Handbook of Zoology: Volume 2 Morphology and Systematics (Elateroidea, Bostrichiformia, Cucujiformia partim) 9783110911213, 9783110190755

Table of contents :
Editors’ preface
Contributors
1. Changes in classification and list of families and subfamilies
2. Glossary of Morphological Terms
3. Adephaga (Addendum) Introduction and Phylogeny
4. Elateroidea Introduction, Phylogeny
5. Derodontiformia Introduction, Phylogeny
6. Bostrichiformia Latreille 1802
7. Cucujiformia Lameere, 1938 Introduction, Phylogeny
8. Lymexloidae Fleming, 1821
9. Cleroidea Latreille, 1802 Introduction and Phylogeny
10. Cucujoidea, Latreille, 1802 Introduction, Phylogeny
11. Tenebrionoidea Introduction, Phylogeny
Taxonomic Index

Citation preview

Handbook of Zoology

Arthropoda: Insecta Coleoptera, Beetles Volume 2: Morphology and Systematics (Elateroidea, Bostrichiformia, Cucujiformia partim)

Handbook of Zoology Founded by Willy Kