornea round; cornea of male without distal ornament, or with spine on major side. Major cheliped of male enlarged; minor cheliped of male and both of female small. Afruca Crane, 1975 Diagnosis. Ucinae. Carapace mostly covered with prominent tubercles; front broad. Maximum cw. 35 mm. Intertidal. Temperate Northern and Tropical E Atlantic. 1 species (Wolfrath 1993: behaviour).

Uca Leach, 1814 Diagnosis. Ucinae. Carapace smooth; front narrow. Maximum cw. 40 mm (Pl. 72k). Estuarine, intertidal. Tropical W Atlantic, Tropical Eastern Pacific. 10 species (Rathbun 1918: key to species of Uca sensu lato). References Beinlich B, von Hagen HO (2006) Materials for a more stable subdivision of the genus Uca Leach. Zoölogische Mededeelingen 80, 9–32. Bott R (1973) Die verwandtschaftlichen Beziehungen der Uca Arten. Senckenbergiana Biologica 54, 315–325.

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Chace FA, Jr, Hobbs HH, Jr (1969) Bredin-Archbold-Smithsonian Biological survey of Dominica. The freshwater and terrestrial decapod crustaceans of the West Indies with special reference to Dominica. Bulletin of the United States National Museum 292, 1–258. doi:10.5479/ si.03629236.292.1 Côrtes LHdO, Zappes CA, Di Beneditto APM (2019) Sustainability of mangrove crab (Ucides cordatus) gathering in the southeast Brazil: A MESMIS-based assessment. Ocean and Coastal Management 179, 104862. doi:10.1016/j.ocecoaman.2019.104862 Crane J (1975) Fiddler crabs of the World. Ocypodidae: genus Uca. Princeton University Press, Jersey. George RW, Jones DS (1982) A revision of the fiddler crabs of Australia (Ocypodinae: Uca). Records of the Western Australian Museum Supplement 14, 1–99. Kim TW, Christy JH, Choe JC (2004) Semidome building as sexual signaling in the fiddler crab Uca lactea (Brachyura:Ocypodidae). Journal of Crustacean Biology 24, 673–679. doi:10.1651/C-2471 Kristensen E (2008) Mangrove crabs as ecosystem engineers; with emphasis on sediment processes. Journal of Sea Research 59, 30–43. doi:10.1016/j.seares.2007.05.004 Landstorfer RB, Schubart CD (2010) A phylogeny of Pacific fiddler crabs of the subgenus Minuca (Crustacea, Brachyura, Ocypodidae: Uca) with the description of a new species from a tropical gulf in Pacific Costa Rica. Journal of Zoological Systematics and Evolutionary Research 48, 214–218. Lucrezi S, Schlacher TA (2014) The ecology of ghost crabs. Oceanography and Marine Biology - an Annual Review 58, 201–256. Naderloo R, Türkay M, Chen HL (2010) Taxonomic revision of the widefront fiddler crabs of the Uca lactea group Crustacea Decapoda Brachyura Ocypodidae in the Indo-West Pacific. Zootaxa 2500, 1–38. doi:10.11646/zootaxa.2500.1.1 Rathbun MJ (1918) The grapsoid crabs of America. Bulletin of the United States National Museum 97, 1–461, pls 1–161. Rosenberg MS (2019) A fresh look at the biodiversity lexicon for fiddler crabs (Decapoda: Brachyura: Ocypodidae). Part 1: Taxonomy. Journal of Crustacean Biology 39, 729–738. doi:10.1093/jcbiol/ruz057 Rosenberg MS (2020) A fresh look at the biodiversity lexicon for fiddler crabs (Decapoda: Brachyura: Ocypodidae). Part 2: Biogeography. Journal of Crustacean Biology 40, 364–383. doi:10.1093/jcbiol/ruaa029 Sakai K, Türkay M (2013) Revision of the genus Ocypode with the description of a new genus, Hoplocypode (Crustacea: Decapoda: Brachyura). Memoirs of the Queensland Museum – Nature 56, 665–793. Sayão-Aguiar B, Pinheiro MAA, Colpo KD (2012) Sediment bioturbation potential of Uca rapax and Uca uruguayensis as a result of their feeding activity. Journal of Crustacean Biology 32, 223–229. doi:10.1163/193724011X615451 Shih HT (2015) Uca (Xeruca), a new subgenus for the Taiwanese fiddler crab Uca formosensis Rathbun, 1921 (Crustacea: Decapoda: Ocypodidae), based on morphological and molecular evidence. Zootaxa 3974, 151–169. doi:10.11646/zootaxa.3974.2.1 Shih HT, Poupin J (2020) A new fiddler crab of Austruca Bott, 1973, closely related to A. perplexa (H. Milne Edwards, 1852) (Crustacea: Brachyura: Ocypodidae), from the South Pacific Islands. Zoological Studies 59, e26. doi:10.6620/ZS.2020.59-26. Shih H-T, Mok H-K, Chang H-W, Lee S-C (1999) Morphology of Uca formosensis Rathbun, 1921 (Crustacea: Decapoda: Ocypodidae), and endemic fiddler crab from Taiwain, with notes on its ecology. Zoological Studies 38, 164–177. Shih H-T, Mok H-K, Chang H-W (2005) Chimney building by male Uca formosensis Rathbun, 1921 (Crustacea: Decapoda: Ocypodidae) after pairing: a new hypothesis for chimney function. Zoological Studies 44, 242–251.

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Shih HT, Naruse T, Ng PKL (2010) Uca jocelynae sp. nov., a new species of fiddler crab (Crustacea: Brachyura: Ocypodidae) from the Western Pacific. Zootaxa 2337, 47–62. doi:10.11646/zootaxa.2337.1.4 Shih HT, Komai T, Liu MY (2013) A new species of fiddler crab from the Ogasawara (Bonin) Islands, Japan, separated from the widely-distributed sister species Uca (Paraleptuca) crassipes (White, 1847) (Crustacea: Decapoda: Brachyura: Ocypodidae). Zootaxa 3746, 175–193. doi:10.11646/zootaxa.3746.1.8 Shih HT, Ng PKL, Christy JH (2015) Uca (Petruca), a new subgenus for the rock fiddler crab Uca panamensis (Stimpson, 1859) from Central America, with comments on some species of the American broad-fronted subgenera. Zootaxa 4034, 471–494. doi:10.11646/zootaxa.4034.3.3 Shih HT, Lee JH, Ho PH, Liu HC, Wang CH, et al. (2016a) Species diversity of fiddler crabs, genus Uca Leach, 1814 (Crustacea: Ocypodidae), from Taiwan and adjacent islands, with notes on the Japanese species. Zootaxa 4083, 57–82. doi:10.11646/zootaxa.4083.1.3 Shih HT, Ng PKL, Davie PJF, Schubart CD, Türkay M, et al. (2016b) Systematics of the family Ocypodidae Rafinesque, 1815 (Crustacea: Brachyura), based on phylogenetic relationships, with a reorganization of subfamily rankings and a review of the taxonomic status of Uca Leach, 1814, sensu lato and its subgenera. Raffles Bulletin of Zoology 64, 139–175. Shih HT, Chan BKK, Ng PKL (2018) Tubuca alcocki, a new pseudocryptic species of fiddler crab from the Indian Ocean, sister to the southeastern African T. urvillei (H. Milne Edwards, 1852) (Crustacea, Decapoda, Brachyura, Ocypodidae). ZooKeys 747, 41–62. doi:10.3897/zookeys.747.23468 Tavares MS (1990) Considerations sur la position systematique du genre Ucides Rathbum, 1897 (Crustacea, Decapoda, Ocypodidae). Boletim do Museu Nacional, Rio de Janeiro (nova série, Zoologia) 342, 1–8.

Wolfrath B (1993) Observations on the behaviour of the European fiddler crab Uca tangeri. Marine Ecology Progress Series 100, 111–118. doi:10.3354/meps100111

Xenophthalmidae Stimpson, 1858 Figure 14.201 Xenophthalmids are small, ovate free-living crabs from intertidal and shallow muddy marine and estuarine environments. They are characterised by the pronounced swelling of the pterygostomian region. Xenophthalmids resemble pea-crabs and were long considered to belong to Pinnotheridae. They were excluded from this family by Serène and Umali (1972) who recognised two subfamilies. Ng et  al. (2008) recognised the crabs’ thoracotreme affinities and placed the family within Ocypodoidea. Only a five species are known (Khot et al. 2019). Diagnosis. Carapace subtrapezoidal, thickened, elliptical in sagittal section. Orbit placed dorsally, forming longitudinal slits perpendicular to carapace front. Maxilliped 3 merus and ischium with distinct oblique setose ridge. Chelipeds in both sexes weak. Pleon narriow in both sexes; pleonites free.

Key to subfamilies and genera of Xenophthalmidae 1. – 2.

Buccal cavity closed anteriorly by median piece of buccal frame; epistome small (Fig. 14.201d)������������������������������������������������������������������������������������� Anomalifrontinae … Anomalifrons Buccal cavity open anteriorly to antennular fossae, buccal frame without median piece; epistome absent (Fig. 14.201e)�����������������������������������������������������������������������������������������������������������Xenophthalminae … 2 Carapace covered by long fine setae, pereopods setose. Carapace without transverse rim across branchial and cardiac regions; orbit directed posteriorly (Fig. 14.201c, e). Maxilliped 3 with large setose palp (Fig. 14.201g). Gonopod 1 apex with spiniform setae����������������������������������������������������������������������������������� Xenopthalmus

Fig. 14.201.  Xenophthalmidae. Carapace: a, Anomalifrons; b, Neoxenophthalmus; c, Xenophthalmus. Front, orbits, antennular fossae (shaded): d, Anomalifrons; e, Xenophthalmus (with epistome). Maxilliped 3: f, Neoxenophthalmus; g, Xenophthalmus.

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861

Carapace, antennae, and pereopods almost bare. Carapace with transverse rim across branchial and gastric regions; orbit directed obliquely posterolaterally (Fig. 14.201b). Maxilliped 3 with small palp (Fig. 14.201f). Gonopod 1 apex without spiniform setae������������������������������������������Neoxenophthalmus Anomalifrons Rathbun, 1931

Diagnosis. Carapace, antennae and pereopods almost bare; with transverse rim across branchial and cardiac region; orbit directed obliquely posterolaterally. Buccal cavity closed anteriorly by median piece of buccal frame; epistome small. Maxilliped 3 with small palp. Gonopod 1 apex without spiniform setae. Maximum cl. 10 mm. Intertidal, subtidal (0–7 m). Western and Central Indo-Pacific. 2 species (Khot et  al. 2019; descriptions, figures: Serène and Umali 1972).

Neoxenophthalmus Serène & Umali, 1972 Diagnosis. Carapace, antennae and pereopods almost bare; with transverse rim across branchial and cardiac region; orbit directed obliquely posterolaterally. Buccal cavity open anteriorly to antennular fossae, buccal frame without median piece; epistome absent. Maxilliped 3 with small palp. Gonopod 1 apex without spiniform setae. Maximum cl. 9 mm. Subtidal (6–10 m). Western and Central Indo-Pacific. 1 species (Serène and Umali 1972).

Xenophthalmus White, 1846 Diagnosis. Carapace covered by long fine setae, pereopods very setose; without transverse rim across branchial and cardiac regions; orbit directed posteriorly. Buccal cavity open anteriorly to antennular fossae, buccal frame without median piece; epistome absent. Maxilliped 3 with large setose palp. Gonopod 1 apex with spiniform setae. Maximum cl. 13 mm. Estuarine, intertidal, subtidal (0–10 m). Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate SE Australia. 2 species (Khot et al. 2019; descriptions, figures: Serène and Umali 1972). References Khot M, Jaiswar AK, Ng PKL (2019) The taxonomy of two species of Xenophthalmidae from Maharashtra, India, and the generic placement of Xenophthalmus garthii Sankarankutty, 1969 (Decapoda, Brachyura). Crustaceana 92, 1337–1348. doi:10.1163/15685403-00003947 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Serène R, Umali AF (1972) The family Raninidae and other new and rare species of brachyuran decapods from the Philippines and adjacent regions. Philippine Journal of Science 99, 21–105.

Pinnotheroidea De Haan, 1833 Pea crabs Figures 14.202–14.209 The pinnotheroids are here restricted to a single family Pinnotheridae. Ahyong and Ng (2009) and Davie et al. (2015)

tentatively included Aphanodactylidae in the pinnotheroids, but Tsang et al. (2018) showed them to be more closely allied to ocypodoids; aphanodactylids are treated here as a superfamily (Tsang et al. 2018). The pinnotherids are best known for the species commensal in bivalve molluscs, which have a smooth subglobular carapace, earning them the common name, pea crab. Although not all pinnotherids live inside bivalves are subglobular, most if not all are obligate commensals of other invertebrates (molluscs, echinoderms, ascidians, various worm phyla), either living inside the body of the host, or as ectocommensals within the host burrow (see de Gier and Becker 2020 for a detailed review). The obligate commensal (even parasitic) lifestyle of pinnotherids has resulted in reduction of body structures such as spines and other ornamentation, and modification for host occupation leading to a usually simplified subglobular habitus among those primarily infesting bivalve molluscs or the transversely slender, sometimes subcylindrical body form among those that dwell within the tubes of infaunal worms or other burrowing invertebrates. The more subglobular forms have traditionally been placed in the subfamily Pinnotherinae, and most of the more slender forms in Pinnotherelinae, Asthenognathinae and Xenophthalminae (Schmitt et  al. 1973). Commensalism with burrowing infauna, however, has occurred more than once in brachyurans, so similar adaptations have evolved independently in several lineages leading several unrelated groups to have been incorrectly placed in Pinnotheridae (Ahyong and Ng 2009; Ng et al. 2008). The asthenognathines are now placed in Varunidae (Grapsoidea) and the xenophthalmines form a family within Ocypodoidea (Ng et al. 2008). The status and composition of Pinnotherelinae has long been problematical. Palacios Theil et al. (2016) showed that the type genus, Pinnotherelia, is not a pinnotherid but a varunid, and referred most remaining pinnothereline genera to a new subfamily Pinnixulalinae and a reconstituted Pinnixinae, leaving the placement of Parapinnixa, Sakaina and Tetrias ambiguous. We herein formally place Pinnotherelinae (restricted to its type genus, Pinnotherelia) in Varunidae. Parapinnixa, Sakaina, Tetrias are retained as pinnotherids but not referable to any of the existing subfamilies, which themselves are probably not monophyletic (Tsang et al. 2018). Although the limits of Pinnotheridae are now well constrained, the phylogenetic structure within the family is insufficiently known to underpin a workable subfamily system for the group and further research is required to properly place the three ‘unaligned’ genera. Although the

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subfamily groupings remain useful monikers for discussion of broad morphological patterns, at present, they are not sufficiently defined for key purposes. Therefore, for convenience, we do not structure this section by subfamily. However, many of the former pinnothereline genera, namely Parapinnixa, Sakaina, Tetrias, and those assigned to Pinnixinae and Pinnixulalinae by Palacios Theil et  al. (2016) and Palacios Theil et  al. (2016) key out in couplets 1–13, and the pinnotherine genera key from couplet 14 onwards. The biology of only a few pinnotherids has been studied in detail (Castro 2015; Christensen and McDermott 1958) but as far as is known, most pinnotherids that live within the host (i.e. pinnotherines) show significant sexual dimorphism. Compared to males, adult females are usually considerably larger, have different carapace form, less robust chelipeds, more pronounced differences in pereopodal article proportions, different patterns of setation and weaker cuticle chitinisation. Conversely, pinnotherids that live within host burrows (i.e. primarily pinnixines, pinnixulalines, Tetrias) are usually less strongly sexually dimorphic and more heavily chitinised than pinnotherines.

Most pinnotherid species are currently known only from females, so the taxonomy of the group is largely based on female characters. Therefore, the features used in the key below refer to those of adult females except where otherwise indicated. Note that some species currently assigned to large genera such as Pinnixa and Pinnotheres are poorly known and after revision, may prove to belong to other genera. Diagnosis. Carapace cuticle thin, often soft, poorly calcified in females; carapace firm in males. Antennal gland opening on margin of buccal cavern. Maxilliped 3 ischium and merus fused, or ischium and merus free (Alarconia). Pereopods 2–5 propodi lower distal margin unarmed, without teeth opposing flexor margin of dactylus.

Pinnotheridae De Haan, 1833 Implicit generic attributes. Surface smooth. Maxilliped 3 ischium and merus fused; palp 3-articled; propodus longer than carpus; exopod without angular lobe on midlateral margin. Cheliped fingers distally pointed. Pereopods 2–5 bilaterally symmetrical, similarly slender; pereopodal dactyli with simple apex.

Key to females of genera of Pinnotheridae 1. – 2. – 3. – 4. – 5. – 6. – 7. – 8.

Maxilliped 3 dactylus articulating at end of propodus extending to base of ischiomerus when palp folded (Fig. 14.206m)������������������������������������������������������������������������������������������������������������������������������������������������������������������� Tetrias Maxilliped 3 dactylus, when present, articulating proximal to end of propodus or at end of propodus; if at end of propodus, when folded, extending at most to distal quarter of ischiomerus������������������������������������������������2 Maxilliped 3 ischium and merus separate (Fig. 14.205e)������������������������������������������������������������������������������������������� Alarconia Maxilliped 3 ischium and merus fused, sometimes demarcated by groove���������������������������������������������������������������������������3 Maxilliped 3 exopod with angular lobe on midlateral margin (e.g. Fig. 14.205f, r)��������������������������������������������������������������4 Maxilliped 3 exopod without angular lobe on midlateral margin (e.g. Fig. 14.206k)�������������������������������������������������������� 12 Carapace posterior surface near cardiac region fully traversed by single distinct continuous ridge (Fig. 14.203e)���������������������������������������������������������������������������������������������������������������������������������������������������������������� Austinixa Carapace cardiac ridge, if present, not fully traversing surface������������������������������������������������������������������������������������������������5 Maxilliped 3 dactylus and propodus oriented at wide angle to each other, often near perpendicular (Fig. 14.205r, u), or (in Glassella costaricana) dactylus strongly reduced, inserted subdistally��������������������������������������6 Maxilliped 3 dactylus and propodus oriented parallel or nearly parallel to each other�������������������������������������������������������7 Male pleon usually with gonopodal plate (Fig. 14.209d), if plate absent, pereopod 5 dactylus stout, length 3 times as long as wide or less. Western Atlantic, Eastern Pacific������������������������������������������������������������Glassella Male pleon without gonopodal plate. Pereopod 5 dactylus slender, 4 times as long as wide or greater (Fig. 14.208c, d). Indo-West Pacific����������������������������������������������������������������������������������������������������������������������Indopinnixa Mature cheliped fixed finger approximately in line with longitudinal axis of palm, at most, weakly deflected ventrally (Fig. 14.207d)����������������������������������������������������������������������������������������������������������������������������������������������8 Mature cheliped fixed finger strongly deflected ventrally from longitudinal axis of palm, often almost perpendicular����������������������������������������������������������������������������������������������������������������������������������������������������������������������������10 Pereopods 4, 5 meri and ischia lower margins usually with spinules or acute granules, if unarmed, male pleonites 3–5 fused into single plate (Fig. 14.209g, h)����������������������������������������������������������������������������������������Pinnixulala

14 – Brachyura – crabs

– 9. – 10. – 11. – 12. – 13. – 14. – 15. – 16. – 17. – 18. – 19. – 20. – 21. – 22. – 23. –

863

Pereopods 4, 5 meri and ischia lower margins smooth or with rounded granules. Male pleonites all free, or at least clearly demarcated by sutures����������������������������������������������������������������������������������������9 Carapace lateral margins bluntly obtuse, strongly setose (Fig. 14.203q)���������������������������������������������������������������������Pinnixa Carapace lateral margins rounded, without patches of setae (Fig. 14.203v)�����������������������������������������������������������Scleroplax Carapace regions scarcely defined; anterolateral margins smooth except for single lobiform tubercle just anterior to lateral extremity (Fig. 14.203u). Male telson subrectangular, more than twice as wide as long (Fig. 14.209j)�����������������������������������������������������������������������������������������������������������������������������������������������������������������������Sayixa Carapace regions well defined; anterolateral margins smooth or granular, without lobiform tubercle near lateral extreme. Male telson semicircular to subtriangular��������������������������������������������������������������������������������������11 Pereopods 2–4 similarly slender, compressed; dactyli long, slender, nearly as long as propodus. Male telson subtriangular to broadly rounded (Fig. 14.209i)���������������������������������������������������������������������������Rathbunixa Pereopods 2 and 3 slender, pereopod 4 swollen, markedly stouter, than pereopods 2, 3; dactyli stout, shorter than propodus (Fig. 14.204e). Male telson semicircular������������������������������������������������������������������������ Tubicolixa Pereopod 2 or pereopods 2 and 3 distinctly larger, more inflated, longer than remaining pereopods����������������������������13 Pereopods 2–4 of varying length but similarly slender; pereopod 2 never more inflated than other pereopods�����������������������������������������������������������������������������������������������������������������������������������������������������������������������������������14 Pereopod 2 larger than pereopod 3���������������������������������������������������������������������������������������������������������������������������Parapinnixa Pereopods 2, 3 subequal (Fig. 14.203t)����������������������������������������������������������������������������������������������������������������������������� Sakaina Pereopods 2–4 dactylus apex bifurcate, pereopod 5 dactylus simple (Fig. 14.207r, s)���������������������������������������������������������15 Pereopods 2–5 dactylus apex simple�������������������������������������������������������������������������������������������������������������������������������������������16 Maxilliped 3 palp stout, as long as or longer than maximum width of ischiomerus (Fig. 14.205n)��������������� Dissodactylus Maxilliped 3 palp slender, shorter than maximum width of ischiomerus (Fig. 14.205l)�������������������������Clypeasterophilus Maxilliped 3 palp 2-articled (e.g. Fig. 14.205h, m), or if dactylus present, as a minute nub, as long as wide, placed on lower subdistal margin (some species of Calyptraeotheres)�������������������������������������������������17 Maxilliped 3 palp 3-articled; dactylus, if inserted on lower subdistal margin, longer than wide (e.g. Figs 14.205o, 14.206f)������������������������������������������������������������������������������������������������������������������������������������������������������ 26 Carapace, at least anteriorly, with upturned margins or broad, thin, upraised, angled lamella (Fig. 14.202o, p, s)����������������������������������������������������������������������������������������������������������������������������������������������������������������������18 Carapace without upturned margins or anterior lamella��������������������������������������������������������������������������������������������������������19 Carapace transversely ovate, margins upturned, dorsal surface with prominent median mushroom-shaped tubercle (Fig. 14.202s)��������������������������������������������������������������������������������������������������������������Xanthasia Carapace pentagonal, dorsal surface with broad, upraised V-shaped anterior lamella, without median tubercle (Fig. 14.202o, p)�����������������������������������������������������������������������������������������������������������Serenotheres Maxilliped 3 propodus shorter than carpus (Fig. 14.205k)���������������������������������������������������������������������������������������������������� 20 Maxilliped 3 propodus longer than carpus (Figs 14.205m, 14.206b)�������������������������������������������������������������������������������������21 Carapace dorsally flattened, with 2 anterior longitudinal grooves. Pereopod 5 dactylus longer than pereopods 2–4 dactylus (Fig. 14.203f). Maxilliped 3 palp with or without minute nub-like dactylus on lower subdistal margin (Fig. 14.205k)������������������������������������������������������������������������������������ Calyptraeotheres Carapace subglobular, smooth, without longitudinal grooves (Fig. 14.203m). Pereopods 2–5 dactyli subequal. Maxilliped 3 palp without dactylus��������������������������������������������������������������������������������������������������� Nannotheres Carapace widest posteriorly. Cheliped fingers distally spatulate on occlusal surface (Fig. 14.207a). Pereopod 3 bilaterally asymmetrical (Fig. 14.202e)��������������������������������������������������������������������������������������� Discorsotheres Carapace widest at or near midlength. Cheliped fingers distally pointed (Fig. 14.207b). Pereopod 3 bilaterally symmetrical��������������������������������������������������������������������������������������������������������������������������������������� 22 Carapace subpentagonal, angled anteriorly (Fig. 14.204f)����������������������������������������������������������������������������������� Tunicotheres Carapace rounded or transversely ovate to subrectangular��������������������������������������������������������������������������������������������������� 23 Carapace wider than long, transversely ovate to subrectangular (Fig. 14.203k)��������������������������������������������������� Latatheres Carapace about as long as wide, subcircular����������������������������������������������������������������������������������������������������������������������������� 24

864

Marine Decapod Crustacea

24. Carapace soft, weakly sclerotised, anterolateral margins undefined. Eyes not visible in dorsal view (Fig. 14.203h). Maxilliped 3 propodus tapering to narrow but blunt point. Pereopod 3 dactylus bilaterally asymmetrical (Fig. 14.208a, b)���������������������������������������������������������������������������������������������������������Gemmotheres – Carapace firm to hard, anterolateral margins well defined. Eyes visible in dorsal view. Maxilliped 3 propodus spatulate, apex blunt, subtruncate to rounded (Fig. 14.205h, 14.206g). Pereopod 3 dactylus bilaterally symmetrical (Fig. 14.202b)����������������������������������������������������������������������������������������������������������������������������������� 25 25. Carapace dorsal surface glabrous (Fig. 14.202b). Gonopod 1 with lanceolate tip (Fig. 14.209k)�������������������� Austrotheres

Fig. 14.202.  Pinnotheridae. a, Austinotheres angelicus (Lockington, 1877); b, Austrotheres pregenzeri Ahyong, 2018; c, Bonita mexicana Campos, 2009; d, Buergeres deccanensis (Chopra, 1931); e, Discorsotheres subquadratus (Sakai, 1939); f, g, Durckheimia caeca Bürger, 1895 (lateral, dorsal views); h, Holotheres danielae Ahyong, 2010; i, Magnotheres globosus (Hombron & Jacquinot, 1846); j, Mesotheres strombi (Rathbun, 1905); k, Ostracotheres tridacnae (Rüppell, 1830); l, Plenotheres coarctatus (Bürger, 1895); m, Pinnaxodes chilensis (H. Milne Edwards, 1837); n, Pinnotheres hickmani (Guiler, 1950); o, p, Serenotheres besutensis (Serène, 1967) (dorsal, anterior views); q, Trichobezoares villosissimus (Doflein, 1904); r, Tridacnatheres whitei (De Man, 1888); s, Xanthasia murigera White, 1846.

14 – Brachyura – crabs

– 26. – 27. – 28. – 29. – 30. – 31.

865

Carapace dorsal surface finely setose (Fig. 14.202k). Gonopod 1 tip blunt, with short spine-like, anterolaterally directed papilla (Fig. 14.209n)��������������������������������������������������������������������������������������������������Ostracotheres Carapace with prominent subtriangular rostrum and 3 longitudinal epigastric grooves (Fig. 14.203l)��������������Limotheres Carapace without prominent rostrum, without longitudinal postfrontal grooves������������������������������������������������������������ 27 Carapace with long thin median ridge; margins with thin upturned rims, anterior margin with median fissure or notch (Fig. 14.202f, g)���������������������������������������������������������������������������������������������������Durckheimia Carapace margins blunt, not forming thin, upturned rim, without anterior median notch or fissure; surface smooth or variously ornamented, but without long, thin median ridge����������������������������������������������������������� 28 Carapace with central peak, either as large tubercle or short ridge�������������������������������������������������������������������������������������� 29 Carapace without median tubercle or ridge������������������������������������������������������������������������������������������������������������������������������ 30 Carapace margins with blunt rounded rims; surface with large median tubercle (Fig. 14.202r)������������������ Tridacnatheres Carapace margins with well defined edges; surface with short, low median ridge (Fig. 14.204h, i)�����������������������Visayeres Maxilliped 3 propodus shorter than carpus������������������������������������������������������������������������������������������������������������������������������31 Maxilliped 3 propodus as long as or longer than carpus�������������������������������������������������������������������������������������������������������� 36 Pereopods 2–5 stout, pereopod 3 merus length exceeding 3 times width; dactyli subequal����������������������������������������������32

Fig. 14.203.  Pinnotheridae. Carapace, dorsal unless noted: a, Alain; b, Alainotheres; c, Alarconia; d, Amusiotheres; e, Austinixa; f, Calyptraeotheres; g, Fabia; h, Gemmotheres; i, Glassella, with pereopods; j, Holothuriophilus; k, Latatheres; l, Limotheres; m, Nannotheres, anterior; n, Nepinnotheres; o, Orthotheres; p, Parapinnixa; q, Pinnixa; r, Rathbunixa; s, Raytheres; t, Sakaina, with pereopods; u, Sayixa; v, Scleroplax.

866

Marine Decapod Crustacea



Pereopods 2–5 slender, pereopod 3 merus length exceeding 5 times width; pereopod 3 or 4 dactylus longer than dactyli of other walking legs������������������������������������������������������������������������������������������������������������������������������������������33 32. Maxilliped 3 dactylus inserting at distal end of propodus (Fig. 14.206c). Setae of pereopods and carapace margins not obscuring body outline (Fig. 14.202j). Cheliped fingers distally pointed (Fig. 14.207c)������������Mesotheres – Maxilliped 3 dactylus inserting on proximal half of propodus (Fig. 14.206n). Pereopods and carapace densely covered with ‘shaggy’ setae obscuring body outline (Fig. 14.202q). Cheliped fingers distally spatulate on occlusal surface (Fig. 14.207g)��������������������������������������������������������������������������������������������������� Trichobezoares

Fig. 14.204.  Pinnotheridae. Carapace, dorsal unless noted: a, Sindheres; b, Solenotheres; c, Tacitotheres; d, Tetrias; e, Tubicolixa, with pereopods; f, Tunicotheres; g, Viridotheres; h, i, Visayeres (lateral, anterior views); j, Waldotheres.

Fig. 14.205.  Pinnotheridae. Maxilliped 3: a, Abyssotheres; b, Afropinnotheres; c, Alain; d, Alainotheres; e, Alarconia; f, Austinixa; g, Austinotheres; h, Austrotheres; i, Bonita; j, Buergeres; k, Calyptraeotheres; l, Clypeasterophilus; m, Discorsotheres; n, Dissodactylus; o, Durckheimia; p, Enigmatheres; q, Fabia; r, Glassella; s, Holotheres; t, Holothuriophilus; u, Indopinnixa.

14 – Brachyura – crabs

33. –

867

Pereopod 4 longest of walking legs, its dactylus longer than other walking leg dactyli. Carapace with pair of deep longitudinal grooves anteriorly (Fig. 14.202c)������������������������������������������������������������������������������������������������ Bonita Pereopod 3 longest of walking legs, its dactylus longer than dactyli of other walking legs. Carapace without pair of anterior grooves�������������������������������������������������������������������������������������������������������������������������� 34

Fig. 14.206.  Pinnotheridae. Maxilliped 3: a, Juxtafabia; b, Latatheres; c, Mesotheres; d, Nepinnotheres; e, Opisthopus; f, Orthotheres; g, Ostracotheres; h, Pinnaxodes; i, Pinnotheres; j, Rathbunixa; k, Sakaina; l, Sayixa; m, Tetrias; n, Trichobezoares; o, Xanthasia.

Fig. 14.207.  Pinnotheridae. Pereopod 1 chelae: a, Discorsotheres; b, Latatheres; c, Mesotheres; d, Pinnixulala; e, Rathbunixa; f, Sayixa ; g, Trichobezoares. Pereopods (p): h, p5, Alain; i, j, p2, p3, Amusiotheres; k, p5, Arcotheres; l–o, p2–p5, Bonita; p, q, p4, p5, Buergeres; r, s, p4, p5, Dissodactylus; t–w, right p2–p5, Enigmatheres; x, y, p3, p4, Ernestotheres.

868

34. – 35. – 36. – 37. – 38. – 39. – 40. – 41. – 42. – 43. – 44. – 45. – 46. – 47. – 48. – 49. –

Marine Decapod Crustacea

Maxilliped 3 dactylus articulating at subdistal third of propodus (Fig. 14.205p)��������������������������������������������Enigmatheres Maxilliped 3 dactylus articulating at distal end of propodus (Fig. 14.205a, g)��������������������������������������������������������������������35 Pereopod 3 bilaterally symmetrical in length��������������������������������������������������������������������������������������������������������� Abyssotheres Pereopod 3 bilaterally asymmetrical (longer pereopod 1.1–1.2 times as long as shorter pereopod; Fig. 14.202a)�����������������������������������������������������������������������������������������������������������������������������������������������������������Austinotheres Maxilliped 3 dactylus inserting at or near tip of propodus (Fig. 14.206f)����������������������������������������������������������������������������37 Maxilliped 3 dactylus inserting on lower margin of propodus, near or proximal to midlength (Figs 14.205t, 14.206i)�������������������������������������������������������������������������������������������������������������������������������������������������������������� 38 Carapace transversely ovate (Fig. 14.203o). Pereopods 2–5 dactyli with lower surface hollowed (Fig. 14.208e, f)���������������������������������������������������������������������������������������������������������������������������������������������������������Orthotheres Carapace subcircular (Fig. 14.204c). Pereopods 2–5 dactyli lower surface entire, subcircular in crosssection����������������������������������������������������������������������������������������������������������������������������������������������������������������������� Tacitotheres Carapace with 2 longitudinal grooves anteriorly (Figs 14.202l, 14.203g)����������������������������������������������������������������������������� 39 Carapace without 2 longitudinal grooves anteriorly��������������������������������������������������������������������������������������������������������������� 40 Pereopods 2–5 bilaterally symmetrical, or pereopod 3 asymmetrical; pereopods 2 and 5 dactyli subequal��������������� Fabia Pereopod 4 bilaterally asymmetrical, longest of walking legs; pereopod 5 dactylus about twice as long as pereopod 2 dactylus (Fig. 14.202l)��������������������������������������������������������������������������������������������������������������������������Plenotheres Maxilliped 3 dactylus broadly spatulate (Fig. 14.205s, t, 14.206e, h, j)����������������������������������������������������������������������������������41 Maxilliped 3 dactylus digitiform (Figs 14.205c, 14.206a)������������������������������������������������������������������������������������������������������� 46 Maxilliped 3 dactylus inserting on lower proximal margin of propodus, nearly at base (Fig. 14.206e)������������������������� 42 Maxilliped 3 dactylus inserting near midlength of propodus (Fig. 14.206h)���������������������������������������������������������������������� 44 Maxilliped 3 ischium and merus demarcated by groove (Fig. 14.206e)����������������������������������������������������������������Opisthopus Maxilliped 3 ischium and merus fused (Fig. 14.205s, t)���������������������������������������������������������������������������������������������������������� 43 Carapace transversely rectangular to broadly ovate, wider than long (Fig. 14.203j)���������������������������������Holothuriophilus Carapace subcircular, about as long as wide (Fig. 14.202h)��������������������������������������������������������������������������������������Holotheres Maxilliped 3 dactylus reaching or overreaching end of propodus (Fig. 14.206h). Pereopods 2–5 dactyli subequal, or pereopod 5 dactylus shorter than others (Fig. 14.202m)������������������������������������������������������������� Pinnaxodes Maxilliped 3 dactylus not reaching beyond end of propodus (Fig. 14.205j). Pereopod 5 dactylus longer than pereopod 2 and 4 dactyli (and usually also that of pereopod 3)��������������������������������������������������������������������������������������� 45 Carapace surface glabrous. Pereopod 5 dactylus 2–3 times length of pereopods 2–4 dactyli (Fig. 14.202d)����������������������������������������������������������������������������������������������������������������������������������������������������������������Buergeres Carapace surface densely, finely setose. Pereopod 5 dactylus up to ~1.5 times length of pereopods 2 and 4 dactyli (and usually also that of pereopod 3) (Fig. 14.208n, o)��������������������������������������������������������Tumidotheres Pereopod 5 dactylus longer than pereopod 2 dactylus������������������������������������������������������������������������������������������������������������ 47 Pereopod 5 dactylus shorter than or as long as pereopod 2 dactylus�������������������������������������������������������������������������������������51 Pereopod 2 propodus expanded distally. Pereopod 3 dactylus longer than pereopod 5 dactylus (Fig. 14.208r–u)������������������������������������������������������������������������������������������������������������������������������������������������������������������� Zaops Pereopod 2 propodus not expanded distally. Pereopod 3 dactylus shorter than pereopod 5 dactylus������������������������������ 48 Pereopod 4 bilaterally symmetrical. Pereopod 5 ischium and merus with line of articulation diagonal to longitudinal axis (Fig. 14.207h)��������������������������������������������������������������������������������������������������������������������������������������������� 49 Pereopod 4 bilaterally asymmetrical. Pereopod 5 ischium and merus with line of articulation perpendicular to longitudinal axis (Fig. 14.207k)��������������������������������������������������������������������������������������������������������������� 50 Maxilliped 3 dactylus inserted slightly proximal to midlength of propodus (Fig. 14.205c). Male pleon elongate-pentagonal, widest at pleonite 5 and 6, all pleonites free (Fig. 14.209a)�����������������������������������������������������Alain Maxilliped 3 dactylus inserted near base of propodus (Fig. 14.206a). Male pleon narrowly triangular, widest at pleonite 3, abruptly narrowing distally, pleonite 4 and 5 fused (Fig. 14.209e)�������������������������������� Juxtafabia

14 – Brachyura – crabs

869

50. Pereopod 4 usually distinctly asymmetrical, dactyli usually distinctly unequal in length. Male telson semicircular (Fig. 14.209b) to transversely ovate, widest basally or at midlength������������������������������������������� Arcotheres – Pereopod 4 weakly asymmetrical, dactyli subequal (Fig. 14.202i). Male telson trapezoidal, widest distally (Fig. 14.209f)�������������������������������������������������������������������������������������������������������������������������������� Magnotheres 51. Pereopod 3 dactylus of one or both sides longer than pereopod 4 dactylus�������������������������������������������������������������������������52 – Pereopod 3 dactylus subequal to pereopod 4 dactylus������������������������������������������������������������������������������������������������������������ 56 52. Carapace transversely rectangular, ~1.5 times as wide as long (Figs 14.203s, 14.204b)������������������������������������������������������53 – Carapace subcircular to subhexagonal, less than 1.3 times as wide as long������������������������������������������������������������������������ 54 53. Pereopods 2, 3 propodi distally swollen; dactyli widest near midlength. Pereopods 3 left-right equal in length (Fig. 14.208j, k)����������������������������������������������������������������������������������������������������������������������������������Raytheres – Pereopods 2, 3 propodi not distally swollen; dactyli widest proximally, tapering. Pereopods 3 left-right unequal in length (Fig. 14.208l, m)���������������������������������������������������������������������������������������������������Solenotheres 54. Carapace subhexagonal, widest behind midlength (Fig. 14.204j)������������������������������������������������������������������������Waldotheres – Carapace subcircular, widest at midlength��������������������������������������������������������������������������������������������������������������������������������55 55. Carapace subsquare to rounded, anterior margin transverse (Fig. 14.203d). Pereopod 3 dactylus subequal to or shorter than pereopods 4, 5 dactyli (Fig. 14.207i, j)��������������������������������������������������������������Amusiotheres – Carapace subcircular to transversely ovate, anterior margin rounded (Fig. 14.204g). Pereopod 3 dactylus longer than pereopods 4, 5 dactyli (Fig. 14.208p, q)�����������������������������������������������������������������������Viridotheres

Fig. 14.208.  Pinnotheridae. Pereopods (p): a, b, left and right p3, Gemmotheres; c, d, p4, p5, Indopinnixa; e, f, p3, with dactylus, Orthotheres; g, h, p4, p5, Pinnixa; i, p4, Pinnixulala; j, k, p2, p3, Raytheres; l, m, right and left p3, Solenotheres; n, o, p4, p5, Tumidotheres; p, q, p3, p4, Viridotheres; r–u, p2–p5, Zaops.

870

Marine Decapod Crustacea

Fig. 14.209.  Pinnotheridae. Male pleon: a, Alain; b, Arcotheres; c, Ernestotheres; d, Glassella (ventral); e, Juxtafabia; f, Magnotheres; g, h, Pinnixulala; i, Rathbunixa; j, Sayixa. Gonopod 1: k, Austrotheres; l, Hospitotheres; m, Nepinnotheres; n, Ostracotheres.

56. – 57. – 58. – 59. – 60. – 61. –

Maxilliped 3 dactylus inserted proximal to midlength of propodus (Figs 14.205b, 14.206i)���������������������������������������������57 Maxilliped 3 dactylus inserted near midlength of propodus (Figs 14.205d, 14.206d)������������������������������������������������������� 58 Maxilliped 3 dactylus overreaching end of propodus; propodus conical (Fig. 14.205b)��������������������������� Afropinnotheres Maxilliped 3 dactylus not reaching end of propodus; propodus flattened (Fig. 14.206i)����������������������������������Pinnotheres Carapace subrectangular or trapezoidal, distinctly wider than long�������������������������������������������������������������������������������������59 Carapace subcircular or subsquare, about as wide as long (Fig. 14.203n)���������������������������������������������������������������������������� 60 Carapace trapezoidal, widest posteriorly (Fig. 14.204a)����������������������������������������������������������������������������������������������Sindheres Carapace subrectangular (Fig. 14.203b)������������������������������������������������������������������������������������������������������������������� Alainotheres Pereopods 2–5 stout in females (Fig. 14.207x, y), pereopod 3 shorter than carapace length. Male pleonites 5 and 6 fused (Fig. 14.209c)������������������������������������������������������������������������������������������������������� Ernestotheres Pereopods 2–5 slender in females, pereopod 3 longer than carapace length. Male pleonites and telson all free����������������������������������������������������������������������������������������������������������������������������������������������������������������������61 Pereopods 2–5 propodus in females clavate, distally slightly expanded. Gonopod 1 distally kinked (Fig. 14.209l)�������������������������������������������������������������������������������������������������������������������������������Hospitotheres Pereopods 2–5 propodus in females upper and lower margins parallel or slightly convergent distally, not expanded. Gonopod 1 straight or curved to approximately a right-angle (Fig. 14.209m)������������������ Nepinnotheres Abyssotheres Manning & Galil, 2000

Diagnosis. Carapace of female subcircular, little wider than long. Maxilliped 3 dactylus digitiform, articulating at distal end of propodus. Pereopod 3 longest of walking legs; merus 5 times as long as wide. Maximum cl. 11 mm. Slope (722–787 m, in bivalve molluscs (Acesta spp.). Western and Central Indo-Pacific. 2 species (Komatsu and Ohtsuka 2009: rediagnosis, species).

Afropinnotheres Manning, 1993 Diagnosis. Carapace of female subcircular, little wider than long. Maxilliped 3 propodus as long as or slightly longer than carpus, conical; dactylus broadly digitiform, inserting near base of and

overreaching end of propodus. Pereopod 3 or pereopod 4 longest of walking legs. Pereopods 2–5 dactyli subequal. Maximum cl. 12.1 mm. Intertidal, subtidal (0–19 m; in bivalves). Tropical E Atlantic, Western Indo-Pacific, Temperate Southern Africa. 6 species (Ahyong 2019; Manning 1993b: diagnosis, key to 3 species; Ng and Kumar 2015: discussion, distribution).

Alain Manning, 1998 Diagnosis. Carapace of female subcircular, about as wide as long; surface simple, hard, strongly setose. Maxilliped 3 dactylus digitiform, inserting slightly proximal to midlength of propodus. Pereopods 4, 5 dactyli longer than pereopod 2, 3 dactyli. Pereopod 3

14 – Brachyura – crabs

longest of walking legs; pereopod 5 ischium and merus with line of articulation diagonal to longitudinal axis. Male pleon elongatepentagonal, widest at pleonites 5 and 6, all pleonites free. Maximum cl. 12 mm. Slope (399–787 m, in holothurians). Central Indo-Pacific. 2 species (Ahyong and Ng 2008: rediagnosis, species comparison; Manning 1998).

Alainotheres Manning, 1993 Diagnosis. Carapace subrectangular, wider than long. Maxilliped 3 dactylus digitiform, inserting near midlength of propodus, not reaching end of propodus. Pereopods 2–5 bilaterally symmetrical, dactyli subequal; stout in male. Pereopod 3 longest of walking legs. Maximum cl. 2.8 mm (male only known). Subtidal (20 m). Tropical E Atlantic. 1 species (Manning 1993b).

Alarconia Glassell, 1938 Diagnosis. Carapace of female transversely ovate-fusiform, 1.6 times as wide as long; surface regions well defined, cardiac ridge, not fully traversing carapace width. Maxilliped 3 ischium and merus separate; dactylus articulating proximal to midlength of propodus, oriented nearly parallel to each other. Pereopod 4 markedly stouter, longer than pereopods 3 and 5; pereopods 4, 5 merus and ischium lower margins with spinules or acute granules. Maximum cl. 7 mm. Subtidal (11–25 m). Tropical W Atlantic, Eastern Indo-Pacific. 2 species (Coelho 1996; Glassell 1938).

Amusiotheres Ng & Ho, 2016 Diagnosis. Carapace of female transversely subquadrate, or subcircular, little wider than long; surface irregular. Maxilliped 3 dactylus digitiform, articulating slightly proximal to midlength of propodus. Pereopods 3 asymmetrical; right pereopod 3 dactylus longest. Pereopod 3 longest of walking legs. Maximum cl. 11 mm. Subtidal (in bivalve, Amusium). Western and Central IndoPacific. 2 species (Ng and Ho 2016a).

Arcotheres Manning, 1993 Diagnosis. Carapace of female irregularly subquadrate to subhexagonal or transversely ovate, as long as wide to wider than long. Maxilliped 3 dactylus digitiform, inserting near midlength of propodus. Pereopods 2–5 similarly slender; pereopod 4 or 5 dactylus elongated, linear, both longer than falcate pereopod 2 and 3 dactyli. Pereopod 4 longest of walking legs, usually strongly bilaterally asymmetrical, dactyli usually distinctly unequal in length; pereopod 5 ischium and merus with line of articulation perpendicular to longitudinal axis. Male telson semicircular to transversely ovate, widest basally or at midlength. Maximum cl. 10 mm. Intertidal, subtidal (0–18 m, in bivalve molluscs). Western and Central Indo-Pacific. 23 species (Ahyong and Ng 2020, 2021: recent species; Komai et al. 2020: recent species; Manning 1993a; Ng and Ahyong 2022: generic attributes, species descriptions, figures).

871

Austinixa Heard & Manning, 1997 Diagnosis. Carapace of female transversely subelliptical, 1.8–3.3 times as wide as long; surface regions weakly-defined, posterior surface near cardiac region fully traversed by single distinct continuous ridge. Maxilliped 3 dactylus articulating proximal to midlength of propodus, oriented nearly parallel to each other; exopod with angular lobe on midlateral margin. Pereopod 4 markedly stouter, longer than pereopods 3 and 5. Maximum cl. 8 mm. Intertidal–shelf (0–130 m, associated with burrowing callianassid ghost shrimps and tubicolous polychaetes). Tropical W Atlantic, Eastern Indo-Pacific. 14 species (Heard and Manning 1997; Palacios Theil and Felder 2020a: key to 12 species; SalgadoBarragán et  al. 2021: key to 13 species). Pinnixa transversalis (H. Milne Edwards & Lucas, 1842) is herein included owing to its wide transverse carapace ridge.

Austinotheres Campos, 2002 Diagnosis. Carapace of female transversely rounded-subquadrate, 1.3–1.4 times as wide as long. Maxilliped 3 propodus shorter than carpus; dactylus digitiform, inserting on lower subdistal margin. Pereopods 2–5 similarly slender. Pereopod 3 longest of walking legs; pereopod 3 merus more than 5 times as long as wide; longer pereopod 3 1.1–1.2 times as long as shorter. Maximum cl. 9 mm. Intertidal (in bivalve molluscs, Ostreidae). Tropical Eastern Pacific. 1 species (Campos 2002).

Austrotheres Ahyong, 2018 Diagnosis. Carapace of female about as long as wide; surface glabrous, firm to hard, anterolateral margins well defined. Maxilliped 3 palp 2-articled; propodus longer than carpus, spatulate, apex subtruncate to rounded. Pereopodal dactyli bilaterally symmetrical, subequal or pereopod 5 dactylus the longest. Pereopod 2 or pereopod 3 longest of walking legs. Gonopod 1 tip lanceolate. Maximum cl. 11 mm. Subtidal, shelf (3–92 m, in ascidians (Herdmania sp.) and holothurians). Temperate Australia. 2 species (Ahyong 2018: diagnosis, key to species).

Bonita Campos, 2009 Diagnosis. Carapace of female subhexagonal, wider than long; surface soft, weakly sclerotised, with pair of deep longitudinal grooves anteriorly. Maxilliped 3 propodus shorter than carpus; dactylus digitiform, articulating at subdistal one-third of propodus. Pereopods 2–5 similarly slender. Pereopods 4 bilaterally asymmetrical, longer one being longest of walking legs, its dactylus longer than other walking leg dactyli; pereopod 3 merus more than 5 times as long as wide; pereopod 5 propodus clavate. Maximum cl. 6 mm. Intertidal (in bivalve molluscs, Pseudochama exogyra). Tropical Eastern Pacific (Mexico). 1 species (Campos 2009).

Buergeres Ng & Manning, 2003 Diagnosis. Carapace of female subcircular to rounded-­ subquadrate, little wider than long. Maxilliped 3 dactylus broadly

872

Marine Decapod Crustacea

spatulate, inserting near midlength, not overreaching end of propodus. Pereopod 5 dactylus 2–3 times length of pereopods 2–4 dactyli. Maximum cl. 21 mm. Intertidal, subtidal (0–17 m, in holothurians). Western and Central Indo-Pacific. 4 species (Ahyong and Ng 2020: key to species; Ng and Manning 2003).

Calyptraeotheres Campos, 1990 Diagnosis. Carapace of female dorsally flattened, slightly wider than long; surface with 2 anterior longitudinal grooves. Maxilliped 3 palp 2- or 3-articled; propodus shorter than carpus; dactylus minute, nub-like, on lower subdistal margin of propodus, or absent. Pereopods 2–4 similarly slender. Pereopod 4 longest pereopod; pereopod 5 dactylus longer than pereopods 2–4 dactyli. Maximum cl. 6 mm. Intertidal, subtidal (0–6 m, in gastropod molluscs, Calyptraeidea). Tropical W Atlantic, Tropical Eastern Pacific. 6 species (Ayón-Parente and Hendrickx 2014: key to species; Campos 1990).

Clypeasterophilus Campos & Griffith, 1990 Diagnosis. Carapace of female subpentagonal, 1.2–1.6 times as wide as long. Maxilliped 3 palp slender, shorter than maximum width of ischiomerus. Pereopods 2–4 similarly slender; pereopods 2–4 dactyli apices bifurcate, pereopod 5 dactylus simple. Pereopod 3 or pereopod 4 longest of walking legs. Maximum cl. 7 mm. Subtidal–slope (2–281 m, on echinoids, Clypeaster sp.). Tropical W Atlantic, Tropical Eastern Pacific. 4 species. The four species now in Clypeasterophilus were previously in Dissodactylus (Campos and Griffith 1990) (Campos and Griffith 1990; Griffith 1987: key to species as Dissodactylus, couplet 2).

Discorsotheres Ahyong, 2018 Diagnosis. Carapace of female rounded-subquadrate, little wider than long, margins blunt, widest posteriorly. Maxilliped 3 palp 2-articled. Cheliped fingers distally spatulate on occlusal surface. Pereopodal dactyli falcate, apex simple. Pereopod 3 bilaterally asymmetrical, longest of walking legs. Maximum cl. 15 mm. Subtidal–slope (6–45 m, in bivalve molluscs). Western and Central Indo-Pacific. 4 species (Ahyong 2018: diagnosis, key to species).

Dissodactylus Smith, 1870 Diagnosis. Carapace of female subpentagonal, 1.1–1.8 times as wide as long. Maxilliped 3 palp stout, as long as or longer than maximum width of ischiomerus. Pereopods 2–4 similarly slender; pereopodal dactyli apex bifurcate, pereopod 5 dactylus simple. Pereopod 3 or pereopod 4 longest of walking legs. Maximum cl. 7 mm. Intertidal–shelf (0–73 m, on echinoids). Tropical W Atlantic. 9 species (Griffith 1987: key to species includes at couplet 2, 4 species now in Clypeasterophilus). WoRMS lists 15 species, but six are unidentifiable larval names of Aikawa (1933).

Durckheimia De Man, 1889 Diagnosis. Carapace of female ovate to trapezoid, slightly wider than long; surface with long thin median ridge; margins with thin upturned rims, anterior margin with median fissure or notch. Maxilliped 3 dactylus digitiform, inserting on lower subdistal margin. Pereopods 2–5 similarly slender; pereopodal dactyli falcate, apex simple. Pereopod 3 or pereopod 4 longest of walking legs. Maximum cl. 9 mm. Subtidal (9–11 m; associated with bivalves, Limidae). Western and Central Indo-Pacific. 3 species (Ahyong and Ng 2005: rediagnosis, key to species).

Enigmatheres Campos, 2009 Diagnosis. Carapace of female subcircular, little wider than long; surface soft, weakly sclerotized. Maxilliped 3 propodus shorter than carpus; dactylus digitiform, articulating at subdistal onethird of propodus. Pereopods 2–5 similarly slender. Pereopods 3 bilaterally asymmetrical, longer of these the longest of walking legs, its dactylus longer than other walking leg dactyli; pereopod 3 merus more than 5 times as long as wide. Maximum cl. 6 mm. Subtidal (on gastropod mollusc, Megathura crenulata). Temperate Northern Atlantic (California). 1 species (Campos 2009).

Ernestotheres Manning, 1993 Diagnosis. Carapace of female subcircular, little wider than long. Maxilliped 3 propodus longer than carpus, spatulate, apex subtruncate to rounded; dactylus digitiform, inserting near midlength of propodus, not reaching end of propodus. Pereopods 2–5 stout, dactyli subequal. Pereopod 3 longest of walking legs, shorter than carapace length. Male pleonites 5–6 fused. Maximum cl. 15 mm. Intertidal (on gastropods, Conus). Tropical E Atlantic. 1 species (Manning 1993b: diagnosis; Manning and Holthuis 1981: species description)

Fabia Dana, 1851 Diagnosis. Carapace of female subcircular, little wider than long or about as wide as long; surface with 2 anterior longitudinal grooves. Maxilliped 3 propodus longer than carpus or as long as carpus; dactylus digitiform, inserting near midlength of propodus. Pereopods 2–5 similarly slender, bilaterally symmetrical or 3 asymmetrical; pereopodal dactyli falcate, apex simple, pereopod 2 and 5 dactyli subequal. Pereopod 3 longest of walking legs. Maximum cl. 10 mm. Intertidal–shelf (0–80 m, in bivalve molluscs). Tropical W Atlantic, Tropical Eastern Pacific. 9 species (Campos 1996b: rediagnosis, species diagnoses; Campos 2013: species separation; Gore 1986: species comparison).

Gemmotheres Campos, 1996 Diagnosis. Carapace of female subcircular, about as wide as long; surface soft, weakly sclerotised, surface simple, anterolateral margins undefined; eyes not visible in dorsal view. Maxilliped 3 palp

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2-articled; propodus longer than carpus, tapering to narrow, blunt point. Pereopods 2–5 similarly slender; pereopod 3 dactylus bilaterally asymmetrical, pereopod 5 dactylus longer than pereopod 2–4 dactyli. Maximum cl. 7 mm. Subtidal (in bivalve molluscs, Chama congregata). Tropical W Atlantic. 1 species (Campos 1996a).

Glassella Campos & Wicksten, 1997 Diagnosis. Carapace of female transversely subelliptical, 2–3 times as wide as long; surface regions weakly defined, cardiac ridge, if present, obsolescent, not fully traversing carapace width. Maxilliped 3 dactylus strongly reduced, inserted subdistally (in G.  ­costaricana), or and propodus oriented at wide angle to each other, often near perpendicular; exopod with angular lobe on midlateral margin. Pereopod 4 markedly stouter, longer than pereopods 3 and 5; pereopod 5 dactylus stout, less than 3 times as long as wide (if gonopodal plate absent). Male pleon usually with gonopodal plate. Maximum cl. 6 mm. Intertidal–shelf (0–55 m, associated with tubicolous polychaetes, holothurians and sipunculans). Tropical W Atlantic, Tropical Eastern Pacific. 11 species (Felder and Palacios Theil 2020: key to 9 W Atlantic species; Palacios Theil and Felder 2020b: rediagnosis).

Holotheres Ng & Manning, 2003 Diagnosis. Carapace of female subcircular, about as wide as long. Maxilliped 3 dactylus broadly spatulate, inserting near base of propodus. Pereopods 2–5 bilaterally symmetrical, densely setose. Maximum cl. 10 mm. Intertidal, subtidal (in holothurians). Central Indo-Pacific. 5 species (Ahyong 2010: key to species; Ng and Manning 2003).

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Indopinnixa Manning & Morton, 1987 Diagnosis. Carapace of female transversely ovate-fusiform, 1.6–2.4 times as wide as long; surface regions weakly defined, cardiac ridge, when present, distinct but not fully traversing carapace width. Maxilliped 3 dactylus and propodus oriented at wide angle to each other, often near perpendicular; exopod with angular lobe on midlateral margin. Pereopod 5 dactylus slender, 4 or more times as long as wide. Pereopod 4 markedly stouter, longer than pereopods 3 and 5. Male telson about twice as wide as long, wider than pleonite 6. Maximum cl. 3 mm. Intertidal (associated with sipunculans and possibly other tubicolous WoRMS). Central Indo-Pacific. 8 species (Manning and Morton 1987; Naruse and Maenosono 2012; Komai et al. 2022).

Juxtafabia Campos, 1993 Diagnosis. Carapace of female subcircular, about as wide as long; surface simple, weakly sclerotized, glabrous or sparsely setose. Maxilliped 3 dactylus digitiform, inserting near base of propodus. Pereopod 5 dactylus longer than pereopod 2–4 dactyli. Pereopod 5 ischium and merus with line of articulation diagonal to longitudinal axis. Male pleon narrowly triangular, widest at pleonite 3, abruptly narrowing distally, pleonites 4 and 5 fused. Maximum cl. 7 mm. Subtidal (in bivalve molluscs). Tropical Eastern Pacific. 1 species (Campos 1993).

Latatheres Ahyong, 2018 Diagnosis. Carapace of female transversely ovate to subrectangular, 1.2–1.3 times as wide as long, widest at midpoint. Maxilliped 3 palp 2-articled. Pereopod 2 or pereopod 3 longest of walking legs. Maximum cl. 7 mm. Subtidal (in bivalve molluscs). Western and Central IndoPacific. 2 species (Ahyong 2018: diagnosis, key to species).

Holothuriophilus Nauck, 1880 Diagnosis. Carapace of female transversely rectangular to subovate, 1.2–1.6 times as wide as long. Maxilliped 3 dactylus broadly spatulate, inserting near base of propodus. Pereopods 2–5 bilaterally symmetrical, densely setose. Maximum cl. 12 mm. Subtidal (in holothurians). Tropical Eastern Pacific. 2 species (Ahyong and Ng 2007a: diagnosis of type species).

Hospitotheres Manning, 1993 Diagnosis. Carapace of female subcircular, little wider than long. Maxilliped 3 dactylus digitiform, inserting near midlength of propodus, not reaching end of propodus. Pereopods 2–5 propodus in females clavate, distally slightly expanded in females, dactyli subequal. Pereopod 3 longest of walking legs, longer than carapace length. Male pleonites and telson free. Gonopod 1 distally kinked. Maximum cl. 6.5 mm. Estuarine (in burrows with alpheid shrimps and callichirid ghost shrimps, but specific host unknown, possibly galeommatoid bivalves (de Gier and Becker 2020)). Tropical E Atlantic. 1 species (Manning 1993b)

Limotheres Holthuis, 1975 Diagnosis. Carapace of female subhexagonal, with prominent subtriangular rostrum; surface with 3 longitudinal epigastric grooves. Maxilliped 3 dactylus digitiform, articulating at end of propodus. Pereopodal dactyli falcate, apex simple. Pereopod 3 longest of walking legs. Maximum cl. 4 mm. Subtidal (10 m, in bivalve molluscs, Lima). Tropical Atlantic (Caribbean Sea). 1 species (Holthuis 1975).

Magnotheres Ng & Ahyong, 2022 Diagnosis. Carapace of female irregularly rounded-subquadrate to ovate-subhexagonal, as long as wide to wider than long; surface smooth. Maxilliped 3 dactylus digitiform, inserting near midlength of propodus. Pereopods 2–5 similarly slender; pereopod 4 or 5 dactylus elongated, linear, both longer than pereopod 2 and 3 dactyli. Pereopod 4 longest of walking legs, weakly bilaterally asymmetrical, dactyli subequal; pereopod 5 ischium and merus with line of articulation perpendicular to longitudinal axis. Male telson trapezoidal, widest distally. Maximum cl. 14 mm.

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Intertidal, subtidal (0–10 m, in bivalve molluscs, Pinnidae). Western and Central Indo-Pacific. 1 species (Ng and Ahyong 2022).

Mesotheres Ng, Ahyong & Campos, 2019 Diagnosis. Carapace of female transversely ovate, width 1.3–1.5 times length; surface unornamented, setose. Maxilliped 3 propodus shorter than carpus; dactylus minute, inserting at distal end of propodus. Pereopods 2–5 similar, setose, symmetrical, stout; pereopodal dactyli subequal, falcate, lower surface entire, subcircular in cross-section, apex simple. Pereopod 3 merus less than 3 times as long as wide. Maximum cl. 13 mm. Intertidal, subtidal (0–10 m, in gastropod molluscs). Tropical W Atlantic, Tropical Eastern Pacific. 4 species (Ng et al. 2019).

Nannotheres Manning & Felder, 1996 Diagnosis. Carapace of female subglobular, as long as wide. Maxilliped 3 palp 2-articled; propodus shorter than carpus. Pereopods 2–5 similarly slender, dactyli simple, subequal. Pereopod 3 longest of walking legs. Maximum cl. 1.5 mm. Shelf (43 m, in bivalve molluscs, Malleus). Tropical Atlantic (Caribbean Sea). 1 species (Manning and Felder 1996).

Nepinnotheres Manning, 1993 Diagnosis. Carapace of female subcircular, little wider than long. Maxilliped 3 dactylus inserting on proximal half of lower margin of propodus. Pereopods 2–5 propodus in females upper and lower margins parallel or slightly convergent distally, not expanded; dactyli subequal. Pereopod 3 longest of walking legs, longer than carapace length. Male pleonites and telson free. Gonopod 1 distally straight or curved to approximately a right-angle. Maximum cl. 20 mm. Intertidal–shelf (0–75 m, in bivalve molluscs). Tropical Atlantic, Temperate Northern W Pacific, Western and Central IndoPacific. 20 species (Ahyong and Ng 2007a: species redescriptions; Ahyong and Ng 2020; Manning 1993b: diagnosis, key to 7 W African species).

Opisthopus Rathbun, 1894 Diagnosis. Carapace of female transversely rounded-subquadrate, width 1.3 times as wide as long. Maxilliped 3 ischium and merus fused but demarcation indicated by shallow groove; dactylus broadly spatulate, inserting near base of propodus. Pereopods 2–5 bilaterally symmetrical, densely setose. Maximum cl. 14 mm. Subtidal (in holothurians, gastropod and bivalve molluscs). Temperate Northern Pacific (California). 1 species (Campos and Manning 2000: figures; Rathbun 1894).

Orthotheres Sakai, 1969 Diagnosis. Carapace of female transversely ovate, 1.4–1.6 times as wide as long. Maxilliped 3 propodus as long as carpus or longer

than carpus; dactylus digitiform, articulating at end of propodus. Pereopods 2–5 dactyli with lower surface hollowed, apex simple. Pereopod 3 or pereopod 4 longest of walking legs. Maximum cl. 10 mm. Intertidal, subtidal (in gastropod molluscs, Haliotis spp., Turbo spp.). Central Indo-Pacific. 3 species (Ng and Ho 2016b: rediagnosis, species separation).

Ostracotheres H. Milne Edwards, 1853 Diagnosis. Carapace of female subcircular, about as long as wide; surface simple, finely setose, firm to hard, anterolateral margins well defined; eyes visible in dorsal view. Maxilliped 3 palp 2-­articled; propodus longer than carpus, spatulate, apex subtruncate to rounded. Pereopodal dactyli bilaterally symmetrical, subequal or pereopod 5 dactylus longer than others. Pereopod 2 or pereopod 3 longest of walking legs. Gonopod 1 blunt, with short spine-like, anterolaterally directed papilla. Maximum cl. 15 mm. Intertidal, subtidal (0–35 m, in tridacnid clams and ascidians). Western Indo-Pacific (Red Sea). 2 species.

Parapinnixa Holmes, 1895 Diagnosis. Carapace of female transversely ovate, 1.3–2.3 times as wide as long. Pereopod 2 larger than pereopod 3, both distinctly larger, more inflated, longer than other pereopods. Maximum cl. 5 mm. Intertidal–shelf (0–89 m, associations unknown, possibly sponges). Tropical W Atlantic, Tropical Eastern Pacific. 9 species (Thoma et al. 2005).

Pinnaxodes Heller, 1865 Diagnosis. Carapace of female subcircular to subquadrate, little wider than long. Maxilliped 3 ischium and merus fused, with at most a faint trace of demarcation; dactylus broadly spatulate inserting near midlength and overreaching end of propodus. Pereopods 2–5 dactyli subequal, or pereopod 5 dactylus shorter than others. Maximum cl. 21 mm. Intertidal, subtidal (0–11 m, in echinoids, holothurians, gastropod and bivalve molluscs). Tropical W Atlantic, Tropical Eastern Pacific, Temperate South America (Peru, Chile). 7 species (figures: Garth 1957; Green 1992; Ng and Kumar 2015: species list).

Pinnixa White, 1846 Diagnosis. Carapace of female transversely subelliptical to fusiform, 2–3.5 times as wide as long; surface regions weakly defined, cardiac ridge, if present, not fully traversing carapace surface; lateral margins bluntly obtuse, strongly setose. Maxilliped 3 dactylus and propodus oriented parallel or nearly parallel to each other; exopod with angular lobe on midlateral margin. Cheliped fixed finger of mature individuals approximately in line with longitudinal axis of palm or at most, weakly deflected downwards. Pereopod 4 markedly more inflated and robust than, or almost as

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slender as pereopods 2 and 3; pereopods 4, 5 merus and ischium lower margins smooth, or with round granules. Male pleonites free. Maximum cl. 10 mm. Intertidal, subtidal, shelf (0–121 m, associated with tubicolous worms (Komai et al. 2014) and holothurians (Takeda et al. 1997)). Temperate Northern and Tropical W Atlantic, Temperate South America (W Atlantic), Temperate Northern Pacific, Central IndoPacific, Tropical Eastern Pacific. 32 species (Komatsu and Takeda 2009: key to 9 species from Japan; Palacios Theil et  al. 2016: relationships).

Pinnixulala Palacios Theil, Cuesta & Felder, 2016 Diagnosis. Carapace of female transversely ovate or fusiform, about twice as wide as long; surface regions well defined. Maxilliped 3 dactylus articulating proximal to midlength of propodus, oriented nearly parallel to each other; exopod with angular lobe on midlateral margin. Pereopod 4 markedly stouter, longer than pereopods 3 and 5; pereopods 4, 5 merus and ischium lower margins with spinules or acute granules, or smooth (if male pleonites 3–5 fused). Male pleonites free, or pleonites 3–5 fused. Maximum cl. 9 mm. Intertidal–shelf (0–54 m, commensal with polychaete worms and upogebiid mud shrimps). Temperate Northern Atlantic, Tropical Atlantic. 4 species.

Pinnotheres Bosc, 1801 Diagnosis. Carapace of female subcircular, little wider than long. Maxilliped 3 dactylus digitiform, inserting proximal to midlength of propodus. Pereopods 2–5 dactyli subequal. Pereopod 3 longest of walking legs, longer than carapace length. Male pleonites and telson free. Gonopod 1 distally sharply curved at approximately a right-angle. Intertidal–shelf (0–150 m, in bivalve molluscs). Temperate Northern Atlantic, Tropical Atlantic, Temperate Northern W Pacific, Western and Central Indo-Pacific, Temperate Southern Africa, Temperate Australasia. 35 species (Campos and VargasCastillo 2013).

Plenotheres Ng & Ahyong, 2022 Diagnosis. Carapace of female subcircular, as wide as or little wider than long; surface with 2 longitudinal grooves anteriorly. Maxilliped 3 dactylus digitiform, inserting near midlength of propodus. Pereopods 2–5 similarly slender; pereopodal dactyli falcate, apex simple. Pereopods 2 and 5 dactylus subequal, pereopod 4 bilaterally asymmetrical, longest of walking legs. Maximum cl. 14 mm. Estuarine, intertidal, subtidal (0–3 m, in bivalve molluscs, Glauconomiidae). Central Indo-Pacific. 1 species (Ng and Ahyong 2022).

Rathbunixa Palacios Theil & Felder, 2020 Diagnosis. Carapace of female transversely subelliptical to fusiform, about twice as wide as long; surface with cardiac ridge

875

not fully traversing surface; regions well defined; anterolateral margins smooth or granular, without lobiform tubercle near lateral extreme. Maxilliped 3 dactylus and propodus parallel or nearly parallel to each other. Cheliped fixed finger of mature individuals strongly def lected ventrally from longitudinal axis of palm, almost perpendicular. Pereopods 2–4 similarly slender; pereopodal dactyli slender, about as long as propodus. Male telson broadly rounded to subtriangular. Maximum cl. 10 mm. Intertidal–slope (0–436 m, associated with tubicolous polychaetes and echiurans). Temperate Northern and Tropical W Atlantic, Temperate Northern Pacific, Tropical Eastern Pacific, Temperate South America (W and E coasts). 5 species (Palacios Theil and Felder 2020a).

Raytheres Campos, 2004 Diagnosis. Carapace of female subrectangular, wider than long. Maxilliped 3 dactylus inserting on proximal half of lower margin of propodus. Pereopods 2, 3 propodi distally swollen; dactyli widest near midlength. Maximum cl. 2.2 mm. Intertidal (in bivalve, Lithophaga). Tropical Eastern Pacific. 1 species (Campos 2002: diagnosis, figures; Campos 2004: replacement name for Raymondia Campos, 2002).

Sakaina Serène, 1964 Diagnosis. Carapace of female transversely ovate, 1.6–1.8 times as wide as long. Maxilliped 3 propodus as long as carpus; dactylus little longer than wide, attached at end of propodus. Pereopods 2, 3 subequal, distinctly larger, more inflated and longer than other pereopods. Maximum cl. 6 mm. Intertidal–shelf (0–47 m, associations unknown). Temperate Northern Pacific (China, Japan, South Korea). 7 species (Jiang and Liu 2011: remarks, key to species).

Sayixa Palacios Theil & Felder, 2020 Diagnosis. Carapace of female transversely subelliptical, ~1.7 times as wide as long; surface cardiac ridge weak, short; regions scarcely defined; anterolateral margins smooth except for single lobiform tubercle just anterior to lateral extremity. Maxilliped 3 dactylus and propodus parallel or nearly parallel to each other. Cheliped fixed finger of mature individuals strongly deflected ventrally from longitudinal axis of palm, almost perpendicular. Pereopod 4 markedly stouter, longer than pereopods 3 and 5. Male telson subrectangular, more than twice as wide as long. Maximum cl. 8 mm. Subtidal, shelf (1–39 m). Tropical W Atlantic. 1 species (Palacios Theil and Felder 2020b).

Scleroplax Rathbun, 1894 Diagnosis. Carapace of female transversely oblong to subelliptical, 1.3–2.5 times as wide as long; surface regions weakly defined, cardiac ridge, if present, not fully traversing carapace surface; lateral margins rounded, without patches of setae. Maxilliped 3

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Marine Decapod Crustacea

dactylus and propodus parallel or nearly parallel to each other; exopod with angular lobe on midlateral margin. Cheliped fixed finger of mature individuals approximately in line with longitudinal axis of palm or at most, weakly deflected downwards. Pereopod 4 markedly more inflated and robust than, or almost as slender as, pereopods 2 and 3; pereopods 4, 5 merus and ischium lower margins smooth, or with round granules. Male pleonites all free, or at least clearly demarcated by sutures. Maximum cl. 16 mm. Intertidal–slope (0–311 m, associated with bivalve molluscs, tubicolous worms, holothurians, callianassid and upogebiid mud shrimps). Temperate Northern E Pacific Tropical Eastern Pacific. 7 species (Palacios Theil and Felder 2020b). The deep-water species Pinnixa scamit Martin & Zmarzly, 1994 is here included in Scleroplax.

Serenotheres Ahyong & Ng, 2005 Diagnosis. Carapace of female pentagonal; surface pitted, with broad, upraised V-shaped anterior lamella, without median tubercle. Maxilliped 3 palp 2-articled. Pereopods 2–5 similarly slender. Pereopod 3 longest of walking legs. Maximum cl. 8 mm. Intertidal, subtidal (in lithophagine clams). Central IndoPacific. 2 species (Ahyong and Ng 2005; Ng and Meyer 2016).

Sindheres Kazmi & Manning, 2003 Diagnosis. Carapace of female trapezoidal, widest posteriorly. Maxilliped 3 dactylus digitiform, inserting near midlength of propodus. Pereopods 2–4 dactyli subequal, slightly longer than pereopod 5 dactylus. Pereopod 2 longest of walking legs. Maximum cl. 3.25 mm. Intertidal (in bivalves). Western Indo-Pacific (Pakistan). 1 species (Kazmi and Manning 2003).

Solenotheres Ng & Ngo, 2010 Diagnosis. Carapace of female subquadrate, 1.5 times as wide as long. Maxilliped 3 propodus shorter than carpus; dactylus broadly spatulate, inserting near base of propodus. Pereopods 2, 4, 5 bilaterally symmetrical, right pereopod 3 longer than left. Maximum cl. 4.4 mm. Intertidal (in bivalve, Solen). Central Indo-Pacific. 1 species (Ng and Ngo 2010).

Tacitotheres Ng, Ahyong & Campos, 2019 Diagnosis. Carapace of female subcircular, little wider than long, or subcircular, about as wide as long. Maxilliped 3 propodus as long as carpus, or longer than carpus; dactylus digitiform, articulating at end of propodus. Pereopods 4, 5 propodi clavate; pereopods 2–5 dactyli subequal, falcate, apex simple, lower surface entire, subcircular in cross-section. Pereopod 3 longest of walking legs. Maximum cl. 8 mm. Intertidal, subtidal (in bivalve molluscs). Central Indo-Pacific. 3 species (Ng et al. 2019).

Tetrias Rathbun, 1898 Diagnosis. Carapace of female transversely ovate to suboblong. Maxilliped 3 ischium and merus fused but demarcation indicated by shallow groove; dactylus articulating at end of propodus, widened distally, when folded, extending to proximal midlength or as far as base of ischiomerus. Cheliped, pereopods 2–5 with spinules. Maximum cl. 7 mm. Intertidal–shelf (0–51 m, commensal with tubicolous polychaetes, Chaetopterus). Western and Central Indo-Pacific Tropical Eastern Pacific. 2 species (Rathbun 1898).

Trichobezoares Ng, 2018 Diagnosis. Carapace of female subcircular to transversely ovate, 1.1–1.3 times as wide as long; surface unornamented, margins and surface densely setose, obscuring body outline. Maxilliped 3 propodus shorter than carpus; dactylus inserting on proximal half of lower margin of propodus. Cheliped fingers distally spatulate on occlusal surface. Pereopods 2–5 similar, densely setose, symmetrical, stout; pereopodal dactyli subequal, falcate, lower surface entire, subcircular in cross-section, apex simple. Pereopod 3 merus less than 3 times as long as wide. Maximum cl. 9 mm. Subtidal (in cloaca of holothurians). Western and Central Indo-Pacific. 2 species (Chopra 1931: figures; Ng 2018: diagnosis, species).

Tridacnatheres Ahyong & Ng, 2005 Diagnosis. Carapace of female irregularly subcircular, little wider than long; margins with blunt rounded rims; surface with large median tubercle and blunt curved, thickened ridge on each side close to lateral margins. Maxilliped 3 dactylus digitiform, inserting near midlength of propodus. Pereopods 2–5 similarly slender; pereopodal dactyli falcate, apex simple. Pereopod 3 longest of walking legs. Maximum cl. 14 mm. Intertidal, subtidal (in bivalve molluscs, Tridacna, Pinna). Central Indo-Pacific. 1 species (Ahyong and Ng 2005).

Tubicolixa Palacios Theil & Felder, 2020 Diagnosis. Carapace of female transversely subelliptical to fusiform, about twice as wide as long; surface with cardiac ridge not fully traversing surface; regions well defined; anterolateral margins granular, without lobiform tubercle near lateral extreme. Maxilliped 3 dactylus and propodus oriented parallel or nearly parallel to each other. Cheliped fixed finger of mature individuals strongly deflected ventrally from longitudinal axis of palm, almost perpendicular. Pereopodal dactyli stout, shorter than propodus. Pereopods 2, 3 slender, pereopod 4 swollen, markedly stouter than pereopods 2, 3. Male telson semicircular, not more than 1.5 times as wide as long. Maximum cl. 7 mm. Intertidal–shelf (0–79 m, associated with callianassid shrimps and tubicolous polychaetes). Tropical W Atlantic. 3 species (Palacios Theil and Felder 2020b).

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Tumidotheres Campos, 1989

Xanthasia White, 1846

Diagnosis. Carapace of female subcircular to subquadrate, little wider than long; surface unornamented, setose. Maxilliped 3 dactylus broadly spatulate, inserting near midlength, not overreaching end of propodus. Pereopods 2–5 similarly slender; pereopod 5 dactylus up to 1.5 times as long as pereopods 2 and 4 dactyli (usually also that of pereopod 3). Pereopod 3 bilaterally asymmetrical. Maximum cl. 12 mm. Intertidal, subtidal (0–46 m, in bivalve molluscs). Temperate Northern and Tropical W Atlantic, Tropical Eastern Pacific Temperate South America. 4 species (Campos 1989: diagnosis; Campos and Vargas-Castillo 2013: rediagnosis; Palacios Theil and Felder 2019: species comparison, figures, hosts, phylogeography).

Diagnosis. Carapace of female transversely ovate, margins upturned; surface with prominent median mushroom-shaped tubercle. Maxilliped 3 palp 2-articled. Pereopods 2–5 similarly slender; pereopodal dactyli falcate, apex simple. Pereopod 3 longest of walking legs. Maximum cl. 14 mm. Intertidal, subtidal (in tridacnid clams). Indo-West Pacific. 1 species (Ahyong and Ng 2005: rediagnosis, figures).

Tunicotheres Campos, 1996 Diagnosis. Carapace of female subpentagonal, about as wide as long, widest at or near midlength; surface finely setose. Maxilliped 3 palp 2-articled. Pereopod 5 dactylus longest. Pereopod 5 articulated to body above level of pereopods 2–4. Maximum cl. 9 mm. Subtidal (1–5 m, in ascidians). Tropical W Atlantic. 1 species (Campos 1996a).

Zaops Rathbun, 1900 Diagnosis. Carapace of female irregularly subcircular to subhexagonal, little wider than long. Maxilliped 3 propodus longer than carpus; dactylus digitiform, inserting near midlength of propodus. Pereopods 2–5 similarly slender; 2 propodus expanded distally; pereopod 3 dactylus longest, pereopods 4, 5 dactyli subequal, longer than pereopod 2 dactylus. Pereopod 3 bilaterally asymmetrical, longest of walking legs. Maximum cl. 12 mm. Intertidal, subtidal (0–3 m, in bivalve molluscs, Ostreidae). Temperate Northern and Tropical W Atlantic. 3 species (Campos and Manning 2000: species separation; Manning 1993a: rediagnosis, figures; Palacios Theil and Felder 2019: phylogeography, species separation).

Viridotheres Manning, 1996 Diagnosis. Carapace of female subcircular to transversely ovate, as wide as or little wider than long. Maxilliped 3 dactylus digitiform, inserting near midlength of propodus. Pereopod 3 dactylus longest. Pereopods 4, 5 dactyli shorter than or as long as pereopod 2 dactylus. Pereopod 3 longest of walking legs. Maximum cl. 13 mm. Subtidal, shelf (0–100 m, in bivalve molluscs). Tropical E Atlantic, Temperate Northern E Pacific, Central Indo-Pacific. 11 species (Ahyong 2020: hosts; Ahyong et al. 2012: key to 7 species; Manning 1996).

Visayeres Ahyong & Ng, 2007 Diagnosis. Carapace of female rounded-subquadrate, little wider than long; margins with well defined edges; surface ascending to short, low median ridge. Maxilliped 3 dactylus digitiform, inserting near midlength of propodus. Pereopods 2–5 dactyli subequal, falcate, apex simple. Pereopod 3 longest of walking legs. Maximum cl. 3 mm. Shelf (34–82 m, in bivalve molluscs, Lithophaga). Central Indo-Pacific (Philippines). 1 species (Ahyong and Ng 2007b).

Waldotheres Manning, 1993 Diagnosis. Carapace of female anteriorly rounded, with posterolateral and posterior angles. Maxilliped 3 dactylus digitiform, inserting near midlength of propodus, not reaching end of propodus. Pereopods 2, 4, 5 dactyli subequal, shorter than pereopod 3 dactylus. Pereopod 3 longest of walking legs. Maximum cl. 7 mm. Intertidal (in bivalve, Donax). Tropical E Atlantic. 1 species (Manning 1993b).

References Ahyong ST (2010) Holotheres danielae, a new species of pinnotherid crab from the Indo-West Pacific (Decapoda: Brachyura), with a key to the genus. In: Castro P, Davie PJF, Ng PKL, Richer de Forges B (Eds) Studies on Brachyura: a homage to Danièle Guinot. Crustaceana Monographs 11, 35–40. Ahyong ST (2018) Revision of Ostracotheres H. Milne Edwards, 1853 (Crustacea: Brachyura: Pinnotheridae). Raffles Bulletin of Zoology 66, 538–571. Ahyong ST (2019) First Indonesian Viridotheres Manning, 1996, and redescription of male Afropinnotheres dofleini Manning, 1993, from South Africa (Decapoda, Pinnotheridae). Crustaceana 92, 107–118. doi:10.1163/15685403-00003855 Ahyong ST (2020) Resolution of the identity of Pinnotheres latipes Hombron & Jacquinot, 1846 and description of a new species of Viridotheres Manning, 1996 (Decapoda: Brachyura: Pinnotheridae): two symbionts of bivalve molluscs. Journal of Crustacean Biology 40, 879–886. doi:10.1093/jcbiol/ruaa059 Ahyong ST, Ng PKL (2005) Review of Durckheimia and Xanthasia, with descriptions of two new genera (Decapoda: Brachyura: Pinnotheridae). Journal of Crustacean Biology 25, 116–129. doi:10.1651/C-2504 Ahyong ST, Ng PKL (2007a) The pinnotherid type material of Semper (1880), Nauck (1880) and Bürger (1895) (Crustacea: Decapoda: Brachyura). Raffles Bulletin of Zoology Supplement 16, 191–226. Ahyong ST, Ng PKL (2007b) Visayeres acron, a new genus and species of pinnotherid crab (Crustacea: Decapoda: Brachyura) from the Philippines. Raffles Bulletin of Zoology Supplement 16, 187–189. Ahyong ST, Ng PKL (2008) Alain raymondi, a new species of deepwater pinnotherid crab (Crustacea: Decapoda: Brachyura) from the Philippines, commensal with holothurians. Zootaxa 1919, 61–68. doi:10.11646/​ zootaxa.1919.1.5 Ahyong ST, Ng PKL (2009) Aphanodactylidae, a new family of thoracotreme crabs (Crustacea: Brachyura) symbiotic with polychaete worms. Zootaxa 2289, 33–47. doi:10.11646/zootaxa.2289.1.3

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Ahyong ST, Ng PKL (2020) New species of pinnotherid crabs from Southeast Asia and Papua New Guinea (Crustacea: Decapoda: Brachyura). Zootaxa 4816, 333–349. doi:10.11646/zootaxa.4816.3.4 Ahyong ST, Ng PKL (2021) The pinnotherid crabs from the Gulf of Siam described by Rathbun (1909) (Decapoda: Brachyura): revisited and revised. Raffles Bulletin of Zoology 69, 188–211. doi:10.26107/ RBZ-2021-0016 Ahyong ST, Komai T, Watanabe T (2012) First Viridotheres Manning, 1996, from Japan, with a key to species (Decapoda, Brachyura, Pinnotheridae). In: Komatsu H, Okuno J, Fukuoka K (Eds) Studies on Eumalacostraca: a homage to Masatsune Takeda. Crustaceana Monographs 17, 35–47. doi:10.1163/9789004202894_003 Aikawa H (1933) On larval forms of some Brachyura. Paper II: a note on indeterminate zoeas. Records of Oceanographic Works in Japan 5, 124–254. Ayón-Parente M, Hendrickx ME (2014) Calyptraeotheres sp. nov. (Crustacea: Decapoda: Pinnotheridae), symbiont of the slipper shell Crepidula striolata Menke, 1851 (Mollusca: Gastropoda: Calyptraeidae) from the Gulf of California, Mexico. Zootaxa 3872, 89–94. doi:10.11646/ zootaxa.3872.1.8 Campos E (1989) Tumidotheres, a new genus for Pinnotheres margarita Smith, 1869, and Pinnotheres maculatus Say, 1818 (Brachyura: Pinnotheridae). Journal of Crustacean Biology 9, 672–679. doi:10.2307/1548597 Campos E (1990) Calyptraeotheres, a new genus of Pinnotheridae for the limpet crab Fabia granti Glassell, 1933 (Crustacea, Brachyura). Proceedings of the Biological Society of Washington 103, 364–371. Campos E (1993) Systematics and taxonomic remarks on Pinnotheres muliniarum Rathbun, 1918 (Crustacea: Brachyura: Pinnotheridae). Proceedings of the Biological Society of Washington 106, 92–101. Campos E (1996a) Partial revision of pinnotherid crab genera with a twosegmented palp on the third maxilliped (Decapoda: Brachyura). Journal of Crustacean Biology 16, 556–563. doi:10.2307/1548747 Campos E (1996b) Partial revision of the genus Fabia Dana, 1851 (Crustacea: Brachyura: Pinnotheridae). Journal of Natural History 30, 1157– 1178. doi:10.1080/00222939600770621 Campos E (2002) Two new genera of pinnotherid crabs from the tropical eastern Pacific (Decapoda: Brachyura: Pinnotheridae). Journal of Crustacean Biology 22, 328–336. doi:10.1163/20021975-99990239 Campos E (2004) Raytheres, a new name for Raymondia Campos, 2002 (Crustacea: Brachyura: Pinnotheridae) preoccupied by Raymondia Frauenfeld, 1855 (Hexapoda: Diptera: Streblidae). Journal of Crustacean Biology 24, 372. doi:10.1651/C-2443 Campos E (2009) A new species and two new genera of pinnotherid crabs from the northeastern Pacific Ocean, with a reappraisal of the subfamily Pinnotherinae de Haan, 1833 (Crustacea: Brachyura: Pinnotheridae). Zootaxa 2022, 29–44. doi:10.11646/zootaxa.2022.1.3 Campos E (2013) Remarks on the sexual dimorphism and taxonomy of Fabia Dana, 1851 (Crustacea, Brachyura, Pinnotheridae). Zootaxa 3616, 190–200. doi:10.11646/zootaxa.3616.2.7 Campos E, Griffith H (1990) Clypeasterophilus, a new genus to receive the small-palped species of the Dissodactylus complex (Brachyura: Pinnotheridae). Journal of Crustacean Biology 10, 550–553. doi:10.2307/1548346 Campos E, Manning RB (2000) The identities of Pinnotheres nudus Holmes, 1895 and P. nudus sensu Weymouth, 1910 (Crustacea: Decapoda: Pinnotheridae). Proceedings of the Biological Society of Washington 113, 799–805. Campos E, Vargas-Castillo R (2013) Pinnotheres orcutti Rathbun, 1918, a new eastern tropical Pacific species of Tumidotheres Campos, 1989 (Crustacea: Brachyura: Pinnotheridae). Zootaxa 3666, 84–92. doi:10.11646/zootaxa.3666.1.8 Castro P (2015) Symbiotic Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura

(Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 543–981. Brill, Leiden. Chopra B (1931) Further notes on Crustacea Decapoda in the Indian Museum. II. On some decapod Crustacea found in the cloaca of holothurians. Records of the Indian Museum 33, 303–324, pl. 7. Christensen AM, McDermott JJ (1958) Life-history and biology of the oyster crab, Pinnotheres ostreum Say. Biological Bulletin 114, 146–179. doi:10.2307/1538845 Coelho PA (1996) Revisão do gênero Alarconia Glassell, 1938, no Brasil, com descrição de Alarconia guinotae, n. sp. (Crustacea,Decapoda, Pinnotheridae). Trabalhos Oceanograficos da Universidade Federal de Pernambuco, Recife 24, 173–178. doi:10.5914/tropocean.v24i1.2705 Davie PJF, Guinot D, Ng PKL (2015) Chapter 71–18. Systematics and classification of Brachyura. In Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea complimentary to the volumes translated from the French of the Traité de Zoologie. Decapoda: Brachyura (Part 1). (Eds Castro P, Davie PJF, Guinot D, Schram FR, von Vaupel Klein JC) pp. 1049–1130. Brill, Leiden. de Gier W, Becker C (2020) A review of the ecomorphology of pinnotherine pea crabs (Brachyura: Pinnotheridae), with an updated list of symbionthost associations. Diversity 12, 431. doi:10.3390/d12110431 Felder DL, Palacios Theil E (2020) Three new symbiotic crabs of the genus Glassella Campos & Wicksten, 1997 from Atlantic and Gulf of Mexico coasts of Florida, USA (Decapoda: Brachyura: Pinnotheridae). Journal of Crustacean Biology 40, 899–917. doi:10.1093/jcbiol/ruaa072 Garth JS (1957) Reports of the Lund University Chile Expedition 1948–49. 29. The Crustacea Decapoda Brachyura of Chile. Lunds Universitets Årsskrift N.F. Avd. 2 53, 1–129, pls 1–4. Glassell SA (1938) New and obscure decapod Crustacea from the West American coasts. Transactions of the San Diego Society of Natural History 8, 411–454, pls 27–36. Gore RH (1986) Fabia felderi species novum, a new pinnotherid crab from the central eastern coast of Florida (Crustacea: Decapoda: Brachyura). Northeast Gulf Science 8, 143–148. doi:10.18785/negs.0802.05 Green TM (1992) Pinnaxodes gigas, a new species of pinnotherid crab from the Gulf of California (Decapoda: Brachyura: Pinnotheridae). Proceedings of the Biological Society of Washington 105, 775–779. Griffith H (1987) Taxonomy of the genus Dissodactylus (Crustacea: Brachyura: Pinnotheridae) with descriptions of three new species. Bulletin of Marine Science 40, 397–422. Heard RW, Manning RB (1997) Austinixa, a new genus of pinnotherid crab (Crustacea: Decapoda: Brachyura), with the description of A.hardyi, a new species from Tobago, West Indies. Proceedings of the Biological Society of Washington 110, 393–398. Holthuis LB (1975) Limotheres, a new genus of pinnotherid crab, commensal of the bivalve lima, from the Caribbean sea. Zoölogische Mededeelingen 48, 291–295. Jiang W, Liu R (2011) New species and new records of pinnotherid crabs (Crustacea: Decapoda: Brachyura) from the Yellow Sea. Zoologischer Anzeiger 250, 488–496. doi:10.1016/j.jcz.2011.05.001 Kazmi QB, Manning RB (2003) A new genus and species of pinnotherid crab from Karachi, northern Arabian Sea (Crustacea, Decapoda, Brachyura). Journal of Natural History 37, 1085–1089. doi:10.1080/​00222930110108353 Komai T, Naruse T, Yokooka H, Taru M, Shimetsugu M, et  al. (2022) Redescription of Pinnixa haematosticta Sakai, 1934, its transfer to Indopinnixa Manning & Morton, 1987, and a reappraisal of Indopinnixa kumejima Naruse & Maenosono, 2012 (Decapoda: Brachyura: Pinnotheridae). Zootaxa 5100, 361–389. doi:10.11646/zootaxa.5100.3.3 Komai T, Nishi E, Taru M (2014) A new species of Pinnixa (Crustacea: Decapoda: Brachyura: Pinnotheridae) associated with a tube worm, Chaetopterus cautus (Annelida: Polychaeta), from Tokyo Bay, Japan. Zootaxa 3793, 119–132. doi:10.11646/zootaxa.3793.1.5

14 – Brachyura – crabs

Komai T, Kawai K, Ng PKL (2020) On the identity of the poorly known pea crab, Pinnothera obesa Dana, 1852, and description of a new species of Arcotheres Manning, 1993 from the Southwest Pacific (Decapoda: Brachyura: Pinnotheridae). Zootaxa 4822, 221–247. doi:10.11646/zootaxa.4822.2.5 Komatsu H, Ohtsuka S (2009) A new species of the genus Abyssotheres (Crustacea, Decapoda, Brachyura, Pinnotheridae) from the Ryukyu Islands, southwestern Japan, with taxonomic notes on the genus. Bulletin of the National Science Museum, Tokyo, Series A (Zoology) 35, 73–81. Komatsu H, Takeda M (2009) A new species of the genus Pinnixa (Decapoda: Brachyura: Pinnotheridae) from Otsuchi Bay, northeastern Japan. Bulletin of the National Museum of Nature and Science. Series A, Zoology 3, 199–204. Manning RB (1993a) Three genera removed from the synonymy of Pinnotheres Bosc, 1802 (Brachyura: Pinnotheridae). Proceedings of the Biological Society of Washington 106, 523–531. Manning RB (1993b) West African pinnotherid crabs, subfamily Pinnotherinae (Crustacea, Decapoda, Brachyura). Bulletin du Muséum National d’Histoire Naturelle, Paris, 4e série, A 15, 125–177. Manning RB (1996) Viridotheres marionae, a new genus and species of pinnotherid crab from west Africa (Crustacea: Decapoda: Brachyura). Zoölogische Mededeelingen 70, 271–273. Manning RB (1998) A new genus and species of pinnotherid crab (Crustacea, Decapoda, Brachyura) from Indonesia. Zoosystema 20, 357–362. Manning RB, Felder DL (1996) Nannotheres moorei, a new genus and species of minute pinnotherid crabs from Belize, Caribbean Sea (Crustacea: Decapoda: Pinnotheridae). Proceedings of the Biological Society of Washington 109, 311–317. Manning RB, Holthuis LB (1981) West African brachyuran crabs. Smithsonian Contributions to Zoology 306, 1–379. doi:10.5479/si.00810282.306 Manning RB, Morton B (1987) Pinnotherids (Crustacea: Decapoda) and leptonaceans (Mollusca: Bivalvia) associated with sipunculan worms in Hong Kong. Proceedings of the Biological Society of Washington 100, 543–551. Naruse T, Maenosono T (2012) Two new species of Indopinnixa Manning & Morton, 1987 (Decapoda: Brachyura: Pinnotheridae) from the Ryukyu Islands, Japan. Zootaxa 3367, 222–231. doi:10.11646/zootaxa.​ 3367.1.21 Ng PKL (2018) On the identities of Pinnotheres villosissimus Doflein, 1904, P. dofleini Lenz, in Lenz & Strunck, 1914, and P. pilumnoides Nobili, 1906 (Decapoda, Brachyura, Pinnotheridae) from the western Indian Ocean. Crustaceana 91, 611–633. doi:10.1163/15685403-00003785 Ng PKL, Ahyong ST (2022) The pea crabs of the genus Arcotheres Manning, 1993 (Crustacea: Brachyura), from Singapore and Peninsular Malaysia, with a reappraisal of diagnostic characters and descriptions of two new genera. Raffles Bulletin of Zoology 70, 134–248. doi:10.26107/RBZ-2022-0009 Ng PKL, Ho P-H (2016a) A new genus for Fabia obtusidentata Dai, Feng, Song and Chen, 1980, a pea crab (Decapoda: Brachyura: Pinnotheridae) symbiotic with the moon scallop Amusium pleuronectes (Linnaeus, 1758) (Mollusca: Pectinidae). Journal of Crustacean Biology 36, 740–751. doi:10.1163/1937240X-00002474 Ng PKL, Ho P-H (2016b) Orthotheres baoyu, a new species of pea crab (Crustacea: Brachyura: Pinnotheridae) associated with abalones from Tungsha Island, Taiwan; with notes on the genus. Raffles Bulletin of Zoology 64, 229–241. Ng PKL, Kumar AB (2015) A new species of Afropinnotheres Manning, 1993 (Crustacea, Brachyura, Pinnotheridae) from southwestern India, the first record of the genus from the Indian Ocean, with a review of the Pinnotheridae of India and adjacent seas. Zootaxa 3947, 264–274. doi:10.11646/zootaxa.3947.2.8 Ng PKL, Manning RB (2003) On two new genera of pea crabs parasitic in holothurians (Crustacea: Decapoda: Brachyura: Pinnotheridae) from

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the Indo-West Pacific, with notes on allied genera. Proceedings of the Biological Society of Washington 116, 901–919. Ng PKL, Meyer C (2016) A new species of pea crab of the genus Serenotheres Ahyong & Ng, 2005 (Crustacea, Brachyura, Pinnotheridae) from the date mussel Leiosolenus Carpenter, 1857 (Mollusca, Bivalvia, Mytilidae, Lithophaginae) from the Solomon Islands. ZooKeys 623, 31–41. doi:10.3897/zookeys.623.10272 Ng PKL, Ngo VT (2010) Solenotheres prolixus, a new genus and new species of pinnotherid crab (Crustacea: Decapoda: Brachyura) associated with the razor clam, Solen corneus Lamarck, 1818 (Solenidae) in Vietnam. Zootaxa 2570, 61–68. doi:10.11646/zootaxa.2570.1.4 Ng PKL, Guinot D, Davie PJF (2008) Systema Brachyurorum; Part I. An annotated checklist of extant brachyuran crabs of the world. Raffles Bulletin of Zoology Supplement 17, 1–286. Ng PKL, Ahyong ST, Campos E (2019) Two new genera of pinnotherid crabs (Crustacea: Brachyura: Pinnotheroidea) from the Americas and the Western Pacific. Raffles Bulletin of Zoology 67, 337–351. Palacios Theil E, Felder DL (2019) Molecular phylogeography of Tumidotheres maculatus (Say, 1818) and Zaops ostreus (Say, 1817) (Crustacea: Decapoda: Pinnotheridae) in the western Atlantic, with description of a new species and synonymy of Epulotheres Manning, 1993. Marine Biology Research 15, 548–567. doi:10.1080/17451000.2019.1693597 Palacios Theil E, Felder DL (2020a) Phylogeny of the genus Austinixa Heard & Manning, 1997, inferred from mitochondrial and nuclear molecular markers, with descriptions of three new species and redescription of Austinixa felipensis (Glassell, 1935) (Decapoda: Brachyura: Pinnotheridae). Zootaxa 4778, 101–134. doi:10.11646/zootaxa.4778.1.4 Palacios Theil E, Felder DL (2020b) Phylogeny of the genus Pinnixa White, 1846 (Crustacea, Brachyura, Pinnotheridae) and allies inferred from mitochondrial and nuclear molecular markers, with generic reassignment of twenty-one species. Zoosystema 42, 85–103. doi:10.5252/ zoosystema2020v42a6 Palacios Theil E, Cuesta JA, Felder DL (2016) Molecular evidence for nonmonophyly of the pinnotheroid crabs (Crustacea : Brachyura : Pinnotheroidea), warranting taxonomic reappraisal. Invertebrate Systematics 30, 1–27. doi:10.1071/IS15023 Rathbun MJ (1894) Scientific results of explorations by the U.S. Fish Commission steamer Albatross. No. XXIV. Descriptions of new genera and species of crabs from the west coast of North America and the Sandwich Islands. Proceedings of the United States National Museum 16, 223–260. doi:10.5479/si.00963801.933.223 Rathbun MJ (1898) The Brachyura collected by the U.S. Fish Commission steamer Albatross on the voyage from Norfolk, Virginia, to San Francisco, California, 1887–1888. Proceedings of the United States National Museum 21, 567–616, pls 41–44. Salgado-Barragán J, Raymundo-Huizar AR, Ayón-Parente M (2021) A new species of Austinixa Heard and Manning 1997 (Decapoda: Pinnotheridae) and new records of A. felipensis (Glassell, 1935) from the Mexican Pacific. Nauplius 29, e2021022. doi:10.1590/2358-2936e2021022 Schmitt WL, McCain JC, Davidson E (1973) Decapoda I. Brachyura I. Fam. Pinnotheridae. Crustaceorum Catalogus 3, 1–160. Takeda S, Tamura S, Washio M (1997) Relationship between the pea crab Pinnixa tumida and its endobenthic holothurian host Paracaudina chilensis. Marine Ecology Progress Series 149, 143–154. doi:10.3354/meps149143 Thoma BP, Heard RW, Vargas R (2005) A new species of Parapinnixa (Decapoda: Brachyura: Pinnotheridae) from Isla del Coco, Costa Rica. Proceedings of the Biological Society of Washington 118, 543–550. doi:10.2988/0006-324X(2005)118[543:ANSOPD]2.0.CO;2 Tsang LM, Ahyong ST, Shih H-T, Ng PKL (2018) Further polyphyly of pinnotheroid crabs: the molecular phylogenetic position of the polychaete-associated Aphanodactylidae. Invertebrate Systematics 32, 92–99. doi:10.1071/IS17038

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Glossary

Crustaceans, especially eucaridans, are built on essentially the same body-plan, a head and thorax covered by a carapace, plus a pleon. Homologies exist between the major taxa for all the limbs and for the regions into which the carapace is sculptured. Terminology is much the same for all taxa but there are minor differences according to body type. The glossary includes most of the specialised terms but hybrid words, like gastrofrontal, are not defined. Plurals and adjectival forms are given for some. Alternative terms not used in this volume (italicised) are cross-referred to our preferred term. Serial structures are numbered in sequence, e.g. maxilliped 3, pereopods 2–4, rather than third maxilliped, second to fourth pereopods. The terms spines and teeth are confined to cuticular expansions or protrusions. ‘Movable spine’ is often used to refer to sharp, hard mobile structures especially on pereopods – we usually use ‘spiniform seta’ instead.

Orientation Anterior and posterior refer to the head and tail ends of a typical decapod, dorsal and ventral to the top and bottom surfaces when the animal is oriented normally. In some literature dorsal and ventral, or anterior and posterior, are also used to refer to one or other edge of a limb when the terms upper and lower or extensor and flexor are anatomically more correct. Extensor and flexor refer to the margins corresponding to the action of the internal muscles along each margin. Similarly, strictly speaking, legs do not have a dorsal and ventral, or anterior and posterior, surface or edge because they do not have fixed orientation, but the terms may be applied to describe the position of spines or ridges. Medial refers to a structure in the midline of the animal and mesial to the face of a paired limb facing the midline. Lateral is the opposite of mesial or refers to a position away from the midline. Sublateral is used to indicate, for example, a spine between the medial and lateral spines. Proximal and distal describe the attached and remote ends of a limb. Adjectives describing orientation are frequently combined to indicate intermediate conditions, e.g. posterolateral, anteroventral etc. The body and appendages are divided into segments. The segments of the body are here called somites, arranged in three sections (tagmata), cephalon (head), thorax and pleon (Figs G1, G2). In the decapod literature, pereon and

pleon are terms often interchangeably used for thorax and abdomen, respectively. The segments of appendages or limbs are called articles. The word segment is used for both somite and article in some literature.

Prawns, lobsters and shrimps Identification of families, genera and species of prawns, lobsters and shrimps relies on differences in the sculpture and spination of the carapace and the rostrum. Basic nomenclature is the same for all groups (see figure of typical caridean shrimp). The rostrum may carry dorsal and ventral teeth and may have an adrostral ridge running parallel along each side. The orbit (concavity on each side of the rostrum) is defined at its lower end by an antennal spine. In some carideans, an orbital spine sits between the rostrum and antennal spine. The free side of the carapace, called the pterygostomian, or branchiostegite when it covers the gills, may have a pterygostomial spine at its anteroventral corner and a branchiostegal spine nearby but away from the margin. Running usually obliquely across the carapace is the cervical ridge and groove. The hepatic spine is situated at or near the lower edge of the cervical groove. The cervical groove may run completely across the dorsal part of the carapace or may be visible only laterally. Ridges may run posteriorly from the antennal, orbital and branchiostegal spines (Fig. G1). Other spines and grooves are important in some groups but not others. Spines on the dorsal regions of the carapace are labelled according to the area from which they arise. Males and females differ in the shape and number of the first two pairs of pleopods. This is most obvious in Dendrobranchiata where males possess a petasma, a complex formed from the paired endopods of the first pleopods. In some groups, males may lack or have reduced first pleopods and have an additional lobe, the appendix masculina, on pleopod 2. Secondary sexual characters of the chelipeds or other limbs differentiate the sexes of some groups.

Crabs Identification of crabs to family, genus and species relies on understanding crab anatomy and the special terms applied. The dorsal surface of the carapace is consistently divided into regions more or less obviously divided by grooves. The names of the regions, or areolae, reflects the internal

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Fig. G1.  Typical caridean shrimp (Palaemon serenus). 1–6, pleonites 1–6; a1, antennule; a2, antenna; as, antennal spine; b, basis; B, branchiostegite; bg, branchiostegal groove; bs, branchiostegal spine; c, carpus; d, dactylus; e, eyestalk; i, ischium; m, merus; mp3, maxilliped 3; P, pleon; p, propodus; p1–p5, pereopod 1–pereopod 5; pt, pterygostomial angle; r, rostrum; sc, scaphocerite; st, stylocerite; t, telson; u; uropod. Hepatic spine not shown in this species.

organs – gastric for stomach, cardiac for heart, hepatic for ‘liver’ and intestinal for intestine. Prefixes like epi- and prorefer to divisions of these (see figures of a typical crab (Fig. G2)). Ornamentation of the carapace margin is also critical, especially in Xanthoidea (Fig. 14.141). The front is between the eyes and may be lobed; the eyestalks have a terminal cornea and are in orbits, or sockets, which may be protected by spines; the anterolateral and posterolateral margins run from the orbit to the back of the carapace and are more or less distinct. The anterolateral margin is frequently dentate. The five pairs of pereopods include the chelipeds plus four pairs of walking or ambulatory legs. We use cheliped for the first pair of pereopods and prefer pereopods 2–5 for the others. The buccal field or cavern is the large, usually rectangular opening below the eyes, often defined by ridges, and covered by the pair of third maxillipeds. In most crabs the antennules (first antennae) and antennae (second antennae) are small. The mobile articles of the antennules usually sit in a pair of cavities (fossae) between the eyes. The antennae are often larger and lie lateral to the antennules and next to the orbit. The first two articles of the antenna are usually

fused to the carapace and termed the antennal ‘basal article’. The antennal basal article can be easily overlooked; its relationship to the orbit can be significant. Males and females are easily distinguished. The narrow pleon of the male may have fused segments; the shape of its two pairs of pleopods (rarely more), called gonopods 1 and 2, can be critical. Females have a broader and usually freer pleon and five pairs of biramous pleopods. The sexes may be further differentiated by secondary sexually characters of size, spination or development of chelipeds.

Glossary abdomen see pleon abdominal somite see pleonite abyssopelagic pelagic at abyssal depths, greater than 2000 m depth acicle see scaphocerite adrostral ridge ridge parallel to and separated by narrow groove from rostrum (Penaeoidea) afferent branchial channels see inhalent branchial channel

Glossary

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Fig. G2.  Diagrammatic dorsal and ventral views of typical brachyuran crab: carapace regions and names of articles of pereopods, reproduced from Rathbun MJ (1918) The grapsoid crabs of America. Bulletin of the United States National Museum 97, 1–461, pls 1–161. Rathbun’s ‘ambulatory legs’ are referred to as pereopods 2–5 in this guide, her ‘abdomen’ as pleon.

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Marine Decapod Crustacea

antenna (pl. antennae) second cephalic appendage, usually with peduncle of five articles and multiarticulate flagellum; plural sometimes used to refer to both antennules and antennae ambulatory legs walking legs in crabs, here referred to as pereopods 2–5 antennal scale see scaphocerite antennal spine spine on anterior margin of carapace ventral or lateral to orbit (antennal angle if spine absent); usually positioned above the antenna antennule first cephalic appendage, usually with peduncle of three articles and two multiarticulate flagella, one of which may be duplicate (some Caridea) anterolateral margin margin of carapace between orbit and lateral angle or spine appendix interna (pl. appendices internae) small branch on inner margin of pleopod 2, triangular or rod-like, usually with clusters of hooks to attach to its pair appendix masculina (pl. appendices masculinae) small branch on inner margin of pleopod 2 endopod of males, usually rod-like, proximal to appendix interna arthrobranch gill attaching to membrane between coxa and side of body, 1 or 2 pairs per somite and usually pointing upwards article any of the segments of a limb or appendage (see coxa, basis, ischium, merus, carpus, propodus, dactylus) basis (pl. bases; adj. basial or basal) second article of limb bathypelagic pelagic at bathyal depths, 200 to 2000 m depth benthic living on the sea floor; infaunal species burrow, epibenthic species walk over the surface (cf. pelagic) branchiocardiac (ridge or groove) line along dorsal limit of branchiostegite branchiostegite (adj. branchiostegal) expanded ventrolateral part of carapace covering gills branchia (pl. of branchos, rarely used; adj. branchial) gills, usually thoracic branchial cavity space between thorax and lateral flap of carapace (branchiostegite) enclosing branchia or gills branchial region paired regions of dorsal surface of the carapace between hepatic regions in front and posterior margin behind, sometimes bearing branchial spine(s), defined laterally in crabs by posterolateral margin buccal cavern space, especially in crabs, enclosing the mouthparts and more or less covered by third maxillipeds, defined anteriorly by epistome and laterally by pterygostomial region, sometimes by ridges (sometimes buccal frame) carapace shield arising from posterior margin of head and comprising lateral extensions of thoracic somites covering all or most of cephalothorax carcinisation tendency for decapods to evolve towards a crab-like form cardiac a median region of the carapace, between the gastric and intestinal regions carina (adj. carinate) keel, ridge, crest carpus fifth article of limb (first article of palp in maxilliped 3) cephalon head, body segments bearing eyes, antennules, antennae, mandibles, maxillules and maxillae

cephalothorax all fused somites of the cephalon (head) and thorax, covered in Decapoda by carapace cervical groove groove across the middle of the carapace between gastric and cardiac regions, curving forwards laterally, sometimes incomplete medially, sometimes accompanied by cervical ridge and ending laterally in cervical incision in some lobsters, sometimes with paired postcervical spines chela (adj. chelate) pincer, derived from interacting fixed finger (pollex, extension from palm of propodus) and moveable finger (dactylus); palm refers to the part of the propodus excluding the finger cheliped limb with a chela chromatophore pigmented cell in exoskeleton cincinnuli minute hooked processes interlocking halves of petasma (Dendrobranchiata) continental shelf sea floor between low water level and shelf break, usually about 200 m depth continental slope sea floor between edge of continental shelf, usually about 200 m depth, and top of continental rise, about 2000 m depth (often refers to slope and rise) continental rise sea floor between lower limit of continental slope and abyssal plain (about 1000 to 4000 m) cornea faceted and pigmented part of eyestalk coxa first article of limb (closest to body), usually short dactylus seventh article of limb (last article interacting with fixed finger to form chela in chelate limb) dendrobranchiate gill gill with paired primary branches subdivided efferent branchial channels see exhalent branchial channels endite lobe, usually mesial, of proximal articles of limbs, especially mouthparts endopod inner of two branches of biramous limb, comprising ischium through to dactylus; in many decapods the dominant or only branch endostome plate inside upper part of buccal cavern surrounding mouth endostomial ridges pair of longitudinal ridges inside the buccal cavern and reaching the anterior margin, concealed by the third maxillipeds epipod lateral branch of limb attached sometimes with gill (podobranch) to coxa, coxal endite epistome region of carapace above the mouth between front of carapace and antennules exhalent branchial channels pair of channels venting branchial cavity and opening lateral to endostome (most Brachyura) (also called efferent branchial channels) exopod outer of two branches of biramous limb, comprising one or several short articles arising from basis; absent from many pereopods exorbital tooth tooth lateral to the orbit at the start of the anterolateral margin of the carapace; also known as outer orbital tooth eyestalk articulating eye comprising peduncle and (usually) pigmented cornea fixed finger prolongation of propodus of chelate limb (sometimes pollex)

Glossary

flagellum (pl. flagella) multiarticulate distal extension of antennule (2, rarely 3, present) and antenna (1 only); also at the end of exopods of maxillipeds front anterior median region of upper carapace between inner margins of orbits and gastric region; or in crabs the margin between the orbits gastric region median region of dorsal carapace between frontal region and cardiac region; divided from anterior in midline into mesogastric, metagastric and urogastric regions, and with protogastric and epigastric regions laterally gonopod one of the first two pair of male pleopods modified for copulation hepatic region paired regions of dorsal carapace between orbits and branchial regions, defined laterally in crabs by anterolateral margin, sometimes bearing hepatic spine(s), and sometimes (prawns and lobsters) with a curved hepatic groove or ridge ventrally (see subhepatic) incisor terminal process of mandible, cutting blade, often toothed (more properly, incisor process) inhalent branchial channels pair of channels between ridges, supplying water to branchial cavity and opening in front of chelipeds or alongside epistome (some Brachyura). Also known as afferent branchial channels intestinal region most posterior median region of the dorsal carapace, sometimes with intestinal spine(s) ischiomerus fused ischium and merus, especially of maxilliped 3 and pereopods 1–5 ischium (adj. ischial) third article of limb (first article of endopod) linea anomurica longitudinal lateral groove defining upper margin of branchiostegite in Anomura linea thalassinica longitudinal lateral groove defining upper margin of branchiostegite in Axiidea mandible (adj. mandibular) third limb of cephalon, comprising a thick body with mesial molar process, terminal incisor process and mandibular palp, usually of 3 articles maxilla fifth limb of cephalon, epipod sometimes developed as scaphognathite maxilliped one of three pairs of thoracic mouthparts; maxilliped 1 is membranous like the maxilla, maxilliped 2 a little more robust, and maxilliped 3 is much larger and may be more or less opercular over the other mouthparts in crabs, or pereopod-like in shrimps and prawns maxillule fourth limb of cephalon megalopa terminal larval stage in Brachyura and Anomura merus (adj. meral) fourth article of limb mesopelagic midwater over continental slope, beyond 200 m depth midwater pelagic monophyletic describing a higher taxon comprising all the species descendant from a common ancestor mouthparts all cephalothoracic limbs involved with feeding: mandibles through to maxillipeds 3 obsolete scarcely developed, indistinct; of a spine, tooth or lobe ocular of the eye ocular peduncle see eyestalk ocular scale articulating plate attached mesially on eyestalks (some Anomura)

885

orbitorostral groove paired grooves between median ridge extending to rostrum and ridges above eyes (Alpheidae) palm body of propodus of chelate limb (excluding fixed finger) paraphyletic describing a group comprising some but not all species descendant from of the same common ancestor pectinate (adj.) formed of short, thin, closely spaced upright projections, comb-like peduncle proximal segments of biramous limb, especially of antennule, antenna, pleopods and uropod (sometimes protopod or sympod) pelagic free-swimming in water mass, not associated with sea floor (cf. benthic) pereopod non-maxilliped thoracic limbs; five pairs in Decapoda (pereopods 1–5 are attached to thoracic somites 4–8); includes one pair of chelipeds and four pairs of walking legs in Brachyura. Each pereopod comprises the seven articles, coxa, basis and five endopodal articles, ischium, merus, carpus, propodus, dactylus, plus sometimes an exopod (attached to the basis) petasma male genital structure comprising the enlarged and coupled endopods of pleopods 1 (Dendrobranchiata); usually composed of complex lobes and lobules photophore luminescent integumental organ, sometimes with lens phyllobranchiate gill gill with plate-like or leaf-like branches arranged in paired series along main axis phyllosoma pelagic larva of Achelata (Fig. 11.1), develops eventually into puerulus stage before settlement pleon posterior part of body, of five segments or somites plus telson (also abdomen) pleonite segment or somite of the pleon pleomere see pleonite pleopod one of paired limbs on pleonites 1–5, usually biramous pleotelson fused pleonite 6 and telson pleurobranch gill attached to body wall inside branchial cavity dorsal to coxae of pereopods, usually 1 pair per somite and pointing down between arthrobranchs pleuron (pl. pleura) lateral part of body wall, especially of pleonites, produced ventrally podobranch gill attached to coxa of pereopod or its epipod polyphyletic describing a group comprising unrelated species descendant from different common ancestors post- prefix meaning posterior to, as in postrostral, postcervical etc. propodus sixth article of limb (comprising palm + often fixed finger of chela) prosartema process arising from mesial margin of first article of antennule (Dendrobranchiata) pterygostomial region anteroventral region of carapace (anterior part of branchiostegite of Caridea) or ventral region of carapace (of Brachyura); bearing pterygostomial spine at anteroventral angle (in Caridea and Dendrobranchiata) ramus (pl. rami) branch of limb; endopod is inner branch and exopod is outer branch rostrum (adj. rostral) prolongation of median part of anterior carapace between eyes scaphocerite lateral plate or spine attached to distal margin of second article of antenna, equivalent to exopod; often called

886

Marine Decapod Crustacea

antennal acicle when spiniform, antennal scale when broad, flat scaphognathite expanded epipod of maxilla (Caridea, Axiidea) seamount submarine mountain, usually volcanic and steep-sided, reaching much shallower water than surrounding continental slope or abyssal depths seminal receptacles sacs for storage of spermatophores or sperm, usually on sternite of thoracic somite 7 (sometimes spermathecae), associated with thelycum in Dendrobranchiata shield of hermit crabs, the anterior harder part of the carapace in front of the cervical groove somite body segment, as in thoracic somites 1–8 and pleonites 1–6 spermatophore mass of spermatozoa deposited by gonopods of male on to thoracic sternum or into seminal receptacle of female statocyst cavity filled with fluid and particles acting as organ of orientation sternite ventral plate of integument (cf. tergite) sternum ventral surface of body, to which limbs are attached, or specifically fused thoracic sternites 1–8 stridulating organ sound-producing structure on limbs and/or carapace formed by row of regularly ridged or beaded integument and engaged by opposing structure on another limb (e.g., paired chelipeds in Strigopagurus or cheliped and carapace in ocypodids) stylamblys specialised second endopodal article of pleopod 2 of female Achelata

stylocerite sharp scale arising from lateral base of first article of antennular peduncle (Caridea) sub- prefix meaning almost or close to subchelate chela-like, formed of transverse end of propodus and opposing dactylus (cf. chelate) supraocular or supraorbital spine or projection spine or lobe on carapace above eyes (some Achelata or Caridea) tailfan telson plus pair of uropods telson plate attached medially to terminal segment of pleon tergite dorsal plate of integument (cf. sternite) tergum arched dorsal part of pleonites thelycum female genitalia comprising modified posterior two or three thoracic sternal plates (Dendrobranchiata) thoracic somite: one of the eight segments of the thorax (also pereonite) thoracic sternite plate between maxillpeds and pereopods thoracomere see thoracic somite thorax middle body region of eight somites bearing maxillipeds 1–3 and pereopods 1–5 tomentum mat of fine setae trichobranchiate gill gill with fine branches on a central axis uropod paired limb of sixth pleonite, combining with telson to form tailfan walking leg see pereopod zoea (adj. zoeal) one of several larval stages, first stage hatching from the egg; transforming to final megalopa stage

Credits

Figures Most line illustrations in this work are assembled tracings of f­igures in the literature, all more or less simplified, undertaken by GCBP using Adobe Illustrator®. Others, listed below, are reprinted directly with permission of the publisher and/or author or are in the public domain. The references cited can be found in the relevant chapters. F ­ igure 1.1, Spalding et al. (2006), map supplied by Marine Regions team, Flanders Marine Institute. ­Figure 3.1a, Poore (2004); b, c, e–i, Crosnier (1978); j, Pérez Farfante (1987). ­Figure  3.2a, Crosnier and Vereshchaka (2008); b, e, Pérez Farfante and Kensley (1997); c, Crosnier (1985); d, Joubin (1905). ­Figure  3.4a, b, d–f, i, Pérez Farfante and Kensley (1997); c, g, h, Crosnier (1985, 1991, 1994). F ­ igure 3.5a, Crosnier (1986); b–i, Pérez Farfante and Kensley (1997). ­Figure 3.6, Pérez Farfante and Kensley (1997). F ­ igure 3.7a, Poore (2004); b, Chan (2012). F ­ igure 3.9e–g, Calman (1913). Figure  3.10a, Poore (2004). ­ ­ Figure  3.11a, Poore (2004). F ­ igure 4.1c, Alvarez et al. (2006). F ­ igure 4.2a, Poore (2004). F ­ igure 4.3a, Saito et al. (2009). F ­ igure 5.1a, Chace and Manning (1972). F ­ igure  6.8a, b, d–g, Poore (2004); c, Kemp (1914). F ­ igure  6.10e, Nguyen (2000). ­Figure  6.11a, Komai (2002. ­Figure  6.12, Poore (2004). F ­ igure  6.13a, b, Yokoya (1930); k, Komai and Fujiwara (2012). F ­ igure 6.14a, b, Poore (2004). F ­ igure  6.17a, Bruce (1990). ­Figure  6.19a, Kensley (1983). ­Figure 6.21a, Poore (2004). ­Figure 6.22a, Taylor and Collins (2009); b, Hanamura et al. (1999); c, Poore (2004); d, Ďuriš (1992); e, Komai and Matsuzaki (2016). F ­ igure 6.24a, Poore (2004). F ­ igure 6.25, Boone (1927). F ­ igure 6.26a, Poore (2004). F ­ igure  6.28c, Poore (2004). ­Figure  6.29a, d, f, g, Poore (2004); b, Crosnier and Forest (1973). ­Figure 6.31a, d, Poore (2004); b, Walker and Poore (2003). F ­ igure  6.46a, Poore (2004). F ­ igure  6.47a, b, Poore (2004). F ­ igure  6.48v, Fransen (1990). ­Figure 6.49a, b, d, f, Poore (2004); e, Crosnier & Forest (1973); g, Lin et al. (2001). F ­ igure 6.50a, Chace (1940). F ­ igure 6.51a, Poore (2004). ­Figure 6.54a, Mejía-Ortíz et al. (2017). ­Figure 7.2a, b, Poore (2004). ­Figure 8.1a, Forest and de Saint Laurent (1989); b, c, Forest (2006); d, Richer de Forges (2006). F ­ igure 10.1a, Poore (2004). ­Figure 10.3a, d–g, i–k, n, o, q, Poore (2004); b, Alcock and McArdle (1901); c, Williams (1993); h, Sakai (2006); l, m, p, Sakai (1982). ­Figure  10.4d, Williams and Ngoc-Ho (1990). ­Figure  11.1, Haeckel (1904: Issue 8, pl. 76). ­Figure  11.3a, d, f, g, Poore (2004); b, c, e, h, j, Holthuis (1985, 1991, 2002); i, De Man (1916). F ­igure  11.4b, De Man (1916); q, Poore (2004).

­ igure 12.1a, b, Bate (1888), c, d, Poore (2004). ­Figure 13.1a, F b, e, g–h, l, Poore et al. (2011); f, Poore and Andreakis (2011). F ­ igure 13.2, Poore et al. (2011). ­Figure 13.3b–d, Macpherson et al. (2005). ­Figure 13.4a, Poore et al. (2011). F ­ igure 13.5a–k, Poore et al. (2011); l, Macpherson and Robainas-Barcia (2013). F ­ igure  13.6a–g, i–o, Poore et al. (2011); h, Cabezas and Macpherson (2014). ­Figure 13.7a–e, l, Poore et al. (2011); f–i, Baba and de Saint Laurent (1996); j, Ahyong (2007). F ­ igure  13.9a–d, Poore et al. (2011); e, Poore (2014). ­Figure  13.10c, Poore (2004); d, Miyake (1942); f, Dolorosa and Werding (2014); g, n, Glassell (1938); h, Ng and Nakasone (1993); i, Osawa (2001); q, Osawa (1998). F ­ igure 13.11a, Glassell (1936); b, e, f, i, Poore (2004); c, Werding and Hiller (2017); j, Lemaitre and Campos (2000); n, Glassell (1938). ­Figure 13.16, Poore (2004). F ­ igure 13.17a, Alcock (1905: pl. 14 Fig. 3). F ­ igure  13.18a, d, f, h, Poore (2004); b, Alcock (1905); c, g, McLaughlin (2003); e, Forest (1995). ­Figure  13.20a–d, Schmitt (1921); d, Makarov (1938); f, g, McLaughlin (2003). F ­ igure  13.21a, b, e, Schmitt (1921); c, Haig (1974); d, h, Ahyong (2010); f, Poore (2004); g, Milne Edwards and Lucas (1841); k, Sakai (1971). ­Figure 13.22a, g, j, Milne Edwards and Lucas (1841); i, Poore (2004); l, Bouvier (1895). F ­ igure  13.23a, illustration by J.E. Benedict in McLaughlin (1974); g, Anker and Pauly (2013). ­Figure 13.24c, d, Komai and Poupin (2012). ­Figure  13.30a, Poore (2004). Figure  13.31a, Lemaitre (1995). ­ ­ Figure  14.3a–f, Tavares (1996); g, h, Poore (2004); i, k, l, Tavares (1993). ­Figure 14.4a, f, g, k, l, m, Tavares (1996); c, e, Tavares (1991); p, Poore (2004); b, j, n, o, q–s, Tavares (1993). ­Figure 14.5a, c, MilneEdwards and Bouvier (1902); b, Poore (2004). ­Figure 14.6a, Tavares and Lemaitre (1996); c, d, f, Tavares (1993). ­Figure 14.7a, McLay (1998); b, McLay (2019); d, f, t, u, Poore (2004); f, McLay and Naruse (2019); g, l–n, s, McLay (1993); i, McLay (2001). ­Figure 14.8a, g, q, u, McLay (1993); d, e, i, Poore (2004); v, McLay (2009). ­Figure 14.11a, d, McLay and Ng (2005); b, Alcock (1901); c, McLay (1999); e, Ng and McLay (2010); f, McLay and Ng (2004). ­Figure  14.12a, b, Milne Edwards and Bouvier (1902). F ­ igure 14.13a–c, Poore (2004). F ­ igure 14.15a, Poore (2004); b, Castro et al. (2003). ­Figure 14.18a, Monod (1956); b, c, e–h, Ahyong et al. (2009); i, Takeda and Manuel-Santos (2006). F ­ igure  14.20a, Miers (1877); b, Sakai (1938); c, Griffin (1972); d, Rathbun (1911); e, Holthuis (1959); f, Guinot (1966); g, o, Manning and Holthuis (1981). F ­ igure  14.21b, Castelnau (1855). F ­ igure 14.23d, Takeda et al. (2000); l, Guinot and Segonzac (2018); m, Guinot and Hurtado (2003); n, Hessler and

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Marine Decapod Crustacea

Martin (1989; p, Guinot (1989). F ­ igure 14.24b, Poore (2004); d, g, j, Williams and Child (1989); e, f, Galil and Clark (1996); h, i, Galil (1993). F ­ igure  14.25a, b, e, Poore (2004). Figure  14.26b–e, Galil and Clark (1994). ­ ­ Figure  14.28a, Poore (2004); f, Davie and Ng (2012). ­Figure  14.29a, Sakai (1976). ­Figure 14.30a, b, Benedict (1892). ­Figure 14.33a–c, e, f, h, Holthuis and Manning (1990); d, Monod (1933); g, Poore (2004); i, Monod (1956). ­Figure  14.36a, b, Rathbun (1930). ­Figure 14.37, Poore (2004). ­Figure 14.38a, Bianconi (1851). ­Figure  14.39a, Alcock (1899); b, g, Poore (2004); e, Yang et al. (1998); h, McCoy (1890). ­Figure  14.40a, Garth (1940); b–g, Thoma et al. (2012). ­Figure  14.41a, b, Ng and Rahayu (2014). ­Figure 14.46a, Shen (1932); b, Alcock (1902). Figure  14.49a, Poore (2004); c, d, Guinot et al. (2013). ­ ­Figure  14.51a, Tavares and Melo (2010); c, Castro and Ng (2008). ­Figure 14.52a, b, Ng and Castro (2013). F ­ igure 14.54a, Ng and Manuel-Santos (2007). F ­ igure  14.55a, Rahayu and Ng (2014); c, h, Huang et al. (2002); d, f, Poore (2004); i, Rahayu and Widyastuti (2018). F ­ igure  14.57a, d, o, r, Poore (2004); b, i, Poore et al. (2016); c, Ng and Lukhaup (2015); s, Ng (2019). F ­ igure 14.58a, b, k, n, o, r, Poore (2004); b, Guinot (2011); c, g, k, p, s, Poore et al. (2016); i, Guinot and de Mazancourt (2020); j, Husana et al. (2011); l, Naruse et al. (2008); m, Tavares and Santana (2015); g, h, t, Komai and Poore (2016). ­ Figure  14.59i, k, Poore et al. (2016). F ­ igure 14.61c, Ng et al. (2017). F ­ igure 14.62a, b, Galil and Ng (2020); c, Walker (1887); d, e, Bell (1855). F ­ igure  14.63a, i, Rathbun (1937); e, h, r, Poore (2004); g, Capart (1951); j, l, Tan and Ng (1995); k, Komai and Ng (2012); m, Miers (1886); n, Naruse et al. (2017); o, Lanchester (1900); p, Komai and Tsuchida (2014); q, Tan (1996); r, Chen (1989); t, Ovaere (1989); u, Miers (1886). F ­igure  14.64a, Miers (1886); c, Komatsu and Ng (2011); d, e, Galil and Innocenti (2019); f, l, Rathbun (1937); h, Melo and Torres (1996); i, j, Komatsu and Takeda (2001); k, Poore (2004); l, Rathbun (1937); m, Ng et al. (2018); n, Komatsu and Takeda (2003); r, Dai and Yang (1991); s, t, Tan and Ng (1995); u, Sakai (1980). ­Figure 14.65a, Ng et al. (2018); b, k m, n, Rathbun (1937); c, d, Galil (2009); e, Bell (1855); f, Galil (2015); g, Tan and Ng (1993); h, Capart (1951); i, Komai et al. (2019); j, Galil (2009); n, o, Melo (1998); p, Tan and Ng (1995); q, Poore (2004); r, s, Rathbun (1937). Figure  14.66x, Komai and Ng (2012); z, Tan (1995). ­ Figure  14.66x, Komai and Ng (2012); z, Tan (1995). ­ Figure  14.70a, f, g, Poore (2004); d, Chen (2002). ­ Figure  14.71b–d, Rathbun (1925); e, Lee and Richer de ­ Forges (2019); f, Miers (1880); g, Garth (1973); h, Alcock and Anderson (1898); i, Poore (2004). ­Figure  14.72a–c, e, g, h, Rathbun (1925); d, Miers (1886); f, Hendrickx and Cervantes (2003). F ­ igure 14.73a, g, Rathbun (1925); b, Garth (1940); c, Miers (1886); d, j, A. Milne-Edwards (1873–1881); e, Stimpson (1860); f, Colavite et al. (2020); h, Ng & Richer de Forges (2013); i, Colavite et al. (2019); l, Garth (1952). ­Figure 14.74a,

Tavares and Santana (2011); b, Rathbun (1925); c, Griffin and Tranter (1986). ­Figure  14.75b, g, j, k, w, Griffin and Tranter (1986); l, Wagner (1986); p, q, Richer de Forges and Ng (2009). ­Figure 14.76a, d, h, p, Griffin and Tranter (1986); b, Richer de Forges and Ng (2009); c, Targioni-Tozzetti (1877); f, Hendrickx and Cervantes (2003); i–k, Poore (2004); n, Tavares and Santana (2018); u, Lee et al. (2019); v, w, Hosie and Hara (2016). ­Figure 14.77d, Forest (1966); f, p, q, Griffin and Tranter (1986); g, Tavares and Santana (2018); j, Tavares and Santana (2018); l, Richer de Forges and Ng (2009); m, Rathbun (1925); s, Lee et al. (2019). ­Figure 14.78a, c–e, g–i, k, p, r–v, Griffin and Tranter (1986); q, Lee et al. (2020); w, Ng and Richer de Forges (2013). ­Figure 14.79m, n, Tavares and Santana (2011). ­Figure 14.80a, e, i, Poore (2004); b, c, Milne-Edwards and Bouvier (1878); d, Garth (1939); g, Carpart (1951); h, Alcock and Anderson (1895); j, MilneEdwards (1873–1878). ­ Figure  14.81b, Forest and Guinot (1966); d, Garth (1935); p, r, t, Griffin and Tranter (1986). ­Figure  14.82d, Griffin (1974); e, g, h, Griffin and Tranter (1986). F ­ igure  14.83a, Rathbun (1925); b, Guinot and Van Bakel (2020); c, Guinot (2012); d, Poore (2004); e, Monod (1956). ­ Figure  14.84o, Guinot and Van Bakel (2020). ­Figure 14.85a, Baker (1905); b, Forest and Guinot (1961); e, f, h, l, Poore (2004); f, Rathbun (1925); g, Sakai (1965); i, k, Ng and Richer de Forges (2015). ­Figure 14.86b, Rathbun (1925); c, j, k, m, n, s, Poore (2004); f, h, Richer de Forges and Ng (2007); i, q, Griffin and Tranter (1986). ­Figure  14.88c, Ng and Richer de Forges (2015). ­Figure 14.89a, d, f, g, i, Rathbun (1925); b, Windsor and Felder (2011); c, e, Wagner (1990); h, Lemos de Castro (1953). F ­ igure  14.91h, Poore (2004). F ­ igure  14.94a, Shen (1932). F ­ igure  14.96d, Moosa and Serène (1981); h, Poore (2004). ­Figure  14.99a, Poore (2004). F ­ igure 14.103c, McLay (2009). F ­ igure 14.104a, e, Ng et al. (2015); d, Guinot (1985). F ­ igure  14.106a, d, e, Poore (2004). F ­ igure 14.108a, d, f, Poore (2004); b, Monod (1956); c, Garth (1986); e, Williams (1984). F ­igure  14.109h, Ng (2010). F ­ igure 14.112a, Ng & Lin (2015). F ­ igure 14.115, Crosnier and Moosa (2002). F ­igure  14.116a, Poore (2004). Figure  14.117a, Alcock and Anderson (1899); b, Poore ­ (2004). ­Figure  14.118a, Spiridonov (2020). ­Figure  14.119a, Poore (2004). F ­ igure  14.121a, k, l, Manning and Holthuis, 1981; b, e, Poore (2004); i, Guinot (1961). ­Figure  14.123j, Menoza and Devi (2017). F ­igure  14.125c, Leene (1940). Figure  14.127a, Rathbun (1930); b, Alcock (1899). ­ F ­ igure 14.128a, Zupolini et al. (2017); b, Capart (1951); d, i, Nguyen and Ng (2010); f, Koch et al. (2015). ­Figure 14.131, McCoy (1890: pls 179, 180). F ­ igure 14.132a, Naruse and Ng (2014). ­Figure  14.133a, Capart (1951); b, Ng and Ahyong (2013). F ­ igure  14.137a, Garth (1939); b, Borradaile (1902); d–f, Serène (1984). F ­ igure  14.140a, Rathbun (1930); b, d–f, Tavares and Cleva (2010); c, Dell (1972). F ­ igure  14.142a, Mendoza et al. (2012). ­Figure  14.145a, Vázquez-Bader and

Credits

Gracia (1991); b, Chace (1940). ­Figure  14.149a, b, Poore (2004); f, Serène (1984); g, Ng (2014); j, Odhner (1925); k, Komatsu and Takeda (2011). F ­ igure 14.150a, Mendoza and Ng (2012); c, Takeda & Miyake (1968); d, Milne-Edwards (1865); f, Davie (1997); i, j, Guinot (1969). F ­ igure  14.151b, Naruse et al. (2020); c, d, Ng & Chia (1994). F ­ igure 14.153f, Clark (1999); i, Dai and Yang (1998). ­Figure 14.155a, Poore (2004); c, Crosnier (1994). ­Figure 14.156b, Williams (1984); c, d, Ng (1993); h, i, Mendoza et al. (2009); j, k, Mendoza and Ng (2008); m, Ng and Clark (2003). F ­ igure 14.157a, ManuelSantos and Ng (2007); b, Mendoza et al. (2009); e, Forest (1976); k, Takeda and Komatsu (2011); n, p, Mendoza and Ng (2008); o, Mendoza and Ng (2012). F ­ igure 14.159a, Mendoza and Manuel-Santos (2012). ­ Figure  14.161a, Hsueh (2020); c, Poore (2004). F ­ igure  14.162d, Naruse and Fujita (2015). ­ Figure  14.163d, Mendoza and Ng (2011). ­Figure 14.164a, b, d, Garth (1986); c, Boone (1927); f, Monod (1956); i, Lasley et al. (2010); j, Devi et al. (2019). ­Figure  14.165e, Ng and Clark (2003); f, Poore (2004); l, Alcock (1899). ­Figure  14.169e, Naruse and Yoshida (2018). ­Figure 14.170a, e, f, Kropp and Manning (1987); b–d, g–m, o, q–s, Kropp (1990); n, Badaro et al. (2012); p, Kropp and Manning (1996). F ­ igure  14.171a–e, h, i, Kropp (1990); f, g, Kropp (1987); j, k, Kropp and Manning (1996). ­Figure 14.172a, g, j, Kropp and Manning (1987); b, c, e, f, h, i, k–m, o–s, Kropp (1990); d, Badaro et al. (2012); n, Kropp and Manning (1996). ­Figure  14.174a–d, f, Poore (2004). ­Figure 14.176a, c, d, Poore (2004); e, edited from Schubart et al. (2010). ­Figure 14.177a, Abele (1992); b, f, Naruse and Ng (2020); c, Lee et al. (2015); d, Shahdadi et al. (2018); e, Davie and Ng (2007). F ­ igure  14.178a, Shahdadi et al. (2018); b, Schubart et al. (2020); c, d, Naruse and Ng (2020); e, Abele (1992); f, Davie and Ng (2007); g, Nagai et al. (2011). ­Figure  14.179c, Naruse and Ng (2012); d, Schubart et al. (2020); g, Ng and Davie (2011); k, Poore (2004); l, Ng and Schubart (2017); n, q, Naruse and Ng (2020); m, Davie and Ng (2007); p, McLay and Ryan (1990). ­Figure  14.180a, u, Abele (1992); c, Lee et al. (2015); d–f, h, l–o, t, Shahdadi et al. (2017); g, q, r, s, Naruse and Ng (2020); p, Schubart et al. (2020); q, Davie (1992); v, Naruse and Ng (2019); w, Ng et al. (2019); z, Nagai et al. (2011). ­Figure 14.181a, Ng and Wowor (2019); b, Schubart et al. (2009); n, Schubart et al. (2020). ­Figure  14.183c–e, g, h, Poore (2004); i, Ng et al. (2018); j, Rathbun (1918). F ­igure  14.187, Stebbing (1893: pl. iii). ­Figure  14.191a, Ahyong (2014); b, Manning and Holthuis (1981). F ­ igure  14.195a, b, d, e, Kemp (1919). ­Figure  14.196, Poore (2004). ­Figure 14.197a, b, Poore (2004). ­Figure 14.199, Poore (2004). F ­ igure 14.200a, Shih et al. (2010); b, Shih et al. (2016); c, Shih et al. (2015); d, Shih (2015). ­Figure 14.201a, c, Khot et al. (2019). ­Figure  14.202a, Campos (2002); b, e, k, Ahyong (2018); c, Campos (2009); d, q, Chopra (1931); f, g,

889

Ahyong & Ng (2007); h, Ahyong (2010); j, Ng et al. (2019); m, Garth (1957); n, Poore (2004); o, p, r, s, Ahyong & Ng (2005). Unless in the public domain, cleared for copyright or uncontactable, permission were obtained from these journal publishers: Allan Hancock Foundation, Bishop Museum Press, Brill, CSIRO, Elsevier Masson SAS, Museu de Ciências Naturalis, Porto Alegre, Brazil, Royal Belgian Institute of Natural Science, Lee Kong Chian Natural History Museum, Singapore, Muséum Nationale d’Histoire Naturelle, Paris, Magnolia Press, Marine Biological Association of the United Kingdom, National Science Museum, Tokyo, Institut de recherche pour le développement, Marseille, Oxford University Press, Taylor & Francis, and University of Hawaii Press. Some ­ Figures from Poore’s (2004) Marine decapod Crustacea of Southern Australia are reproduced here. Those by Kate Nolan are: Figs 3.1a, 3.7a, 3.10a, 3.10a, 3.11a, 4.2a, 6.8d–g, 6.12, 6.14a, b, 6.21a, 6.24a, 6.26a, 6.28c, 6.29a, d, f, g, 6.31a, d, 6.46a, 6.49a, b, d, f, 6.51a, 14.3g, h, 14.4p, 14.5b, 14.7d, f, t, u, 14.8d, e, i, 14.13, 14.15a, 14.24b, 14.25a, b, e, 14.28a, 14.33g, 14.37, 14.39b, g, 14.55d, f, 14.58a, b, k, n, o, r, 14.63e, h, r, 14.64j, k, 14.65q, 14.70a, f, g, 14.71i, 14.76i–k, 14.80 a, e, i, 14.85e, f, h, l, 14.86c, j, k, m, n, s, 14.99a, 14.106a, d, e, 14.108a, d, f, 14.116a, 14.119a, 14.149a, b, 14.155a, 14.165f, 14.174a–d, f, 14.176a, c, d, 14.179j, q, 14.183c–e, g, h, 14.196, 14.197a, b, 14.202n. Those by Graham Milledge are: Figs 7.2a, b, 10.1a, 10.3a, d–g, i–k, n, o, q, 11.3d, f, 12.1c, d, 13.11b, e, f, i, 13.16, 13.18a, d, f, h, 13.21f, 13.22i, 13.30a. Photographs or drawings were provided by the following photographers and are reproduced in grayscale with their kind permission. All photographers retain copyright unless noted: Arthur Anker, 14.61b, 14.63b, c, f, 14.64g, m, q, 14.65l, t, 14.70b, 14.77e, 14.143h, 14.166d. Shane T. Ahyong, 14.85c, d. R. Cleva/J. Poupin/MNHN©, 14.26a. Darryl L. Felder, 14.18d. T.H. Hui and T. Naruse, 14.165d. Tin-Yam Chan/ MNHN©, 3.4j, 14.149e, h, l, 14.150b, e, g, 14.152b, 14.157j, 14.162c, e, 14.165i, k, 14.168c, g–i. R. Lasley, 14.153a–e, 14.157m, 14.165j. B.Y. Lee, 14.71e. J.C. Mendoza, 14.157f, g. M. Nattapat, 14.165a, 14.177f. Peter K.L. Ng, 14.162a. Max Orchard, 14.165c. Gustav Paulay, 14.143e, 14.164k. Joseph Poupin, 14.143b, d, g, 14.149d, 14.156e, 14.162b, f, 14.164e, 14.165h, 14.166c, f, g, 14.168e, f. E. Lazo-Wasem, 14.143a. Ondřej Radosta, 14.32a, 14.143c, 14.149i, 14.164g, 14.168a, b, d. S.H. Tan, 14.99b–d, 14.100a–l, 14.101a, c–l, 14.102a–j, 14.103c, 14.165d. Ling-Kuang Tseng, 14.149c, 14.165b. Photographs of type specimens in the Muséum Nationale d’Histoire Naturelle, Paris, are reproduced with permission as follows: Figs 14.23c, 14.73k, 14.152a, 14.156a, f, l, 14.157c, h, l, 14.164h, 14.165g, 14.166e.

890

Marine Decapod Crustacea

Plates Photographs were provided by the following photographers for the colour plates and are reproduced with their kind permission. All photographers retain copyright unless noted: Shane T. Ahyong: Pls 20c, 28a, d, 31c, 41d, 42e, 43l, 44i, 45c, e, 53c, g, 54a, 60d, i, 61d, e, o, 69i, 71d. Arthur Anker: Pls 1e, 4d, 5a–e, 6a, b, f–n, 7d, f–j, m, 9d, 10a–d, l, m, o, 11a–c, e, g, h, 12h, i, 13b, c, 17d, h, i, 18c, i, 24b, h, 25a–d, g, k, 28f, 29a, 30f, 31g, 33l, 34a, 39h, j, 40c, g, 41a, 43m, 44c, 46b, 47i, 48b, e, j, n, 50h, 51a, c, 52g, 54h, 57e, f, h, i, 58a, 60c, 61m, 64c, 65b, 67c, 69a, 71j, l. Christopher Boyko: Pls 29d, 30b. S. Campbell: Pl. 62e. Tin-Yam Chan/NTOU©: Pls 2f, 3f–h, 12e, l. Tin-Yam Chan (some by C.W. Lin)/MNHN©: Pls 2a–c, h, 3a, c–e, g, 4a–c, e–j, 5f, g, 7l, n, 8e, g, i, k, l, 9a, c, g, i, j, 10g, i, k, n, 11d, f, k, 12a–d, f, g, m, 13a–c, e, g, h, 14a, c–h, 15a, g, 16g, p, 17a, f, g, 18a, f, g, 19c, f, 20a, b, e, f, 21b, c, 22c, e, 24a, e, g, i, 25b–f, 26b–f, 27a, j, l, 28c, e, g, 29b, e, 30g, 31f, 32c, d, h, j, k, 33b, c, f, g, 34c, e, h, j, 36a–g, i, j, 37c, d, i, j, l, n, 38a., b, d, e, g, h–k, m–o, 39a–c, e, 40a, b, f, 41b, h, i, k, 42l, 43d, e, g, n, o, 44a, b, d, f–h, k–n, 45g–i, 46a, d–g, l–n, 47e, f, j, k, 48a, d, g, i, l, m, o, 49b, e, i–l, 50d, e, g, j, l, 51d, g, i–k, 52a, b, d, 53a, b, i, l, 54b, 55a–e, h, 56c, d, f, 57a, b, d, g, j, k, 59c–g, i, j, 60a, f, g, 61a, j, n, 62d, f, h, 63c–j, 64f–h, k, m, o–q, 65a, h, i, 66a, b, e, g, i, l–o, 67a, d–f, h–k, m, o, r, 68j, 69g, h, j, 70h. Laure Corbari/MNHN©: Pls 8c, 9m, 11j, 15b, 19d, 21e, 24f, 31a, 35a–d, 38c, f, l, 44j, 49c, 50i, 51e, h, 56b, 63b, k, 67b. Laure Corbari & Joseph Poupin/MNHN©: Pls 10h, j, 19e, 21d, f, 27f, i, 28b, h, 33e, j, 34b, 39i, 40d, e, 41e, g, h, 46c, i, 47a, c, 48f, 49a, g, 50a, c, f, k, 54d, f, 58c, e, 64d, e, l,

66d, 67n, 69d, 69e, 70a, b, 72g. Bernd Hoppe: Pl. 10f. Serena Cox/NIWA©: Pl. 1a. P.D.F. Davie: Pl. 72c. Zdeněk Ďuriš: Pls 14b, 15c–f, 16d, e, 18d, h, 18h. Darryl L. Felder: 16a–c, f, l–o, q, 17b, c, e, 39f A. Fifis/IFREMER©/courtesy M. Segonzac, MNHN: 23b. Julian K. Finn/Museums Victoria©: Pls 1f, 9e, 10e, 11i, 45f. Charles Fisher/WHOI©: Pl. 1c. Yoshihisa Fujita: Pl. 8b. Karen Gowlett–Holmes/CSIRO©: Pls 2d, e, 3b, 7e, 8d, h, j, 9b, h, k, l, n, 12j, k, 13e, f, 20d, 21a, 23d, 26a, 29c, 32l, 33i, k, 34d, i, k, 43f, 51b, 53e, k, 55g, 56e. K. Fujikura/JAMSTEC© and S.W. Lin/NTOU©/courtesy T.-Y. Chan: Pl. 22a, b. Hans Hillwaert: Pls 7k, 8f, 37g, 42a, c, j, 46j, 52e, 55f, 61 g, i, l, 62l, 72h. P.W. Hsueh: Pl. 30a. Tomo Komai: Pl. 8a. Robert Lasley: Pl. 67 g. Michael Marmach/Museums Victoria©: Pls. 6c, d, 7a–c, 9f, 17j, 18e, 24d, 30d, 32e, 37h, 44o, 45a, d, 46h, 47l, 52 g, 53f, 60b, e, g, j, 61c, 69f, 70k, 71a–c, f, o. G. Millen/Australian Museum©: Pl. 22d. Mos Natapatt: Pls 44e, 59a, 66h, 70c, d, e, g, i, l–o, 71e, g, h, i, k, m n, 72a, b, e, j. Paul Ng: Pl. 52c. P.K.L. Ng: Pl. 40h. S.E.T. van der Meij & B.T. Reijnen: Pl. 68a–i. Mark Norman/Museums Victoria©: Pls 37a, 61f. B. Ogata: Pls 33a. 50b. N. Ohtsuchi: Pl. 42i. Gustav Paulay: Pls 16i, 30c, 42d, 54e, i–k, 60h, 61k, 62c, j, k, 65g. Gary C.B. Poore: Pls 18b, 69b. Joseph Poupin: Pls 16h, 30e, 67p. J. Poupin/R. Cleva: Pl. 24c. Ondřej Radosta: Pls 31b, d, e, 36h, k, 37b, e, f, 39d, g, k, 41c, f, j, l, 42b, f, g, k, 43a–c, i–k, 46k, 47b, d, g, h, m, 48c, h, k, 49f, h, 51f, 53d, h, j, 54c, g, 56a, 57c, l, 58b, d, 59b, 60k, 61b, h, 62a, b, g, i, 63a, 64a, b, i, j, r, 66f, 67l, q, 69c, 70f, j, 72d, f, i, k, l. Andrey Ryanskyi: Pls 1d, 27e, h, 28i, 32a, b, f, g, i, 33d, h, 34f, g, 37j, k, 42h, 49d, 59h, 66k. Kareen Schnabel/NIWA©: Pl. 45b. Sven Thatje: Pl. 23a, c. Steve Thurston/AMNH©: Pl. 29f.

Index to genus-group, family-group and higher taxonomic names

Bold numbers refer to colour plates. Abyssotheres 866, 868, 870 Acanthacaris 175, 177–179, 13 Acanthanas 67, 69, 70 Acantharctus 254, 256 Acanthaxius 191–195, 14 Acanthephyra 116, 9 Acanthilia 556, 559, 560, 563, 567, 46 Acanthocarpus 482–484, 41 Acanthocyclus 477, 478, 40 Acanthodromia 455, 456, 36 Acanthodromiinae 455 Acantholithodes 312, 314, 315 Acantholobulus 737–740 Acanthonyx 586, 593, 594 Acanthopagurus 321, 326 Acanthophrys 586, 591, 592, 594 Acetes 41–44, 46, 47 Achaeopsis 609, 611–613 Achaeus 610–614 Achelata 9, 11, 12, 14, 15, 183, 247, 19–21 Achelouinae 701, 703, 704, 61 Achelous 702–704, 61 Acidops 512, 513 Acidopsidae 429, 510, 511, 43 Acidopsinae 513 Acmaeopleura 828, 829 Acrothoracica 12 Actaea 752–756, 64 Actaeinae 751–754, 64 Actaeodes 753–755 Actaeomorpha 474, 475 Actinimenes 127, 135, 137, 139, 142 Actiomera 774, 775 Actites 775 Actumnus 669, 675, 676 Acutigebia 242, 243, 18 Aegaeon 105, 106, 8 Aeglidae 263 Aegloidea 263 Aepinus 616–619 Aethogebia 242, 243 Aethra 474, 475, 40 Aethridae 424, 474, 40 Aethroidea 424, 474, 733

Afrophila 558, 559, 564 Afropinnotheres 866, 870 Afruca 856, 859 Agaricochirus 323, 324, 326 Ageitomaia 625–628 Agolambrus 653, 657 Agononida 278, 280, 25 Agostocaris 96, 173 Ala 635–637 Alain 865–868, 870 Alainius 273–275 Alainodaeus 765, 767–769, 66 Alainodromia 445, 447–449 Alainopaguroides 322, 326 Alainopagurus 322, 327 Alainopasiphaea 166, 12 Alainotheres 865, 866, 870, 871 Alainthesius 515, 516, 518 Alarconia 862, 865, 866, 871 Albunea 297, 29 Albuneidae 295, 296, 29 Albuneinae 296 Alcockdromia 445, 448, 449 Alcockia 595 Alcockpenaeopsis 30, 33 Alcomaja 624, 626–628, 53 Aldrovandiopanope 778, 781, 782 Aletheiana 542, 543, 545 Alfredalcockia 586, 590, 595 Aliaporcellana 288, 290, 27 Alienaxiopsis 189, 192, 194, 14 Alionectes 701, 707, 710, 711 Allactaea 752, 753, 755 Allaxiopsis 189, 192, 194, 14 Allaxius 192, 194, 195 Allodardanus 305, 307 Allogalathea 274, 275, 24 Allograea 479, 480 Allomonomia 702, 710, 711 Allomunida 274, 275 Allopetrolisthes 286–268, 290, 27 Allopontonia 125, 138, 140, 142 Allosergestes 43, 44, 46 Allotriocarida 10, 12

892

Marine Decapod Crustacea

Alope 83–85, 7 Alox 555, 560, 563, 564, 46 Alpheoidea 61, 63, 65 Alpheopsis 67, 69, 70, 72, 6 Alpheus 65, 68, 69, 70, 6 Altelatipes 24, 25 Altopontonia 130, 135, 138, 142 Alvinocarididae 62, 96, 8 Alvinocaridinides 97–99 Alvinocaris 96–98, 1, 8 Amakusaxius 191, 192, 194, 195 Amalopenaeus 24 Amarinus 541–545, 549, 45 Ambiaxiopsis 189, 192, 195, 196 Ambiaxius 189, 192, 194–196 Ambidexter 168, 169 Amboplax 515–517 Amphibetaeus 68, 69, 71 Amphionidacea 61 Amphionides 61 Amphipoda 11, 13, 14 Amphipontonia 130, 135, 138, 139, 141 Amphithrax 635–637 Amusiotheres 865, 867, 869, 871 Anacalliacidae 186, 187 Anacalliax 187 Anachlorocurtis 157 Anacinetops 622, 623, 626, 628 Anamathia 582, 594, 595 Anapagrides 321, 327 Anapagurus 321, 327 Anapontonia 128, 135, 139 Anaptychus 637 Anasimus 618, 619 Anaspidacea 11, 13, 14 Anatolikos 489, 490, 42 Anchialocarididae 63, 172 Anchialocaris 172, 173 Anchiopontonia 130, 135, 139, 140, 142, 10 Anchistioides 122, 132, 133 Anchistioididae 119 Anchistus 130, 140, 10 Ancylocaris 127, 135, 139, 140, 10 Ancylocheles 288–290 Ancylodactyla 554, 560, 563, 564, 46 Ancylomenes 127, 138, 140, 10 Andamancalliax 229, 230 Andamanium 812, 816 Angustopelta 515–517, 44 Aniculus 305–307, 32 Aniptumnus 670, 676, 677 Anisomenaeus 128, 140, 141

Anisonotus 610, 612, 613 Anisopagurus 323, 325–327 Ankerius 793, 794 Anomala 263 Anomalifrons 860, 861 Anomalifrontinae 860 Anomalothir 609, 610, 613 Anomoeomunida 277, 279 Anomura 3, 11, 12, 15, 185, 263 Anophthalmaxius 189, 192, 194–196 Anoplonida 277, 278, 280 Anostraca 11, 12 Antarctus 253, 254, 256, 20 Antecaridina 95 Antilibinia 586, 593, 595 Antrocarcininae 750, 757, 64 Antrocarcinus 757, 758 Apechocinus 542, 544, 545 Aphanodactylidae 422, 430, 792, 793, 861, 68 Aphanodactyloidea 422, 430, 792 Aphanodactylus 794 Aphareocaris 48 Apias 586, 589, 595 Apiomithrax 583, 584, 586, 595 Apograpsus 850, 851 Apopontonia 128, 135, 139, 140 Aqaballianassa 205, 211 Arabigebicula 244 Arachnochium 120, 122, 132, 133 Arachnopsis 618–620 Arafurapenaeopsis 29, 30, 33 Araiopontonia 129, 135, 138–140, 142 Aratus 809, 815, 817, 70 Arcania 552, 557, 560, 563, 565, 46 Arcotheres 867, 869–871 Arctides 253, 255 Arctidinae 252, 253, 255, 20 Arenaeus 702, 709–711, 62 Arenallianassa 205–207, 210, 211 Areopaguristes 305, 307, 12 Arete 67, 69, 71 Aretopsis 67, 69, 71, 72 Argis 104–106 Aristaeomorpha 21–23, 2 Aristaeopsis 21, 22, 2 Aristeidae 19–21, 27, 2 Aristeus 21, 22, 2 Aristotelopanope 778, 781, 782 Armases 810, 812, 817, 70 Artemesia 29, 31, 33 Ascidiophilus 445–447, 449 Ascidonia 130, 135, 139, 140

Index to genus-group, family-group and higher taxonomic names

Ascothoracica 12 Ashtoret 486, 41 Astacidae 175 Astacidea 11, 12, 15, 175, 183, 13 Astacus 175 Asthenognathinae 825, 861 Asthenognathus 825 Atelecyclidae 426, 477, 487, 488, 493, 42 Atelecyclus 487, 488, 42 Atergatis 787, 788, 67 Atergatopsis 787, 788, 67 Athanas 67, 69, 73 Athanopsis 67, 69, 71, 73, 6 Atlantolocia 558, 564, 565 Atlantopandalus 162 Atlantophila 558, 564, 565 Atlantotlos 554, 560, 564, 565 Atoportunus 701, 705 Atyidae 61, 62, 95 Atyoidea 62, 95 Atypopenaeus 27, 29–31, 33, 3 Audacallichirus 220–222 Aulacolambrus 653, 657, 660, 57 Austinixa 862, 865, 866, 871 Austinogebia 241–243 Austinograea 479–481 Austinotheres 864, 866, 868, 871 Australeremus 325, 331 Australocarcinus 514, 515, 518 Australocaris 190, 192, 195, 197 Australoplax 850–852 Austrodromidia 445, 447–449, 37 Austrohelice 826, 827 Austrolepidopa 296, 297, 29 Austrolibinia 586, 591, 595 Austropandalus 161–163 Austropenaeus 21, 22, 2 Austrotheres 864, 866, 870, 871 Austruca 856, 857, 72 Automate 68, 69, 71, 73 Axianassa 238, 239, 18 Axianassidae 237 Axiidae 185, 187, 14, 15 Axiidea 12, 14, 15, 185, 237, 342, 14–17 Axiopsis 192, 193, 196, 197 Axiorygma 190, 192, 195, 197 Axius 188, 190, 192, 197 Babamunida 278–290 Baccadromia 449 Balssaxius 198 Balsscallichirus 221, 222

Balssia 123, 135, 141 Balssopilumnus 668, 675, 677 Banareia 758, 759, 64 Banareiinae 750, 758, 64, 65 Bannereus 66, 69, 73 Baptozius 506, 507 Barbouria 80, 81 Barbouriidae 64, 80 Barbouriinae 81 Barnardromia 445, 447, 449 Baruna 832–842, 71 Batella 66, 69, 72, 73 Batepenaeopsis 30, 33 Bathyarctus 254, 256 Bathycalliax 218 Bathycheles 243, 244 Bathyhippolyte 82 Bathymenes 127, 135, 142 Bathymunida 278–280, 25 Bathynarius 306, 307 Bathynectes 697–699 Bathynellacea 13, 14 Bathypaguropsis 319, 323, 327, 33 Bathypalaemonella 102, 103, 8 Bathypalaemonellidae 63, 102, 8 Bathypalaemonetes 102, 103 Bathypilumnus 669, 676–678 Bathyplax 427, 510, 523–527 Bathypluma 724, 725, 63 Bathyrhombila 742–745 Bathystylodactylus 171 Batodaeus 765, 767–769 Batrachonotus 617, 619, 620 Bellia 477, 478 Bellidilia 558, 560, 563–565, 46 Belliidae 422, 425, 432, 477, 40 Bellioidea 425, 428, 477 Belzebub 41, 42 Bentheogennema 24, 25 Benthesicymidae 19, 20, 23, 27, 2 Benthesicymus 24–26 Benthochascon 689, 691–693, 697 Benthonectes 24–26 Benthopanope 670, 675, 677, 678 Bermudacaris 68, 69, 71, 73 Betaeopsis 68, 69, 72, 73 Betaeus 66, 69, 71, 73, 6 Beuroisia 531, 532 Biarctus 254–256 Biffarius 206, 211, 16 Birgus 302, 303, 30 Birulia 92, 93

893

894

Marine Decapod Crustacea

Bitias 161–163 Bivalvopagurus 340, 341 Blepharipoda 299, 29 Blepharipodidae 295, 296, 298, 29 Blepharocaris 126, 135, 143 Boasaxius 190, 192, 194, 197 Bochusacea 13, 14 Boninpagurus 321, 322, 324, 327 Bonita 864, 866, 867, 871 Bossacarcinus 670, 675, 677, 678 Bothromaia 578, 639–643 Bottoxanthodes 778, 781, 783 Bountiana 506–507 Bouvieraxius 189, 192, 195–197, 14 Brachycarpus 121, 133, 10 Brachygnatha 421 Brachynotus 831–833, 71 Brachypoda 12 Brachyrhyncha 421 Brachyura 3, 9, 11, 12, 15, 263, 421 Branchiopoda 9, 10, 12 Branchiura 10, 12 Brankocleistostoma 828, 829 Bresedium 815–817 Bresilia 100, 8 Bresiliidae 62, 95, 99, 100 Bresilioidea 61, 62, 95 Bruceaxius 200 Brucecaris 126, 135, 142, 143 Bruceonia 120, 129, 135, 143 Bruceopsis 67, 70, 72, 73 Bruciana 774, 775, 66 Brusinia 689, 697, 61 Brusiniidae 426, 687–689, 697, 61 Buergeres 864, 866–868, 871 Bythiopagurus 319, 322, 323, 326, 327 Bythocarides 82 Bythocarididae 64, 82 Bythocaris 82 Bythograea 479, 480, 40 Bythograeidae 426, 479, 40 Bythograeoidea 426, 478 Caecopilumnus 512–514 Caenopedia 534, 535 Cainonia 130, 135, 143 Calabarium 840–842 Calappa 482–484, 41 Calappidae 424, 474, 481, 482, 41 Calappoidea 422, 424, 481, 646 Calappula 483, 484 Calastacus 189, 192, 196, 197 Calathaemon 121, 132, 133

Calaxiopsis 189, 192, 198 Calaxius 190, 192, 194, 195, 198, 14 Calcinidae 301, 302, 304 Calcinus 302, 304, 306, 307, 32 Caligoneus 68, 70–72, 74 Callianassa 205–207, 209, 211–214, 223, 224 Callianassida 185 Callianassidae 185, 186, 204 Callianassoidea 185 Callianidea 216, 217, 16 Callianideidae 186, 215, 231, 16 Callianopsidae 186, 217, 228 Callianopsis 218 Calliapagurops 219, 220, 222 Calliasmata 80, 81 Calliasmatinae 81 Calliax 228–230 Calliaxina 229, 230, 17 Callidactylus 556, 560, 565, 46 Callinectes 687, 700, 702, 709–711, 62 Calmania 667, 672 Calmaniinae 666, 667, 672 Calocarcininae 728 Calocarcinus 728–730, 63 Calocarides 191, 192, 195, 198 Calocarididae 188 Calocaris 188, 192, 196, 198 Calocarisopsis 198, 200 Calvactaea 758, 759, 64 Calypsachaeus 609, 611, 613 Calyptraeotheres 863, 865, 872 Camatopsis 515–517, 44 Cambaridae 2, 175 Camilohelleria 778, 781–783 Camposcia 611–613, 52 Camptandriidae 430, 432, 433, 837, 838, 71 Camptandrium 839–843, 845, 71 Camptoplax 671, 682 Campylonotidae 63, 102, 103, 8 Campylonotoidea 63, 102 Campylonotus 103, 8 Cancellocheles 343–345 Cancellus 302, 305, 306, 308, 32 Cancer 2, 432, 487–491, 42 Cancridae 425, 432, 487, 488, 42 Cancrocaeca 541, 542, 544, 546 Cancroidea 426, 487 Capartiella 610, 612, 613 Caphyra 700, 702, 712–714 Caphyrinae 700, 713 Capilliporcellana 288, 289, 291 Carcinidae 426, 432, 687–690, 694, 696, 61 Carcinoplax 523–526, 533, 736, 44

Index to genus-group, family-group and higher taxonomic names

Carcinus 690, 61 Cardisoma 802 Cardus 259, 260 Caridea 3, 9, 11, 12, 14, 15, 51, 59, 61 Caridion 84, 85 Carinopontonia 124, 135, 143 Carolinaxius 191, 192, 198 Carpiliidae 428, 432, 491, 42 Carpilioidea 428, 491, 733 Carpoporus 764, 767, 769 Carupa 701, 704–706, 61 Carupella 700, 707 Carupinae 700, 701, 704, 708, 61 Cataleptodius 778, 779, 782, 783 Catapaguroides 319, 324, 328 Catapaguropsis 321, 326, 328 Catapagurus 321, 328 Cateios 555, 556, 564, 565 Catoptrinae 700 Catoptrus 701, 705, 706 Caviallianassa 205–207, 209–211 Cavoportunus 702, 709–711 Cebudonus 668, 672, 673 Cecidocarcinus 796, 798, 799 Celatopesia 653, 657, 658, 660 Cephalocarida 9, 10, 12 Cerataspis 21–23, 2 Ceratocarcinus 668, 672, 673, 59 Ceratopagurus 322, 323, 328 Ceratoplax 671, 683, 59 Certolambrus 655, 657, 658, 660 Cervimunida 277–280, 25 Cestopagurus 320, 323, 324, 328, 335 Chacella 123, 135, 143, 151 Chacellus 742, 744, 745 Chaceon 531, 691–693, 61 Chaenostoma 850–852 Chalaroachaeus 611, 612, 614 Challengerosergia 45, 46 Chanopagurus 319, 322, 324, 328 Charybdis 687, 700, 703, 712–714, 62 Chasmagnathus 826, 827 Chasmocarcinidae 429, 431, 510, 514, 44 Chasmocarcininae 514, 515, 517 Chasmocarcinops 515–517, 44 Chasmocarcinus 515–517 Chasmophora 736, 741, 742, 744, 746 Cheiragonidae 425, 432, 492, 42 Cheiragonoidea 425, 492 Cheiragonus 492, 493 Cheiroplatea 343, 344, 35 Chelarctus 254–256, 20 Chelonika 161–163

Cheramoides 206, 207, 210, 211 Cheramus 206, 207, 211, 212, 214 Cherusius 726, 727 Chicosciencea 54 Chinommatia 515–517 Chionoecetes 578, 640–643 Chiromantes 813–815, 817 Chirostylidae 263, 266, 271, 22 Chirostyloidea 263–265, 272 Chirostylus 267, 268 Chlorocurtis 159, 11 Chlorodiella 760–762, 65 Chlorodiellinae 750, 759, 65 Chlorotocella 158, 159, 11 Chlorotocellidae 63, 158, 11 Chlorotocoides 160–163, 12 Chlorotocus 160, 162, 163, 12 Choniognathus 625, 627, 628 Chorilia 582, 591, 593, 595 Chorinachus 611, 612, 614 Chorinus 582, 593, 595, 49 Chorismus 84–86 Chorocaris 99 Christmaplacidae 427, 720 Christmaplax 720, 721 Ciliopagurus 305–308, 32 Cinetorhynchus 114, 9 Circulium 810, 816, 817 Cirripedia 11, 12 Clastotoechus 286–288, 290, 291 Cleistostoma 839–843 Clibanarius 306–308, 32 Climeniperaeus 128, 135, 141, 143 Clistocoeloma 809, 812, 817, 70 Clypeasterophilus 863, 866, 872 Clythrocerus 435–437 Clytomanningus 169 Coelocarcininae 700, 706, 707 Coelocarcinus 687, 688, 700, 707 Coelocerus 582, 590, 595 Coenobita 303, 304, 30 Coenobitidae 264, 302, 303, 30 Coenophthalmus 688, 691–693, 697 Colemonia 130, 135, 143 Colerolumnus 668, 675–678 Coleusia 553, 574, 48 Collodes 595, 617–620 Collodinae 620 Conchodytes 127, 135, 143, 10 Conchoecetes 444, 445, 447, 449, 37 Conleyidae 428, 432, 511, 519 Conleyus 519 Contusarma 815–817

895

896

Marine Decapod Crustacea

Copepoda 11, 12 Coralaxius 190, 192, 194, 198 Corallianassa 219-223, 16 Coralliocaris 127, 135, 142, 143, 10 Coralliocryptus 554, 560, 563, 566 Coralliogalathea 274, 275 Coralliope 778, 781–783 Coriollianassa 205–207, 212 Cornutosergestes 43, 44, 46 Coronalpheus 68, 70, 74, 6 Corycodus 435–437, 36 Coryrhynchus 611, 612, 614 Corystes 421, 493, 494, 42 Corystidae 425, 492, 493, 42 Corystoidea 425, 492 Corystoides 477, 478 Cosmonotus 469–471, 39 Costalambrus 654, 657, 659 Coutieralpheus 67, 70–72, 74 Coutierea 123, 138, 139, 143 Cranaothus 765, 768, 769 Crangon 104–106, 8 Crangonidae 62, 103, 8 Crangonoidea 62, 103 Cranuca 856, 857 Creaseria 120, 121, 132, 133 Crenarctus 253–256 Crinitocinus 512, 513, 43 Crinotonia 126, 135, 138, 141–143 Criocarcinus 586, 589, 596, 49 Cristarma 812, 817 Cristimenes 125, 135, 141, 144 Crocydocinus 582, 593, 596, 49 Cronius 703, 712–714 Crosniera 216, 217 Crosnierita 277–280 Crosnierius 767–769 Crosnierocaris 66, 74 Crossotonotidae 424, 647 Crossotonotinae 647 Crossotonotus 647 Crustaenia 542–544, 546 Cryphiops 122, 132, 133 Cryptocheles 82, 83 Cryptochiridae 430, 432, 795, 68 Cryptochiroidea 430, 795 Cryptochirus 796, 798, 799 Cryptocneminae 552, 553, 559, 46 Cryptocnemus 552, 553, 558, 559, 652, 46 Cryptocoeloma 671, 683 Cryptodromia 445, 447–450, 37 Cryptodromiopsis 445, 447, 448, 450, 451

Cryptolithodes 312, 313, 314, 31 Cryptolutea 671, 682, 683 Cryptopenaeus 39, 40 Cryptopilumnus 670, 675–678 Cryptopodia 652, 653, 657, 659, 660, 57 Cryptosoma 483, 484, 41 Ctenocheles 226, 227 Ctenochelidae 186, 187, 225, 231, 17 Ctenocheloides 226, 227, 17 Ctenopontonia 124, 135, 144 Cuapetes 126, 136, 144 Culexisoma 542, 543, 546 Cumacea 13, 14 Curupironomus 439, 440 Cyanagraea 479, 480 Cycetopagurus 321, 324, 328 Cyclax 623, 626–628, 53 Cycloachelous 702, 709–711 Cyclocoeloma 582, 589, 596, 49 Cyclodius 760–762, 65 Cyclodorippe 436, 437 Cyclodorippidae 423, 431, 434, 435, 36 Cyclodorippinae 435, 436 Cyclodorippoida 421, 423, 434 Cyclodorippoidea 423 Cycloes 483, 484, 41 Cyclograpsinae 431, 432, 825 Cyclograpsus 431, 824, 826, 827, 829, 71 Cyclonyx 581, 586, 596 Cycloplax 736, 741, 742, 744, 746 Cyclorma 810, 812, 815, 817 Cycloxanthops 777, 780, 782, 783, 785 Cyclozodion 483, 485, 41 Cymo 762, 763, 65 Cymoinae 750, 762 Cymonomidae 423, 432, 434, 439, 36 Cymonomoides 440 Cymonomus 439, 440, 36 Cymopolia 648, 650 Cymopolus 439, 440 Cyphocarcinus 586, 589, 589 Cyrtocarcinus 757, 758, 64 Cyrtograpsus 831, 832 Cyrtomaia 578, 639–642, 55 Cyrtoplax 742, 744–747 Cyrtorhina 469–471 Cyrtorhininae 469–471 Dacryomaia 796, 797, 799, 68 Dacryopilumnidae 428, 494, 500, 501 Dacryopilumnus 501, 502 Dactylonia 130, 136, 138, 142, 144

Index to genus-group, family-group and higher taxonomic names

Dagnaudus 460–461 Dairidae 429, 494, 495, 43 Dairoidea 429, 494, 501, 733 Dairoides 650–652, 57 Dairoididae 429, 494, 500, 651, 57 Daldorfia 653, 655, 656, 57 Daldorfiinae 652, 655, 656 Damithrax 638 Danarma 812, 813, 818 Danielea 765, 767–769, 66 Danielella 840–843 Danielethus 508, 509 Danielita 843 Danielum 669, 675, 677, 678 Dardanus 72, 304–308, 32, 33 Dasella 125, 136, 142, 144 Dasycaris 125, 136, 144 Davusia 807, 808 Dawsonius 226, 227, 17 Decaphyllus 318, 319, 324, 328 Dehaanius 594 Deilocerus 434–437 Deioneus 67, 70, 71, 74 Deiratonotus 840–843 Delsolaria 580, 582, 590, 591, 596 Deltopelta 515–517 Demania 777, 778, 780–783, 67 Dendrobranchiata 3, 9, 11, 12, 14, 15, 19 Dentalopagurus 320, 324, 326, 329 Dentoxanthus 665, 666 Deosergestes 43, 44, 46 Derilambrus 656, 657, 660 Dermaturus 312, 314, 315 Desmocarididae 2, 61, 63, 119 Desmocaris 119 Desmodromia 445, 448, 450 Detocarcinus 796, 799 Diacanthurus 324–326, 329 Diapontonia 129, 136, 144 Dichelopandalus 161, 163 Dicranodromia 457, 458, 38 Dinochelus 177–179, 13 Diogenes 306–308, 33 Diogenidae 301, 302, 304 Diplostraca 12 Disciadidae 62, 96, 100 Discias 100, 101 Discoplax 802 Discorsopagurus 325, 326, 329 Discorsotheres 863, 864, 866, 867, 872 Dissodactylus 863, 866, 867, 872 Distolambrus 653, 657, 660

Doclea 586, 596, 49 Dolos 555, 560, 564, 566 Domecia 726, 727, 63 Domeciidae 430, 725, 726, 63 Dorhynchus 609, 611, 614 Dorippe 496, 497, 43 Dorippidae 424, 495, 496, 43 Dorippoidea 424, 495 Dorippoides 496, 497, 43 Dorodotes 61, 160, 161, 163, 12 Dorphinaxius 190, 192, 199 Dotilla 846, 847, 71 Dotillidae 433, 838, 845, 71 Dotilloplax 846, 847 Dotillopsis 846, 847 Drachiella 474, 475, 40 Dromia 443, 444, 445, 447, 450, 453, 37 Dromiacea 421–423, 442 Dromidia 445, 447, 448, 450, 452 Dromidiopsis 444, 445, 447, 448, 450, 452, 37 Dromiidae 423, 432, 442, 443 Dromiinae 443, 449 Dromioidea 423, 442, 443 Dumea 579, 608, 610–612, 614, 52 Durckheimia 864–866, 872 Dynomene 455, 456, 36 Dynomenidae 423, 442, 455, 36 Dynomeninae 455, 456 Dyspanopeus 737, 738 Ebalia 552–554, 560, 563, 566, 571, 46 Ebaliinae 552, 553, 559 Ebaliopsis 556, 560, 566 Echidnocerus 313–315 Echinoecus 668, 672, 673 Echinolatus 688, 691–693, 695 Echinopericlimenes 127, 136, 139, 142, 144 Ecphantor 839–843 Ectaesthesius 728–730, 777 Eduarctus 253, 254, 256, 20 Edwardsium 765, 768, 769, 66 Eiconaxiidae 188 Eiconaxius 188, 189, 191, 192, 195, 196, 199, 14 Elamena 542–544, 546, 45 Elamenopsis 540–542, 546 Elassochirus 325, 326, 329 Elassopodus 439, 440 Emerita 299, 300, 30 Enallopaguropsis 325, 326, 329 Enallopagurus 325, 329 Encantada 100 Encephaloides 610, 613, 614

897

898

Marine Decapod Crustacea

Eneosesarma 812, 816, 818 Engystenopus 53, 54 Enigmaplax 850–852 Enigmatheres 866–868, 872 Enneobranchus 318, 320, 324, 326, 329 Enneopagurus 320, 324, 326, 329 Enneophyllus 320, 322, 330 Enoplolambrus 656, 657, 660, 57 Enoplometopidae 175, 176 Enoplometopoidea 175 Enoplometopus 175, 176, 13 Enosteoides 288, 289, 291 Enriquea 277–280 Entomonyx 625, 628, 629 Entricoplax 524, 526, 527 Eodemus 701, 710, 711 Eodromia 444, 445, 453 Ephippias 609, 610, 614 Ephyrina 115, 116, 9 Epiactaea 752, 753, 755, 64 Epiactaeodes 753–755 Epialtidae 425, 431, 578–580, 49–51 Epialtinae 578, 580 Epialtoides 581, 586, 596 Epialtus 581, 586, 596, 49 Epigodromia 445, 448, 450 Epigrapsus 802 Epiloboceridae 425 Epipedodromia 445, 446, 450 Epipontonia 130, 136, 144 Episesarma 810, 814, 818, 70 Epistocavea 765, 768, 769 Epixanthoides 506, 507 Epixanthops 780, 782, 783 Epixanthus 506, 507, 43 Eplumula 465, 38 Ergasticus 578, 639, 640, 642, 643 Ericerodes 611, 613, 614 Ericiolumnus 669, 675, 678 Erileptus 617–620 Erimacrus 492, 493, 42 Eriocheir 824, 831, 832 Eriphia 502, 503, 43 Eriphides 502, 503 Eriphiidae 428, 432, 500–502, 43 Eriphioidea 427, 428, 500 Ernestotheres 867, 870, 872 eryoneicus 259, 260 Esopus 617–620 Ethusa 498, 499, 43 Ethusidae 424, 495, 498, 43 Ethusina 498–500, 43

Etisinae 750, 763 Etisus 763, 764, 66 Eualus 92, 93, 7 Eubrachyura 421, 422, 473, 495 Eucalastacus 188, 193, 194, 199 Eucalliacidae 186, 217, 228, 17 Eucalliaxiopsis 228–230, 17 Euceramus 286, 288, 291, 27 Euchirograpsus 807, 808, 69 Eucinetops 609, 611, 612, 614 Euclosia 574 Euclosiana 553, 574, 48 Eucrate 520, 521 Eucratodes 741, 742, 745–747 Eucratopsis 736–738 Eudromidia 445, 446, 448, 450 Eugonatonotidae 62, 111, 9 Eugonatonotus 111, 9 Eulenaios 288, 291 Eumanningia 83, 85, 86 Eumedoninae 666, 667, 672, 674 Eumedonus 668, 672, 673, 59 Eumunida 269, 270, 22 Eumunidopsis 270 Eumunididae 266, 269, 271, 22 Eunephrops 178, 179 Eupasiphae 166, 167, 12 Euphausiacea 11, 12, 14, 15 Euphrosynoplax 741, 742, 744, 746 Euphylax 700, 708, 709 Eupilumnus 506, 507 Euplax 851, 852 Eupleurodon 581, 586, 594, 597 Eupontonia 122, 141, 144 Euprognatha 617–620 Eurycarcinus 669, 675, 677, 678 Eurynolambrus 622, 623, 626, 629, 53 Eurynome 625, 626, 629 Euryozius 723, 724 Eurypanopeus 737–740 Euryplacidae 427, 432, 510, 519, 44 Euryplax 520, 521 Eurypodius 578, 609, 611, 612, 614 Euryrhynchidae 2, 61, 63, 119 Eurytium 736, 737, 739, 64 Euryxanthops 778, 781–783, 67 Eusergestes 43, 44, 47 Eutrichocheles 189, 193, 195, 199 Eutrichopagurus 320, 322, 326, 330 Euxanthinae 751, 763 Euxanthus 765, 767, 769 Evibacus 252, 253, 255

Index to genus-group, family-group and higher taxonomic names

Exagorium 839–843 Exhippolysmata 88, 89 Exoclimenella 122, 138, 144 Exodromidia 445, 447–450 Exopalicus 648, 649 Exopheticus 524, 526, 527 Exopontonia 129, 136, 139, 141, 142, 144 Ezaxius 198 Fabia 865, 866, 868, 872 Facetotecta 12 Farfantepenaeus 27, 28 Fasciarma 813, 814, 818, 70 Favus 555, 560, 566 Fennera 127, 136, 142, 145 Fenneralpheus 66, 70, 71, 74 Fennerogalathea 274, 275 Fenneropenaeus 27 Filhollianassa 205, 207, 208, 212 Fizesereneia 795, 796, 799, 68 Flaberhina 469, 471, 19 Flindersoplax 722, 723 Foredromia 445, 448, 451 Forestiana 752–755 Forestocheles 343–345 Forestopagurus 321, 322, 326, 330 Formosaxius 190, 193, 194, 199 Fractosternalia 183 Fragillianassa 207, 209, 212, 16 Fresnerhynchus 111, 112 Frevillea 520, 521 Frodromia 443, 445, 453 Fultodromia 445, 451 Funchalia 27, 28, 30, 31, 33, 3 Fungicola 796, 798, 799, 68 Furtipodia 654, 657, 660, 57 Gaetice 828, 829 Gaeticinae 825, 828 Gaillardiellus 753–755, 64 Galacantha 284, 26 Galathea 273–275, 24 Galatheacaris 111 Galatheidae 263, 272, 273, 24 Galatheoidea 263–265, 272 Galearctus 254, 255, 257 Galene 665, 666, 59 Galenidae 427, 664, 665, 59 Galilia 557, 560, 564, 566, 46 Gandalfus 479, 480, 40 Gandoa 793, 794 Ganjampenaeopsis 30, 33

Gardinerosergia 45–47 Garthambrus 656, 657, 661, 57 Garthasia 773 Garthiella 751, 771, 772 Garthiellinae 751, 771 Garthinia 586, 590, 597 Garthiope 741, 742, 744, 746, 747 Garthopilumnus 668, 675–678 Gastroptychus 267, 268, 271, 22 Gaudichaudia 778, 780, 783 Gebiacantha 241–243, 18 Gebicula 241–243 Gebiidea 11, 12, 14, 185, 237 Gecarcinidae 2, 431, 802, 859 Gecarcinucidae 422, 425 Gecarcinucoidea 422, 425 Gecarcinus 802 Gecarcoidea 802 Gelasiminae 855, 857 Gelasimus 856, 857, 72 Gelastocaris 84–86 Gelastreutes 85, 86 Gemmotheres 864, 865, 869, 872 Genkaia 441, 442 Gennadas 24–26 Geograpsus 803, 804 Georgeoplax 530 Geosesarma 809, 812, 815, 816, 818, 70 Geryon 531, 691–693 Geryonidae 426, 687–689, 691, 697, 61 Gibbularctus 253, 254, 257 Gilvossius 207, 209–212 Giranauria 591, 592, 597 Glabropilumnus 667, 669, 675–678, 682, 60 Glassella 862, 865, 866, 870, 873 Glebocarcinus 489, 490, 42 Globopilumnus 507 Globospongicola 53, 54 Glypheidae 183 Glypheidea 9, 12, 14, 15, 183 Glyphocrangon 110, 8 Glyphocrangonidae 62, 104, 110, 8 Glyphus 166, 167, 12 Glyptocarcinus 757, 758 Glyptograpsidae 2, 422, 433, 802 Glyptograpsus 802 Glyptolithodes 312–314 Glyptoplax 737–739 Glyptoxanthinae 751, 772 Glyptoxanthus 772, 773 Glypturoides 220–223, 16 Glypturus 74, 219–223, 16

899

900

Marine Decapod Crustacea

Gnathophylleptum 121, 131 Gnathophyllidae 119, 120 Gnathophylloides 121, 131 Gnathophyllum 120, 121, 131, 10 Gomeza 493, 494, 42 Gonatonotus 668, 672, 673 Goneplacidae 422, 427–429, 510, 511, 519, 523, 667, 736, 741, 44 Goneplacinae 523 Goneplacoidea 422, 427, 428, 429, 510 Goneplacoides 524–527 Goneplax 524–527 Goniohellenus 703, 713, 714 Gonioinfradens 703, 713, 714 Gonioneptunus 700, 703, 712–714 Goniopsis 803, 804, 69 Goniopugettia 586, 593, 597 Goniosupradens 703, 713, 714, 62 Goniothorax 582, 586, 597 Gonopanope 741, 743–746, 777 Gopkittisak 825, 828, 829 Gordonella 39, 40 Gordonopsis 462, 38 Goreopagurus 318, 321, 323, 330, 33 Gorgonariana 669, 675, 677, 679 Gourretia 226, 227 Gourretiidae 225 Grapsidae 431, 802, 803, 824, 69 Grapsodius 802, 824 Grapsoidea 422, 430, 432, 792, 801, 837, 861 Grapsus 803, 804, 69 Griffinia 586, 593–595, 597 Grynaminna 221–223 Grypachaeus 640, 642–644, 55 Guaplax 542, 543, 546 Guinearma 809, 810, 813, 814, 818 Guinope 741, 743–746 Guinoplax 523–525, 527 Guinotellus 764, 765, 769, 66 Guinotinia 586, 592, 594, 597 Guinusia 807, 808, 69 Guitonia 778, 781, 783 Gustavus 793, 794 Guyanacaris 191, 193, 199 Haberma 811, 812, 815, 818 Hachijopagurus 332, 326, 330 Hadropenaeus 39, 40 Hadroplax 524, 525, 527 Haemocinus 722, 723 Haigiopagurus 323, 325, 330 Haledromia 444, 445, 451 Halicarcinides 542, 544–546

Halicarcinus 542, 545, 547, 45 Halimede 665, 666, 59 Halimena 542, 544, 545, 547 Haliporoides 39, 40 Haliporus 39, 40, 4 Halocaridina 95 Hamiger 129, 136, 141, 145 Hamodactyloides 123, 140, 142, 145 Hamodactylus 123, 136, 142, 145 Hamopontonia 129, 136, 139, 145 Hapalocarcinus 796, 798, 799, 68 Hapalogaster 312, 314, 315 Hapalogastridae 311 Hapalogastrinae 312, 315 Hapalonotinae 666, 668, 672, 674 Hapalonotus 666, 668, 672, 674 Hapaloptyx 267, 268 Harperalpheus 69, 70, 74 Harpiliopsis 125, 136, 138, 145 Harpilius 125, 142, 145 Harrovia 668, 673, 59 Harryplax 720, 721 Hayashidonus 168, 169 Heardaxius 216, 217 Heikea 498 Heikeopsis 496–498, 53 Helicana 826, 827 Helice 826–828 Heloeciidae 433, 868, 849, 71 Heloecius 849, 71 Helograpsus 826, 827, 71 Hemigrapsus 831–833, 71 Hemipagurus 328 Hemipenaeus 21–23, 2 Hemiplax 851, 852 Hemisphaerodromia 444, 445, 447, 448, 451 Hemus 634–637 Hendersonida 278, 279, 281 Henicoplax 520, 521 Hepatella 474–476, 40 Hepatoporus 764, 765, 770 Hepatus 474–476, 40 Hephthopelta 515, 516, 518 Hepomadus 21–23 Heptacarpus 92, 93 Herbstia 585, 586, 591, 597, 49 Heteractaea 752, 753, 755, 64 Heteroaxianassa 238, 239 Heterocarpus 161–163, 12 Heterocrypta 654, 657, 661, 57 Heterogenys 116, 117, 9 Heterolithadia 556, 560, 566, 46

Index to genus-group, family-group and higher taxonomic names

Heteronida 278, 279, 281 Heteronucia 552, 555, 560, 563, 567, 570, 46 Heteropanope 669, 676, 677, 679 Heteropenaeus 27, 28, 31, 33 Heteropilumnus 667, 670, 671, 675–677, 679, 682, 683, 60 Heteroplax 520, 521 Heteropolyonyx 287, 288, 290, 291 Heteroporcellana 288, 289, 291 Heteroptychus 267, 268 Heterotremata 421, 423, 430, 431, 473 Heteroziidae 422, 428, 432, 477, 478 Heterozius 477, 478 Hexagonalia 728–730 Hexagonaloides 728–730 Hexalaughlia 537–539 Hexanauplia 9, 10, 12 Hexapanopeus 737–739 Hexapinus 538–539 Hexaplax 537, 539, 44 Hexapodidae 423, 536, 44 Hexapodoidea 423, 536 Hexapus 537–539 Hiplyra 559, 560, 563, 564, 567, 46 Hippa 299, 300, 30 Hippidae 296, 299, 30 Hippoidea 263, 264, 295 Hippolyte 84–86, 7 Hippolytidae 61, 64, 83, 7 Hiroia 796, 799, 800 Hirsutodynomene 455, 456, 36 Hispidolambrus 655, 658, 660, 661 Holoplites 586, 593, 598 Holotheres 864, 866, 868, 873 Holothuriophilus 865, 866, 868, 873 Holthuija 624–627, 629, 53 Holthuisaeus 128, 136, 138, 139, 145 Holthuispenaeopsis 30, 33 Homalaspis 508, 509 Homalodromia 446, 448, 451 Homarinus 178, 179 Homarus 177–179 Homeryon 259, 260 Homoioplax 523, 736 Homola 462, 463, 38 Homolax 461–463 Homolidae 423, 459, 460, 465, 38 Homolochunia 460, 462, 463 Homolodromia 458, 38 Homolodromiidae 423, 442, 457, 38 Homolodromioidea 423, 442, 457 Homologenus 460, 462, 463, 38 Homoloida 421, 423, 459, 473

Homoloidea 423, 431, 459 Homolomannia 460, 462, 463, 38 Hoplocarida 10, 13, 14 Hoplocypode 858 Hoplophrys 582, 590, 594, 598, 49 Hospitotheres 870, 873, 49 Huenia 581, 582, 594, 598 Hyas 641–644, 55 Hyastenus 586, 591, 592, 598, 49 Hymenicoides 542, 544, 545, 547 Hymenocera 120, 131, 10 Hymenoceridae 119, 120 Hymenodora 115–117 Hymenopenaeus 39, 40 Hymenosoma 541–543, 545, 547 Hymenosomatidae 422, 423, 430–432, 473, 541, 617, 45 Hymenosomatinae 541, 545 Hymenosomatoidea 422, 430, 473, 540 Hypocolpus 764, 765, 767, 770, 66 Hypoconcha 443, 446, 447, 453 Hypoconchinae 443, 453 Hypolambrus 654, 658, 661 Hypothalassia 500, 503 Hypothalassiidae 428, 501, 503 Hypsophrys 463 Ibacinae 252, 253, 255 Ibacus 253–255, 21 Icelopagurus 322, 326, 330 Ichthyostraca 9, 10, 12 Ihleus 554, 560, 563, 565, 567 Ihlopsis 460, 462, 463, 38 Ilia 556, 560, 567 Iliacantha 556, 561, 563, 567, 47 Ilyograpsinae 849 Ilyograpsus 850–852, 72 Ilyogynnis 839, 842, 843 Ilyoplax 846–848, 71 Inachidae 425, 465, 578, 579, 608, 617, 639, 52 Inachinae 577 Inachoides 617, 619, 620 Inachoididae 425, 578, 579, 616, 52 Inachoidinae 577, 620 Inachus 609, 611, 615 Incultus 701, 710, 711 Indopinnixa 862, 866, 869, 873 Ingolfiellida 13, 14 Insulaplax 542, 543, 545, 547 Intesius 531, 532 Iphiculidae 424, 551, 45 Iphiculus 551, 45 Iridopagurus 320, 323, 330

901

902

Marine Decapod Crustacea

Ischnopontonia 128, 136, 139, 145 Isocheles 305, 306, 308 Isopoda 11, 13, 14 Isopontonia 130, 139, 141, 142, 145 Itampolus 667, 669, 675, 677, 679, 682 Ixa 552, 557, 561, 563, 567, 47 Izanami 486, 487 Izucaris 124, 136, 145 Jacforus 764, 765, 767, 768, 770, 777 Jacquinotia 623, 626, 628, 629 Janetogalathea 274, 275 Janicea 81 Janicella 118 Jasus 248–250, 1, 19 Jaxea 238, 239 Jengalpheops 67, 70, 72, 74 Jocaste 124, 136, 142, 145, 10 Jocullianassa 205, 207–209, 212 Jonas 493, 494, 42 Jonesius 327 Justitia 248–250, 19 Juxtafabia 867, 868, 870, 873 Juxtastenopus 56, 5 Juxtaxanthias 778, 781, 784 Kabutos 551, 561, 563, 568 Karstarma 810, 816, 818 Karumballichirus 220–223 Kasagia 624, 625, 629 Kaviengella 127, 136, 146 Keldyshicaris 97, 98 Kemphyra 115–117 Kemponia 144 Ketamia 436–438, 16 Kiictenocheloides 226, 227 Kimbla 625, 627, 629 Kingelus 526, 527 Kirnasia 100, 101 Kishinouyepenaeopsis 30, 33 Kiwa 270, 271, 23 Kiwaidae 265, 270, 23 Krangalangia 436–438, 36 Kraussia 772–774 Kraussiinae 750, 772 Kraussillichirus 220, 223, 540 Kroppcarcinus 796–798, 800 Kume 688, 701, 705, 706 Kumepagurus 318, 319, 323, 324, 330 Labidochirus 322, 325, 330, 340 Labuanium 810, 814, 815, 818, 70

Lachnopodus 778, 782, 784, 67 Ladomedaeus 766–768, 770, 66 Laevicaudata 12 Lahaina 587, 591, 592, 594, 598 Laleonectes 701, 705, 706, 61 Lamarckdromia 444, 446, 448, 451 Lamarckopilumnus 670, 675, 677, 679 Lambdophallus 537–539 Lambrachaeus 653, 654, 658, 661, 57 Lamoha 461–463, 38 Laomedia 238, 239, 18 Laomediidae 237, 18 Laomenes 124, 136, 138, 146, 10 Latatheres 863, 865, 867, 873 Laticallichirus 221–223 Latohexapus 537–539 Latopilumnus 671, 675–677, 679 Latreillia 459, 465, 38 Latreilliidae 423, 459, 465, 38 Latreillopsis 460–463, 38 Latreutes 83–86, 7 Latulambrus 653, 658, 661 Laubierinia 587, 590, 598 Laurentaeglyphea 183, 184 Laurentgourretia 226, 227 Lauridromia 444, 446, 447, 451, 37 Lauriea 274, 276, 24 Lazarocleistostoma 846–848 Leander 121, 133, 10 Leandrites 121, 132–134 Lebbeus 92, 93 Leelumnus 670, 675, 677, 679 Leiogalathea 283, 284, 26 Leiolambrus 653, 658, 661 Lemaitreopsis 346 Lentilumnus 669, 675–677, 679, 681 Lenzicarcinus 112, 113 Leonardsaxius 190, 193, 200 Leontocaris 83, 84, 86, 7 Lepidonaxia 587, 590, 598 Lepidopa 296–298, 29 Lepidophthalmus 219–222, 224, 16 Lepidopinae 296 Leptalpheus 66, 70–72, 74, 6 Leptathanas 67, 71, 72, 74 Lepteces 583, 590, 598 Leptocarpus 120, 122, 132, 134 Leptochela 166, 167, 12 Leptochryseus 840–843 Leptodius 777, 779, 782, 784, 786, 67 Leptograpsodes 805, 806, 69 Leptograpsodidae 431, 432, 802, 805, 806, 69

Index to genus-group, family-group and higher taxonomic names

Leptograpsus 803, 804 Leptomaia 587, 590, 599 Leptomenaeus 126, 136, 138, 141, 146 Leptomithrax 623, 627, 629, 1, 53, 54 Leptopagurus 320, 323, 324, 331 Leptopalaemon 122, 132–134 Leptopisa 583, 592, 599, 50 Leptostraca 10, 11, 13 Leptuca 855–858 Leslibetaeus 68, 70, 71, 75, 6 Leucippa 581, 583, 599 Leucisca 522, 544, 561, 563, 568 Leucolepidopa 296–298 Leucosia 552, 553, 574, 48 Leucosiidae 424, 431, 474, 551, 46–48 Leucosiinae 552, 553, 573 Leucosilia 556, 561, 564, 568 Leucosioidea 424, 550 Leurocyclus 618–620 Levantocaris 190, 193, 194, 200 Levicaris 121, 131 Lewindromia 446, 448, 451, 1 Liagore 779, 782, 784 Libidoclaea 583, 593, 599 Libinia 583, 591, 594, 599, 50 Libystes 426, 688, 700, 701, 705, 706 Lichtylus 793, 794 Ligur 89 Lillyanella 840–843 Limnopilos 542, 544, 545, 547 Limotheres 865, 873 Linnaeoxanthidae 430, 734, 735, 741 Linnaeoxantho 735 Linuparus 248–250, 19 Liocarcinus 697–699, 61 Liocarpilodes 760–762, 65 Liomera 774, 775, 66 Liomerinae 751, 774 Liopetrolisthes 286–288, 291, 27 Lioxanthodes 779, 782, 784 Lipaesthesius 765, 767, 770 Lipkebe 123, 138, 139, 146 Lipkecallianassa 206, 209, 210, 212 Lipkemedaeus 767, 768, 770 Lipkemenes 124, 136, 146 Lipkemera 774, 775 Lipkiidae 63, 111, 9 Lipkius 111, 112, 9 Lissa 581, 587, 599, 600 Lissocarcinus 700, 702, 712–714, 62 Lissocrangon 105–107 Lissomorpha 557, 561, 563, 568

Lissoporcellana 288, 290, 291, 27 Lissosabinea 104, 107, 8 Lithadia 554, 561, 568, 47 Lithodes 311, 313–315, 31 Lithodidae 263, 264, 302, 311, 31 Lithodinae 312–315 Lithodoidea 263, 311 Lithopagurus 319, 323, 331 Lithoscaptus 796, 798, 800, 68 Lithoselatium 810, 812, 819 Litocheira 530 Litocheiridae 427, 511, 530 Litopenaeus 27, 35 Litoraxius 192, 193, 195, 200 Litosus 611, 612, 615 Liusius 553, 574, 575 Lobiactaea 753–755 Lobopilumnus 670, 675, 680 Lomis 263, 264, 300, 301, 30 Lomisidae 263, 300, 30 Lomisoidea 263, 264, 300 Lonchodactylus 441, 442 Lophaxiopsis 188, 193, 200 Lophaxius 188, 193, 200 Lophogastrida 13–15 Lopholithodes 315 Lophomastix 299 Lophopagurus 323, 325, 331 Lophopanopeus 736–739 Lophopilumnus 669, 675, 677, 680 Lophoplax 667, 670, 675, 680, 682 Lophorochinia 582, 583, 599 Lophoxanthus 742, 743, 746 Lophozozymus 787, 788, 67 Loxopagurus 305–308, 31 Loxorhynchus 587, 593, 599 Lucascinus 543–545, 547 Lucaya 101 Lucensosergia 45–47 Luciades 796–798, 800 Lucifer 41, 42 Luciferidae 19, 20, 41 Luniella 760–762 Lupella 702–704, 61 Lupocyclinae 702, 707, 710 Lupocycloporus 702, 707 Lupocyclus 701, 702, 707, 708 Luteocarcinus 671, 682, 683 Lutogemma 430, 851, 853 Lybia 776, 66 Lydia 506, 507 Lyphira 559, 564, 568, 47

903

904

Marine Decapod Crustacea

Lyramaia 587, 590, 599 Lyreididae 423, 467, 468, 39 Lyreidinae 468 Lyreidus 468, 39 Lysirude 438, 39 Lysmata 88–90, 7 Lysmatella 89 Lysmatidae 64, 88, 7 Machaerus 520–522 Macphersonus 112, 113 Macrobrachium 61, 119, 120, 122, 132, 134 Macrocheira 578, 639–641, 643, 644, 719, 55 Macrocheirinae 639 Macrocoeloma 398, 583, 592, 600 Macromaxillocarididae 51, 52, 54 Macromaxillocaris 52 Macromedaeus 777–779, 784, 67 Macropetasma 29–31, 34, 3 Macrophthalmidae 430, 433, 792, 838, 849, 72 Macrophthalminae 849 Macrophthalmus 849, 851–853, 72 Macropipidae 697 Macropipus 697–699 Macropodia 609, 611, 612, 615 Macroregonia 640, 641, 643, 644 Macrothea 274–276, 24 Madangella 125, 136, 141, 146 Madarateuchis 286, 291 Magnotheres 864, 869, 870, 873 Maguimithrax 635–637 Maiopsis 622, 623, 626, 629 Maja 622, 624, 626, 627, 630 Majella 623, 624, 630 Majidae 425, 577–580, 621, 53, 54 Majinae 577, 622 Majoidea 425, 541, 577, 617 Malacoplax 736, 741–744, 747 Malacostraca 2, 9, 10, 12 Maldivia 726, 727 Manarma 815, 816, 819, 70 Manipontonia 126, 136, 146 Manningis 840, 842, 843 Mantisgebia 241–243 Manucomplanus 323, 325, 331 Manuscaris 97, 98 Marcusiaxius 232 Margitonia 126, 136, 142, 146 Marianaxius 189, 193, 200 Mariaplax 537–539, 44 Marratha 777, 779, 782, 784 Marsupenaeus 27

Maryprocessa 169 Mastigochirus 299, 300, 30 Mathildella 531, 532, 44 Mathildellidae 429, 511, 531, 44 Matuta 486, 487, 41 Matutidae 424, 481, 482, 466, 41 Maximiliaeus 39, 40, 4 Mclaydromia 444, 446–448, 451 Mebeli 486, 487 Medaeops 766, 767, 769, 770, 66 Medaeus 766, 768–770, 66 Medorippe 496–498 Megaesthesiinae 514, 515, 518 Megaesthesius 515, 516, 518 Megalobrachium 288, 289, 292, 27 Megametope 777, 779, 784 Megokris 27, 30, 31, 34, 36 Meinertaxius 199 Melicertus 27, 28 Melybia 735 Menaethiops 587, 590, 591, 600 Menaethius 587, 589, 600, 50 Meningodora 116, 117 Menippe 500, 504, 43 Menippidae 428, 500, 501, 504, 43 Menoplax 524–526, 528 Meractaea 752–754, 756 Merguia 90, 91, 7 Merguiidae 63, 90, 7 Merhippolyte 83, 85, 87 Merocryptoides 555, 561, 568 Merocryptus 554, 561, 564, 569, 47 Mertonia 671, 682, 683 Mesocrangon 105, 107 Mesopenaeus 38–40 Mesopontonia 123, 136, 138, 141, 146 Mesorhoea 653, 658, 660, 661 Mesotheres 864, 866, 867, 874 Metabetaeus 69, 70, 72, 75 Metacarcinus 432, 489, 490, 42 Metacrangon 105, 107, 8 Metadromia 444, 446, 447, 452, 37 Metadynomene 455–457 Metadynomeninae 155, 457 Metagrapsus 809, 811, 813, 814, 819 Metalpheus 68, 72, 75 Metanephrops 178, 179, 13 Metapenaeopsis 27, 29–31, 34, 3 Metapenaeus 27, 29–31, 34 Metaplax 826, 827, 71 Metapontonia 130, 136, 146 Metasesarma 810, 815, 819, 70

Index to genus-group, family-group and higher taxonomic names

Metaxanthops 777, 779, 784 Meticonaxius 232 Metopaulias 809 Metopocarcinus 736, 737, 739 Metopograpsus 803, 804 Metoporhaphis 610, 612, 615 Micas 541, 542, 544, 545, 547 Michaelcallianassa 221, 222, 224 Michaelimenes 127, 136, 141, 146 Michelaxiopsis 193, 196, 201 Michelea 231, 232 Micheleidae 186, 231, 17 Michelopagurus 320, 326, 331 Micippa 578, 583, 587, 600, 50 Micippoides 587, 592, 600 Microcassiope 779, 781, 785 Microgoneplax 427, 511, 524, 525, 528 Microhalimus 625, 627, 630 Microlissa 581, 583, 600, 50 Micropagurus 321, 322, 324, 326, 331 Micropanope 743–745, 747, 748, 782–785 Microphrys 634–637 Micropisa 587, 592, 600 Microprosthema 52–54, 5 Microtopsis 515, 516, 518 Mictacea 13, 14 Mictaxius 216, 217 Mictyridae 432, 854, 72 Mictyris 454, 1, 72 Miersarma 811, 813, 819 Miersiella 766–768, 770 Miersiograpsus 807, 808 Mierspenaeopsis 29, 30, 34 Migmarma 815, 818, 819 Milnepanopeus 736–739 Mimilambrus 653, 658, 660, 661 Mimocaris 88–90 Mindanium 811, 816, 819 Minuca 855–858, 72 Minyocerus 287, 288, 292 Minyorhyncha 587, 593, 600, 50 Miopontonia 120, 123, 136, 141, 142, 147 Mirocaris 97, 98 Mirograpsus 850, 851, 853 Miropalicus 649 Miropandalus 159 Mithracidae 425, 478–480, 622, 634, 54 Mithracinae 577, 578, 580 Mithraculus 635–637, 54 Mithrax 636–638, 54 Mixtopagurus 343–345 Mocallichirus 220, 222, 224

Mocosoa 587, 600 Moguai 839–842, 844 Mohocaris 68, 70, 72, 75 Moloha 461, 462, 464, 38 Monodaeus 766–768, 770 Monomia 702, 709–712 Montanaxius 188, 190, 193, 201, 15 Moreiradromia 446–448, 452 Mortensenella 840, 842, 844, 71 Mucrollichirus 220, 221, 224, 16 Multicrustacea 10, 12 Munida 277–279, 281, 25 Munididae 272, 277, 25 Munidopagurus 319, 326, 331 Munidopsidae 272, 283, 26 Munidopsis 283, 284, 26 Muradium 814, 819 Mursia 482–485, 41 Myomenippe 504, 505, 43 Myopilumnus 680 Myra 556, 561, 563, 569, 47 Myrine 556, 569 Myropsis 552, 556, 561, 564, 569, 47 Mysida 11, 13–15 Mystacocarida 9, 10, 12 Namlacium 811, 816, 820 Nancyplax 520, 522 Nannotheres 863, 865, 874 Nanocassiope 779, 781, 785, 67 Nanogalathea 273, 274, 276 Nanopilumnus 671, 676, 680 Nanoplax 743–745, 747 Nanosesarma 810, 812, 815, 820, 70 Nanusia 840, 842, 844 Nasima 840–844, 71 Nasutocarcinus 601 Nasutoplax 543, 545, 548 Natantia 61 Naushonia 238, 239, 18 Nauticaris 83, 84, 86, 7 Nautilocaris 97, 98 Nautilocorystes 493, 688, 694 Nautilocorystidae 422, 425, 426, 493, 688, 494 Nautilocorystinae 422, 688, 694 Naxia 623, 626, 627, 630, 645, 53 Naxioides 585, 587, 601, 50 Neaxiopsis 234 Neaxius 234, 17 Necallianassa 206, 208–210, 212 Necora 698, 699, 61 Necronectes 708

905

906

Marine Decapod Crustacea

Necronectinae 702, 705, 708 Nectipoda 12 Nectocarcinus 688, 691–693, 695, 697, 61 Nectopanope 520–522, 777 Neikolambrus 656, 658, 662, 57 Nematocarcinidae 63, 112, 9 Nematocarcinoidea 61, 62, 110 Nematocarcinus 111–113, 9 Nematopaguroides 321, 324, 331 Nematopagurus 322, 331, 33 Nematopalaemon 121, 132, 134 Nemausa 635–638, 54 Nennalpheus 66, 68, 70, 71, 75 Neoactumnus 669, 676, 680 Neoalpheopsis 75 Neoanchistus 130, 136, 139, 147 Neocallichirus 219–222, 224, 16 Neoclimenes 125, 136, 147 Neocorycodus 435–437 Neocrangon 105–107 Neodoclea 585, 587, 601 Neodorippe 496–498 Neoeriocheir 831–833 Neogebicula 241–243 Neoglyphea 183, 184 Neogoneplax 524–526, 528 Neohelice 826, 827 Neohymenicus 543–545, 548 Neoliomera 774, 775, 66 Neolioxantho 780–782, 785 Neolithodes 313, 314, 316, 31 Neomaja 624, 626, 630 Neommatocarcinus 429, 524, 525, 528 Neonida 278, 279, 281 Neopalaemon 120, 122, 133, 134 Neopalicus 648, 649, 56 Neopanope 737–739 Neopericlimenes 124, 136, 147 Neopetrolisthes 287, 288, 290, 292, 27 Neophrys 587, 594, 601, 50 Neopilumnoplax 531, 532 Neopisosoma 287, 288, 290, 292, 27 Neopontonides 124, 136, 147 Neorhynchoplax 541–545, 548 Neosarmatium 814, 815, 820, 70 Neosergestes 44, 45, 47 Neosesarma 812, 814, 820 Neostylodactylus 171, 172 Neotroglocarcinus 796–798, 800 Neotrypaea 206–208, 210–212, 16 Neoxanthias 779, 781, 785, 67 Neoxanthops 777, 779, 783, 785, 786, 67

Neoxenophthalmus 860, 861 Neparilia 557, 561, 564, 569, 47 Nephropidae 175, 177, 13 Nephropides 178, 179 Nephropoidea 175 Nephrops 178–180 Nephropsis 178–180, 13 Nepinnotheres 865, 867, 870, 874 Nibilia 584, 585, 601 Nicoya 587, 589, 591, 601 Nigmatullinus 112, 113 Nikoides 169, 170 Niobafia 652, 655, 656 Nipponcalaxiopsis 198 Nippontonia 127, 136, 139, 147 Noarograpsus 831–833 Nobiliella 552, 554, 561, 563, 569 Nobilum 497, 498 Nodolambrus 655, 658, 660, 662 Nonala 635, 636, 638 Notalpheus 67, 71, 75 Notiax 205, 208, 210, 213 Notocrangon 105–107 Notogennema 24 Notolopas 584, 585, 591, 601 Notomithrax 625, 627, 630, 53 Notonyx 523, 524, 528 Notopandalus 161, 163 Notopelta 515, 516, 518 Notopodinae 469–471 Notopoides 469, 471, 472 Notopontonia 130, 141, 147 Notopus 469–472, 39 Notosceles 469–472 Notostomus 116, 117, 9 Notostraca 11, 12 Novactaea 752–754, 756 Novorostrum 287, 288, 290, 292, 27 Nucia 552, 555, 561, 569, 47 Nuciops 554, 561, 570 Nupalirus 248–250 Nursia 552, 554, 561, 563, 564, 570, 652 Nursilia 552, 561, 563, 570, 47 Ochtholambrus 655, 658, 660, 662 Ocypode 855, 856, 858, 72 Ocypodidae 431, 433, 838, 845, 855, 72 Ocypodinae 856, 858 Ocypodoidea 432, 792, 801, 837, 860, 861 Odhnea 752, 754, 756, 64 Odhneria 756 Odiomarinae 541, 549

Index to genus-group, family-group and higher taxonomic names

Odiomaris 541, 543–545, 549 Odontonia 129, 136, 138, 147 Odontoplax 736, 737, 740 Odontozona 51, 56 Oedignathus 312, 314, 315 Oediplax 743, 745, 474 Ogyrides 91, 7 Ogyrididae 63, 65, 91, 7 Olenorfia 653, 655, 656, 57 Olenothus 766, 767, 770 Oligostraca 9, 10, 12 Omalacantha 634–636, 638 Ommatocarcinus 431, 524, 525, 528 Onchycaudata 11, 12 Oncinopus 609, 611, 612, 615, 52 Onconida 278, 279, 281 Oncopagurus 340, 341, 34 Onychomorpha 553, 558, 559 Onycocaridella 129, 136, 140, 141, 147, 148 Onycocaridites 128, 136, 141, 142, 147 Onycocaris 129, 139, 141, 142, 147 Onycomenes 123, 138, 148 Opaepele 97, 98 Opaepupu 130, 136, 148 Opecarcinus 796, 797, 800, 68 Opisthopus 867, 868, 874 Oplophoridae 61, 62, 115, 117, 9 Oplophoroidea 62, 114 Oplophorus 118, 9 Oplopisa 587, 593, 594, 601 Opusia 840, 842, 844 Orcovita 830, 832–834 Oregonia 641–644 Oregoniidae 425, 578, 579, 608, 639, 55 Oregoniinae 577, 639 Oreophorus 555, 561, 563, 564, 570 Oreotlos 555, 561, 563, 564, 570 Orientotlos 555, 561, 570 Orisarma 809, 813, 815, 816, 820 Orithyia 646, 56 Orithyiidae 424, 646, 56 Orithyioidea 424, 646 Orphnoxanthus 779, 785 Orthochela 286, 289, 292 Orthopagurus 326, 332 Orthopontonia 129, 136, 138, 148 Orthotheres 865, 867–869, 874 Orygmalpheus 68, 70, 71, 75 Osachila 474–476, 40 Ostracoda 9–12 Ostraconotus 318, 322, 326, 332 Ostracotheres 864, 865, 867, 870, 874

Otmaroxanthus 427, 508, 509 Otognathon 830, 832, 833 Ovalipes 689, 691, 695, 697, 61 Ovalipidae 426, 432, 687–689, 691, 695, 697, 61 Ovatis 780, 782, 785 Ovilyra 558, 561, 564, 570 Ovimaja 623, 624, 627, 630, 53 Oxypleurodon 587, 588, 601, 50 Oxyrhyncha 421 Oxyrhynchaxius 190, 193, 195, 201 Oxystomata 421 Oziidae 428, 500, 501, 504, 505, 43 Ozius 504, 506, 507, 43 Pachelpheus 66, 72, 75 Pachycheles 286, 287, 289, 290, 292, 27 Pachygrapsus 803, 804, 69 Paeduma 538, 539 Pagurellus 320, 324, 332 Paguridae 263, 301, 302, 311, 317, 340, 345, 346, 33, 34 Paguridium 324, 325, 332 Pagurina 324, 325, 332 Paguriscus 318, 319, 324, 332 Paguristes 306–308, 33 Paguritta 323, 326, 332, 33 Pagurixus 324, 325, 332 Pagurodes 320, 332, 326, 332, 33 Pagurodofleinia 321, 324, 326, 333 Paguroidea 3, 263, 264, 301, 312 Pagurojacquesia 320, 323, 324, 333 Paguropsina 302, 306, 307, 309 Paguropsis 302, 306, 307, 309 Paguruncio 318, 319, 324, 333 Pagurus 263, 293, 318, 323, 326, 333, 34 Palaemon 119–121, 132–134, 10 Palaemonella 122, 136, 148 Palaemonidae 61, 63, 119, 10, 11 Palaemoninae 119, 121 Palaemonoidea 61, 63, 119 Palapedia 772–774, 66 Palatigum 766, 767, 771 Palibythus 248–250 Palicidae 424, 647, 648, 56 Palicoidea 424, 514, 647 Palicoides 649, 650 Paliculus 649, 650, 56 Palicus 648, 650 Palinura 14 Palinurellus 248–250 Palinuridae 247–248, 19 Palinuroidea 247 Palinurus 247–250

907

908

Marine Decapod Crustacea

Palinustus 248–250 Palmyria 726, 727 Pancrustacea 9 Pandalidae 61, 63, 158, 160, 12 Pandalina 161, 163 Pandaloidea 63, 158 Pandalus 160, 161, 164, 12 Panopeidae 430, 523, 734, 735, 741, 64 Panopeus 736, 737, 738, 740, 64 Panoplax 743–745, 747, 64 Pantomus 160, 161, 164 Panulirus 247–250, 19 Parabetaeus 66, 70–72, 75 Paracalliacidae 186, 233 Paracallianidea 216, 217 Paracalliax 233 Paracleistostoma 829, 840, 842, 844, 848 Paraclimenaeus 128, 136, 139, 141, 148 Paraclimenes 125, 136, 148 Paracrangon 104, 107 Paractaea 754, 756, 64 Paractaeopsis 754, 756 Paracyclois 483–485, 41 Paradasygyiinae 620 Paradasygyius 617, 619, 620 Paradorippe 496–498 Paradromia 445, 446, 452 Paradynomene 455–457, 36 Paradynomeninae 455, 457 Paraentomonyx 625, 628–630 Paragalene 429, 432, 533, 534 Paragebicula 234 Paragiopagurus 340, 341 Paraglypturus 228–231 Paragoneplax 427, 511, 524–526, 528 Paragourretia 187, 226, 227, 17 Paragrapsus 826, 827, 71 Parahelice 826–828 Parahepomadus 21–23, 2 Parahexapus 538, 539 Paralatreutes 83 Paralbunea 297, 29 Paralebbeus 92, 93 Paraleptuca 856–858, 72 Paraleucolepidopa 296–298 Paraliomera 774, 775 Paralithodes 311, 313, 314, 316, 31 Paralomis 311, 313, 314, 316, 31 Paramaja 624, 626, 627, 631, 53 Paramaya 624, 626, 627, 631, 53 Paramedaeus 766, 768, 771 Paramicippa 628

Paramithrax 626, 628, 631 Paramoguai 839, 841, 842, 844 Paromolopsis 461, 462, 464, 38 Paramunida 278, 279, 281 Paranaxia 587, 589, 601 Paranchistus 125, 136, 141, 142, 148 Paranotonyx 671, 682, 683 Paranursia 552, 553, 558, 559, 46 Parapaguridae 301, 302, 339 Parapagurodes 321, 326, 333 Parapagurus 321, 326, 333 Parapalicus 649, 650 Parapanope 665, 666 Parapasiphae 166, 167 Parapenaeidae 27 Parapenaeinae 27 Parapenaeopsis 27, 30, 31, 33, 34, 3 Parapenaeus 27, 29, 32, 35 Parapetrolisthes 287, 289, 292 Parapilumnus 512, 513, 43 Parapinnixa 861–863, 865, 874 Parapleisticantha 578, 639–644 Parapleurophrycoides 676 Parapontocaris 105–107, 8 Parapontophilus 105–107 Parapylocheles 343–345 Parapyxidognathus 831–834 Parascytoleptus 189, 191, 201, 15 Paraselwynia 671, 682, 683 Parasergestes 44, 45, 47 Parasesarma 812–814, 820, 70 Paraspongicola 52–54 Parastacidae 2, 175 Parastacoidea 175 Parastylodactylus 171, 172 Paratergatis 787, 788, 67 Parathranites 687, 688, 697–699, 61 Paratrichocheles 199 Paratrypaea 208, 210, 213, 16 Paratya 95 Paratylodiplax 840–842, 844 Paratymolus 579, 608, 610–612, 615, 52 Paratypton 123, 136, 138, 148 Paraxanthias 780, 781, 783, 785, 67 Paraxanthodes 766, 767, 769, 771, 777 Paraxanthus 777, 780, 785 Paraxiopsis 189, 193, 201, 15 Paraxius 190, 193, 201, 15 Parethusa 498–500, 43 Parhippolyte 80, 81 Parilia 557, 562–564, 571, 47 Pariphiculus 551, 45

Index to genus-group, family-group and higher taxonomic names

Paromola 459, 461, 462, 464, 38 Parribacus 253, 255, 21 Parthenolambrus 663 Parthenope 654, 658, 660–662, 57 Parthenopidae 429, 474, 651, 652, 57, 58 Parthenopinae 652, 654, 656 Parthenopoidea 429, 500, 650 Parthenopoides 656, 658, 662 Pasiphaea 166, 167, 12 Pasiphaeidae 61, 62, 165, 12 Pasiphaeoidea 62, 165 Patagurus 318, 322, 333, 33 Patonia 125, 136, 148 Patulambrus 656, 659, 662 Paulita 617, 618, 620 Paulitinae 620 Pedroplax 524–526, 528 Peisos 43, 44, 47 Pelagopenaeus 27, 29, 32, 33, 35 Pele 701, 705, 706 Peleianus 667, 669, 677, 680, 682 Pelia 587, 589, 591, 602, 50 Peloeus 508, 509 Peltarion 732, 733 Pelycomaia 797, 798, 800 Penaeidae 19, 20, 21, 27, 3 Penaeinae 27 Penaeoidea 19, 21 Penaeopsis 27, 29, 32, 35, 3 Penaeus 19, 20, 27, 28, 32, 35, 3 Pentacheles 259, 260 Pentastomida 10, 12 Peracarida 13, 14 Percnidae 433, 802, 806, 69 Percnon 806, 807, 69 Periclimenaeus 128, 138, 139, 148, 10 Periclimenella 125, 140, 141, 148 Periclimenes 119, 127, 136, 139, 146, 148, 150, 11 Periclimenoides 130, 138, 149 Perinia 588, 589, 594, 602 Peripandalus 161, 164 Perisesarma 813, 814, 820 Permanotus 667, 673, 59 Persephona 552, 556, 562, 564, 571, 47 Perunorhombila 741, 742, 747 Pestoxanthodes 780, 785 Petalidium 43, 44, 47 Petalomera 445–447, 452, 37 Petramithrax 634, 636, 638 Petrarctus 254, 257 Petrocheles 287, 289, 290, 293 Petrochirus 305, 309, 33

Petrolisthes 286, 287, 289, 290, 293, 28 Petruca 855–858 Phalangipus 584, 588, 602 Philarius 129, 136, 142, 149 Philippicarcinus 728–730 Philippidorippe 496–498 Philocheras 104, 106, 108 Philyra 552, 558, 562, 564, 571, 50 Philyrinae 552 Phimochirus 325, 326, 333, 34 Phlyxia 552, 557, 562, 571, 47 Phorcosergia 45–47 Phycocaris 84, 85, 87 Phycomenes 126, 136, 138, 142, 149 Phylladiorhynchus 273, 274, 276, 24 Phyllocarida 13 Phyllodorippe 496–498 Phyllognathia 120, 131, 132 Phyllolithodes 313, 314, 316 Phyllopoda 12 Phyllotymolinidae 423, 434, 435, 441 Phyllotymolinum 441, 442 Phymodius 762 Physachaeus 610, 611, 615 Physetocarididae 61, 63, 168 Physetocaridoidea 63, 168 Picroceroides 584, 589, 602 Picrocerus 588, 589, 602, 50 Pilbaraxius 190, 191, 194, 201 Pillsburyaxius 191, 193, 195, 202 Pilodius 760–762, 65 Pilomedaeus 766, 767, 771 Piloslambrus 653, 659, 662 Pilumnidae 427, 432, 664–666, 59, 60 Pilumninae 667, 668, 674 Pilumnoidea 426, 664, 733 Pilumnoides 456, 721 Pilumnoididae 427, 429, 720 Pilumnopeus 670, 675, 677, 680, 60 Pilumnus 670, 676, 678, 680, 60 Pinnaxodes 864, 867, 868, 874 Pinnixa 862, 863, 865, 869, 871, 874, 876 Pinnixulala 862, 867, 869, 870, 875 Pinnotherelinae 802, 824, 825, 829, 861 Pinnotheres 862, 864, 867, 870, 875 Pinnotheridae 422, 430–432, 792, 853, 860–862 Pinnotheroidea 430, 792, 825, 830, 861 Pinnotherotonia 120, 127, 149 Pippacirama 622, 625, 626, 631 Pirimela 696, 61 Pirimelidae 426, 687, 688, 696, 61 Pisa 585, 588, 591, 602

909

910

Marine Decapod Crustacea

Pisidia 286, 289, 290, 293 Pisinae 578, 580 Pisoides 584, 589, 591, 602 Pitho 634, 635, 638, 54 Placetron 312, 314, 315 Placopsicrangon 104, 105, 108 Plagusia 807, 808, 69 Plagusiidae 433, 802, 807, 69 Planaja 623, 624, 626, 627, 631 Planaxius 190, 193, 202 Planes 803–805, 69 Planopilumnidae 427, 720, 722 Planopilumnus 722, 723 Planotergum 622, 623, 626, 631, 53 Platepistoma 489, 490, 42 Platyactaea 752, 754, 756, 64 Platycaris 130, 136, 149 Platychelonion 722, 723 Platychirarma 813–815, 820 Platychirograpsus 802 Platydromia 446–448, 450, 452 Platyeriocheir 831–834 Platylambrus 655, 659, 660, 662, 58 Platymaia 578, 639–645, 55 Platymera 483–485 Platyozius 520–522, 530, 44 Platypilumnus 531, 532 Platypodia 787, 788 Platypodiella 787, 788, 67 Platypontonia 130, 136, 149 Platyxanthidae 422, 427, 428, 501, 508 Platyxanthus 508, 509 Pleistacantha 578, 639, 640, 642, 643, 645, 55 Pleistacanthinae 578, 639 Pleisticanthoides 578, 639, 640, 642, 643, 645 Plenotheres 864, 868, 875 Pleoticus 39, 40 Plesiomenaeus 128, 141, 149 Plesionida 278, 279, 281 Plesionika 160–162, 164, 12 Plesiopenaeus 23 Plesiopontonia 128, 136, 139, 149 Pleurocolpus 764, 766, 771 Pleuroncodes 277, 278, 281 Pleurophricus 647, 648 Pliopagurus 323–325, 333 Pliopontonia 129, 137, 142, 149, 11 Pliosoma 580, 584, 590, 602 Pliosomatinae 580 Podocatactes 493, 732, 733, 63 Podochela 611–615 Podophthalminae 700, 708 Podophthalmus 700, 708, 709, 62

Pohleus 584, 589, 603 Polybiidae 426, 431, 687, 688, 697, 61 Polybius 695, 697–699 Polycheles 259–261, 21 Polychelida 9, 11, 12, 14, 15, 183, 259 Polychelidae 259, 21 Polydectinae 750, 776 Polydectus 776 Polyonyx 286, 288, 289, 293, 28 Pomagnathus 66, 70, 72, 75 Pomatocheles 343, 344 Pomatochelinae 344 Pomatogebia 241, 243 Pontocaris 105, 106, 108 Pontonia 129, 137, 139, 149, 11 Pontonides 124, 139, 141, 149 Pontoniinae 119, 121, 122 Pontoniopsides 130, 137, 150 Pontoniopsis 130, 137, 140, 141, 150 Pontophilus 105, 106, 108 Porcellana 286, 289, 290, 293, 28 Porcellanella 287, 289, 290, 293, 28 Porcellanidae 263, 272, 286, 27, 28 Porcellanopagurus 319, 322, 334, 343 Poripontonia 129, 137, 150 Portumnus 690, 691 Portunidae 422, 426, 431, 688, 691, 700, 61, 62 Portuninae 700–703, 707–709 Portunoidea 422, 426, 431, 493, 687 Portunus 687, 700, 702–704, 709, 710, 712, 62 Potamalpheops 67, 70, 72, 76 Potamidae 424 Potamocypoda 846–848 Potamoidea 2, 422, 424 Potamonautidae 424 Poti 207, 209, 213 Poupinia 466 Poupiniidae 423, 459, 466 Praebebalia 554, 562, 571, 48 Praedatrypaea 205, 206, 210, 213 Praosia 554, 562, 564, 571, 48 Prehensilosergia 45–47 Prexylopagurus 347, 35 Priapipilumnus 669, 677, 681 Prionalpheus 66, 70, 72, 76 Prionocrangon 104, 105, 108 Prionoplax 736, 741–744, 747 Prismatopus 625, 628, 631, 53 Probeebei 302, 340, 341 Procarididae 59 Procarididea 12, 15, 59 Procaridoidea 61 Procaris 59

Index to genus-group, family-group and higher taxonomic names

Processa 169, 170 Processidae 61, 63, 168 Processoidea 63, 168 Procletes 161, 162, 164, 12 Proexotelson 828, 829 Progeryon 428, 533, 534 Progeryonidae 428, 429, 432, 511, 533, 44 Projasus 248, 249, 251 Pronotonyx 671, 682, 684 Propagurus 318, 319, 323, 334, 34 Propontonia 123, 137–139, 141, 150 Prosphorachaeus 611, 612, 615 Protoniopagurus 323, 324, 334 Protrachypene 29, 32, 35 Psalidopodidae 62, 170, 11 Psalidopodoidea 62, 170 Psalidopus 170, 11 Psathyrocaris 165–167 Psaumis 751, 764, 766, 767, 771, 66 Pseudactaea 751, 758, 759 Pseudalpheopsis 67, 70, 71, 76 Pseudaristeus 21–23 Pseudathanas 67, 72, 76 Pseudoaxiopsis 192, 194, 202 Pseudocalliax 228–231 Pseudocarcinidae 428, 500, 718, 63 Pseudocarcinoidea 428, 504, 718 Pseudocarcinus 500, 504, 718, 719, 63 Pseudocheles 102 Pseudochelidae 62, 102 Pseudoclimenes 129, 137, 142, 150 Pseudocorystes 488, 493 Pseudocoutierea 124, 137, 150 Pseudocryptochirus 797–800, 68 Pseudocryptocoeloma 671, 680, 684 Pseudodromia 446, 447, 449, 452 Pseudogaetice 831–834 Pseudogelasimus 846-848 Pseudograpsus 831–834 Pseudohapalocarcinus 797–800 Pseudohelice 826–828 Pseudohexapus 537–539 Pseudolambrus 653, 659, 662 Pseudoliomera 752, 754, 756, 64 Pseudolitochira 671, 682–684 Pseudomedaeus 780, 781, 786 Pseudomicippe 622, 623, 626, 632 Pseudomunida 269, 270 Pseudomyra 556, 562, 571 Pseudopaguristes 305, 309 Pseudopagurodes 320, 322, 326, 334 Pseudopagurus 305, 309 Pseudopalaemon 121, 132, 134

Pseudopalicus 648–650, 56 Pseudopandalus 161, 162, 164 Pseudopanopeus 741, 743, 744, 747 Pseudophilyra 557, 562, 571 Pseudopinnixa 828, 829 Pseudopontonia 120, 130, 150 Pseudopontonides 124, 137, 150 Pseudopontophilus 105, 106, 108 Pseudoporcellanella 286, 289, 290, 294 Pseudorhombila 743–745, 748 Pseudorhombilidae 430, 734, 736, 741, 777, 64 Pseudosesarma 809, 812, 813, 815, 816, 820 Pseudothelphusidae 425 Pseudothelphusoidea 2, 422, 425 Pseudoveleronia 124, 137, 140, 150 Pseudoziidae 428, 470, 723 Pseudozioidea 427, 719 Pseudozius 723, 724 Psopheticoides 520, 522 Psopheticus 524–526, 528, 44 Pterocaris 66, 69, 76 Pteromaja 581, 588, 591, 603, 51 Pteropagurus 321, 324, 334 Pteropeltarion 732, 733 Ptychognathus 830, 832–834 Puerulus 248, 249, 251, 19 Pugettia 588, 593, 594, 603 Pugnatrypaea 205, 206, 210, 213, 16 Pulcratis 787, 788, 67 Pumilopagurus 321–324, 326, 334 Pusillopagurus 320, 324, 334 Pycnocaris 121, 131 Pycnoplax 524–526, 529, 44 Pygmaeopagurus 321, 322, 326, 334 Pylocheles 343, 344, 35 Pylochelidae 264, 301, 302, 342, 35 Pylochelinae 343, 344 Pylojacquesia 346 Pylojacquesidae 345 Pylopaguridium 322, 324, 325, 335 Pylopaguropsis 318, 319, 323, 324, 335, 34 Pylopagurus 323, 325, 335 Pyrhila 559, 562, 564, 572, 48 Pyromaia 618, 619, 621, 52 Pyxidognathus 830, 832, 834 Quadrella 728–730, 63 Quadrellinae 728 Racilius 68, 70, 76 Ralumcaris 189, 193, 202, 15 Ralumnia 672 Randallia 552, 556, 562, 564, 572, 573

911

912

Marine Decapod Crustacea

Ranilia 469–472, 39 Ranina 468–470, 472, 39 Raninidae 423, 467, 468, 39 Ranininae 469, 470 Raninoida 421, 422, 466 Raninoidea 422, 431, 467 Raninoides 469–472, 39 Raninoidinae 469, 470, 472 Raoulia 512–514, 43 Raouliinae 513, 514 Raphidopus 287, 289, 290, 294 Rapimenes 126, 137, 140, 142, 150 Rapipontonia 126, 137, 140, 150 Rata 752, 754, 756 Ratha 760–762, 65 Rathbunaja 624, 627, 632 Rathbunaria 722, 723 Rathbunixa 863, 865, 867, 870, 875 Rayapinus 538, 540 Raylilia 552, 557, 562, 564, 565, 572, 48 Rayllianassa 205, 208, 209, 213 Raymanninus 691–693, 697 Raymondia 875 Raymunida 277, 279, 282, 25 Raytheres 865, 869, 875 Rectopalicus 648–650 Remiarctus 254, 255, 257 Remipedia 9, 10, 12 Retropluma 724, 725 Retroplumidae 424, 724, 63 Retroplumoidea 424, 724 Rhabdonotus 667, 673, 674 Rhadinoplax 428, 533, 534, 44 Rhinocarcinus 584, 593, 603 Rhinolambrus 656, 659, 663, 58 Rhinolithodes 313, 314, 316, 31 Rhinospinosa 609, 612, 615, 52 Rhithropanopeus 736, 737, 740, 64 Rhizopa 671, 682–684 Rhizopinae 666–668, 675, 679, 681 Rhizopoides 670, 675–677, 681, 682 Rhodochirus 323, 324, 335 Rhopalaemon 121, 132–134 Rhynchocinetes 114, 9 Rhynchocinetidae 62, 114, 9 Rhynchoplax 543–545, 548 Rhynocrangon 105, 106, 108 Richalpheus 66, 70, 72, 76 Richardina 56, 5 Richerellus 701, 705, 706 Richerius 541, 543–545, 549 Rimapenaeus 27, 30, 32, 35

Rimicaris 96–99 Rizalthus 766, 767, 771 Robertsella 742–744, 748 Robustosergia 45, 46, 48 Rochinia 588, 594, 603 Romaleon 489–491, 42 Rostronia 130, 137, 151 Rudisullianassa 205, 206, 208, 213 Rugathanas 66, 67, 70–72, 76 Ruppellioides 504, 507 Ryphila 558, 562–564, 572, 48 Sabinea 104, 105, 108 Sadayoshia 277, 279, 282, 25 Sagmariasus 248, 249, 251, 19 Saintlaurentiella 238, 239 Sakaija 624, 625, 627, 632, 54 Sakaila 474–476, 40 Sakaina 861–863, 865, 867, 875 Salaciinae 620 Salmoneus 69–71, 76, 6 Samadinia 584, 591, 594, 603, 51 Samarplax 543, 548 Sandimenes 126, 142, 151 Sandyella 123, 137, 151 Sanquerus 700, 702, 708–710, 712, 62 Sarcostraca 12 Sargassocarcinus 581, 588, 603 Sarmatium 812–814, 821, 70 Saron 83, 85, 87, 7 Sayixa 863, 865, 867, 870, 875 Scallasis 206, 208, 209, 213, 16 Scalopidia 534, 535, 44 Scalopidiidae 427, 511, 534, 44 Scammarctus 253, 254, 257, 20 Scandarma 812, 814, 821 Schizophroida 625–627, 632 Schizophrys 625, 627, 632, 54 Scintillosergia 45, 46, 48 Sclerocrangon 104–106, 108 Sclerodora 115 Scleroplax 863, 865, 875 Scopaeopagurus 319, 322, 323, 335 Scopimera 846–848 Scopimerinae 845 Scopolius 741, 743, 748 Sculptolithodes 313–315 Scutumara 830, 832–834 Scylla 687, 700, 702, 705, 708, 712, 735, 61 Scyllaridae 247, 252, 20, 21 Scyllarides 253, 255, 20 Scyllarinae 253, 256

Index to genus-group, family-group and higher taxonomic names

Scyllarus 254–257, 20 Scyra 588, 589, 604, 51 Scyramathia 588, 591, 594, 604 Scytoleptus 189, 193, 202 Segonzacia 479–491 Segonzackomaius 112, 113 Seiitaoides 624, 625, 632 Selatium 809, 812, 814, 815, 821 Selwynia 793, 794, 68 Ser 671, 682–684 Serenella 840–842, 844 Serenepilumnus 671, 676, 681 Serenius 752, 754, 756 Serenolumnus 669, 676, 681 Serenotheres 863, 864, 876 Sergestes 42–44, 48 Sergestidae 19, 20, 41, 42, 4 Sergestoidea 19, 41 Sergia 42, 43, 45, 48 Serpenthusa 498–500 Sesarma 431, 809, 811, 812, 814, 821 Sesarmidae 431, 432, 802, 809, 824, 825, 70 Sesarmoides 811, 812, 816, 821, 70 Sesarmops 809, 812, 816, 821 Sestrostoma 828, 829 Setanida 278, 279, 282 Setozius 721 Seulocia 553, 574, 575, 48 Sheniinae 846 Shenius 432, 837, 846–848 Shinkaia 284, 285, 26 Shinkaicaris 97–99 Shinobium 812, 814, 815, 821 Sicyonella 41, 43, 44, 48, 4 Sicyonia 38, 4 Sicyoniidae 19–21, 27, 38, 4 Siderochinia 588, 594, 604 Silentes 248, 249 Simocarcinus 581, 588, 604, 51 Simodorippe 435, 436, 438 Sindheres 866, 870, 876 Singhaplax 524–526, 529 Sinosesarma 813, 814, 821 Sirpus 696, 697 Soceulia 553, 574, 575 Solenocera 38–40, 4 Solenoceridae 19–21, 27, 38, 4 Solenolambrus 653, 659, 660, 663, 58 Solenopagurus 321, 335 Solenotheres 866, 869, 876 Soliella 760–762 Solinca 584, 593, 604

Solitariopagurus 319, 322, 335, 34 Sotoplacidae 427, 510, 534 Sotoplax 534, 535 Spathapagurus 321, 323, 324, 326, 335 Spelaeogriphacea 13, 14 Speloeophoroides 555, 564, 572 Speloeophorus 555, 562, 564, 572, 48 Speocarcinus 743–745, 747, 748, 64 Speodromia 446, 447, 452 Sphaerodromia 444, 446, 447, 453, 37 Sphaerodromiinae 443, 453, 455 Sphaeropeltarion 732, 733 Sphaerozius 504, 505 Sphenocarcinus 584, 586, 604 Sphenomaia 797, 798, 800 Sphenomerides 728–730 Spinicallianassa 206–209, 214 Spinolambrus 656, 659, 660, 663, 58 Spirontocaris 92, 93 Spiropagurus 321, 335, 34 Spiroplax 537, 538, 540 Spongiaxius 188, 189, 193–195, 202, 15 Spongicola 53, 54 Spongicolidae 51, 52, 5 Spongicoloides 53, 54 Spongiocaris 53, 54 Squillalbunea 297, 298 Statommatia 515–518 Stebbingdromia 444, 446–448, 452 Stegopleurodon 581, 588, 604, 51 Stegopontonia 125, 137, 151 Stelgistra 812, 814, 822 Stemonopa 297, 298 Stenalpheops 66, 70, 72, 77 Stenocionops 588, 592, 604 Stenopodidae 51, 55, 5 Stenopodidea 9, 12, 15, 51 Stenopus 55–57, 5 Stenorhynchinae 621 Stenorhynchus 465, 617–619, 621, 52 Stereomastis 259, 260, 21 Sternodromia 444, 446, 447, 452 Sternostylidae 266, 271, 23 Sternostylus 271, 1, 23 Stevea 537, 538, 540 Stilbognathus 582, 584, 605 Stilbomastax 582, 584, 591, 594, 605, 606 Stimdromia 445, 446, 453 Stimpsoplax 543–545, 548 Stomatopoda 10, 11, 13 Strahlaxiidae 18, 231, 233, 17 Strahlaxius 234, 17

913

914

Marine Decapod Crustacea

Stratiolibinia 585, 591, 594, 605, 51 Strianassa 238, 239 Striatiotes 307 Stridentes 248, 249 Strigopagurus 305–307, 309 Strobopagurus 340, 341 Stygiomysida 13, 14 Stylodactylidae 62, 171, 11 Stylodactyloidea 62, 170 Stylodactyloides 171, 172 Stylodactylus 171, 172, 11 Sulaplax 543, 548 Sulcodius 760–762 Sunipea 609, 611, 616 Symethinae 469, 470, 472 Symethis 469–472, 39 Sympagurus 340–342, 34 Synalpheus 66, 69, 72, 77, 6 Synaxidae 249 Syncarida 13, 14 Syncrangon 105, 106, 109 Systellaspis 117, 118, 9 Systroplax 520, 522 Tacitotheres 866, 868, 876 Takedactylus 793, 794 Takedana 670, 676, 677, 681 Takedax 766, 768, 771 Takedellus 839–842, 844 Takedromia 446, 448, 453 Taliepus 585, 588, 605 Tanaidacea 11, 13, 14 Tanaoa 553, 557, 564, 572, 573, 48 Tanaocheles 687 Tanaochelidae 427, 664, 687, 728 Tantulocarida 12 Tanypenaeus 29, 32, 35 Tarrasopagurus 320, 324, 336 Tasmanida 279, 282 Tasmanoplax 850–853, 72 Tastrypaea 208, 210, 214 Tauropus 667, 673, 674 Tectopontonia 124, 137, 141, 151 Teleophrys 635, 636, 639, 54 Telmatothrix 839, 841, 842, 845 Telmessus 732, 492, 493 Temnonotus 625, 632 Tenagopelta 516, 518 Tenuipedium 120, 122, 132, 134 Teramnonotus 543–545, 549 Teratomaia 622, 625, 628, 632 Tethisea 322, 233, 17

Tetraconata 9 Tetragrapsus 831–834 Tetralia 728, 63 Tetraliidae 427, 725, 727, 63 Tetralobistes 306, 307, 309 Tetraloides 328 Tetraplax 736, 741–744, 748 Tetraxanthus 743, 748 Tetrias 861, 862, 866, 867, 876 Thacanophrys 631 Thailandcallichirus 220, 221, 224 Thalamita 703, 713, 715, 62 Thalamitinae 700, 707, 712 Thalamitoides 703, 713, 715, 62 Thalamonyx 713, 715 Thalassina 240 Thalassinidae 237, 240 Thalassinidea 185, 237 Thalassocarididae 158 Thalassocaris 160–162, 164, 12 Thalassograpsinae 825, 830 Thalassograpsus 830 Thalassoplax 743–745, 748 Thaumastocaris 128, 137, 140, 151, 11 Thaumastocheles 177, 179, 180, 13 Thaumastochelidae 177 Thaumastochelopsis 177, 179, 180 Thaumastoplax 537, 538, 540 Thecaplax 512–514 Thecostaca 12 Theninae 253, 257 Thenus 253, 257, 21 Theoxapus 538, 540 Thermosbaenacea 13, 14 Thersandrus 580, 588, 594, 605 Thiidae 422, 426, 687, 688, 694, 717, 62 Thinora 92, 94 Thoe 634, 636, 639, 54 Thomassinia 216, 217 Thor 92, 94, 7 Thoracica 12 Thoracotremata 421, 430, 431, 473, 492 Thorella 84, 85, 87 Thoridae 64, 91, 7 Thorina 94 Thranita 713, 715 Thusaenys 588, 591, 592, 605 Thuylamea 68, 70, 71, 77 Thymopides 178–180 Thymops 177, 179, 180 Thymopsis 180 Thyraplax 524–526, 529

Index to genus-group, family-group and higher taxonomic names

Thyrolambrus 652, 655, 656, 58 Tiaramedon 667, 673, 674, 59 Tiarinia 580, 588, 589, 591, 592, 605, 51 Tiomanium 811, 814, 815, 822 Tisea 304, 305, 310, 340 Tlos 555, 562, 564, 573 Tmethypocoelis 846–848 Tokoyo 557, 562, 564, 573, 48 Tomopaguroides 319, 324, 336 Tomopaguropsis 319, 336 Tomopagurus 323, 325, 336 Torbenella 277, 279, 282 Toru 557, 564, 572, 573, 48 Tozeuma 85, 87, 7 Trachycaris 84, 85, 87, 7 Trachymaia 588, 593, 605 Trachypenaeinae 27 Trachypenaeopsis 29, 30, 32, 36 Trachypenaeus 30, 32, 35, 36 Trachysalambria 27, 30, 32, 36, 3 Trapezarma 813, 815, 822 Trapezia 728, 729, 731, 63 Trapeziidae 430, 725, 728, 63 Trapeziinae 728 Trapezioidea 427, 430, 725 Trapezioplax 741–744, 748 Triacanthoneus 68, 70, 77 Trichia 758, 759, 64, 65 Trichiinae 758 Trichobezoares 864, 866, 867, 876 Trichodactylidae 425 Trichodactyloidea 2, 422, 425 Trichopagurus 320, 322, 326, 336 Trichopeltariidae 425, 432, 493, 731, 63 Trichopeltarioidea 425, 488, 493, 731 Trichopeltarion 732, 733, 63 Trichoplatus 639, 640, 642, 643, 645 Tridacnatheres 864, 865, 876 Tridiscias 101 Trierarchus 713, 715 Trigonoplax 543–545, 549 Triodonthea 274–276 Trionectes 701, 710, 712 Trissoplax 520–522, 44 Tritodynamia 430, 849–851, 853 Tritodynamiinae 849 Tritoplax 538, 540 Trizocarcinus 520–522 Trizocheles 343–345, 35 Trizochelinae 343–345 Trizopagurus 305, 310 Troglindicus 122, 132, 133, 135

Troglocarcinus 797, 798, 800 Troglocubanus 122, 123, 133, 135 Troglomexicanus 120, 122, 133, 135 Trogloplacinae 514, 515, 518 Trogloplax 514, 515, 518 Trypaea 205, 206, 214 Tsunogaipagurus 340, 342 Tubicolixa 863, 866, 876 Tubuca 856–858, 72 Tuerkayana 802 Tuerkayogebia 241, 242, 244 Tuleariocaris 125, 137, 151 Tumidodromia 444, 446, 447, 453, 37 Tumidotheres 868, 869, 877 Tumulosternum 625–627, 633 Tunebia 776 Tunedromia 444, 446, 453 Tunepugettia 588, 593, 594, 605, 51 Tunicotheres 863, 866, 877 Turleania 320, 324, 336 Tutankhamen 655, 659, 660, 663 Tweedieia 760–762, 65 Tyche 582, 585, 594, 606 Tychinae 578, 580 Tylaspis 340, 342 Tylocarcinus 585, 588, 591, 606, 51 Tylodiplax 840, 842, 844, 845 Tymolus 436–438, 36 Typhlatya 95 Typhlocarcinodes 512–514 Typhlocarcinops 671, 682–684, 59 Typhlocarcinus 671, 682–684 Typhlocarididae 2, 63, 119 Typhlocaris 119 Typhlopagurus 340, 342 Typton 123, 137, 151, 11 Typtonoides 123, 137, 151 Typtonomenaeus 128, 137, 141, 151 Typtonychus 123, 137, 138, 141, 151 Uca 855–859, 72 Ucides 855, 856, 858, 859, 72 Ucidinae 855, 858 Ucinae 856, 859 Uhlias 555, 562, 573 Ulloaia 287, 289, 294 Umalia 469–472, 39 Unesconia 128, 137, 139, 151 Unguicaris 126, 137, 138, 142, 151 Upogebia 241–244, 18 Upogebiidae 77, 237, 241, 18 Urashima 557, 562, 573, 48

915

916

Marine Decapod Crustacea

Urnalana 553, 574, 574, 48 Urocaridella 121, 132, 135 Urocaris 125, 137, 152 Uroptychodes 267, 268, 22 Uroptychus 267, 268, 22 Uruma 793, 794 Utica 831, 832, 834 Utinomiella 797–800, 68 Varuna 831–833, 835, 71 Varunidae 432, 433, 802, 804, 861, 71 Varuninae 431, 825, 830 Veleronia 124, 137–139, 141, 152 Veleroniopsis 124, 137, 152 Vellodius 760–762 Vellumnus 536, 667, 669, 676, 677, 681 Velolambrus 654, 659, 660, 663 Venitus 851–853, 72 Vercoia 104, 105, 109 Vetericaris 59 Vexillipar 69–71, 77 Viaderiana 667, 670, 676, 677, 681 Villoplax 520, 522 Vir 122, 137, 141, 152 Viridotheres 866, 869, 877 Visayax 766, 767, 771 Visayeres 865, 866, 877 Vitjazmaia 578, 639, 640, 642, 644, 645, 55 Vojmirophthalmus 700, 709 Vounopagurus 318, 330 Vulcanocalliax 218 Vultocinidae 429, 536, 44 Vultocinus 536, 44 Waldola 123, 137, 152 Waldotheres 866, 869, 877 Wardoxanthops 777, 780, 786 Willemoesia 259–261 Williamstimpsonia 779, 780, 786 Wolffogebia 241, 242, 244 Xaiva 690, 691 Xanthasia 863, 864, 867, 877 Xanthias 780, 781, 786, 67 Xanthidae 119, 430, 432, 728, 734–736, 741, 749, 64–67 Xanthinae 751, 776 Xantho 780, 781, 786

Xanthodius 778, 780, 782, 786 Xanthoidea 430, 733 Xeinostoma 436–438, 36 Xeinostomatinae 435, 436, 438 Xenocarcinus 585, 589, 591, 606, 51 Xenocrate 520–522, 44 Xenograpsidae 433, 802, 836 Xenograpsus 836, 837 Xenophthalmidae 432, 838, 860 Xenophthalminae 860, 861 Xenophthalmodes 666, 671, 682–684 Xenophthalmodinae 666, 682 Xenophthalmus 682, 860, 861 Xeruca 856–858 Xestopilumnus 668, 676, 677, 681 Xiphocarididae 2, 61, 63, 110 Xiphocaris 110 Xiphonectes 701, 703, 707, 709–712 Xiphopenaeus 27, 29, 32, 36 Xlumnus 669, 676, 677, 681 Xylocheles 243–245 Xylopaguridae 302, 346, 35 Xylopagurus 346, 347 Xynomaia 797–799, 801 Yagerocaris 68, 70, 72, 77, 6 Yaldwynopsis 461, 462, 464, 38 Yemenicaris 128, 137, 139, 152 Zalasiinae 758 Zalasius 759 Zaops 868, 869, 877 Zarenkolambrus 654, 659, 663 Zariquieyon 692, 693 Zebrida 668, 673, 674, 59 Zebridonus 668, 673, 674 Zehntneria 684 Zehntneriana 671, 682, 684 Zenopontonia 125, 140, 142, 152, 11 Zibrovia 797, 801 Zosiminae 751, 786 Zosimus 787–789, 67 Zoukaris 127, 137, 142, 152 Zozymodes 787, 789, 67 Zygita 703, 713, 715, 62 Zygopa 297, 298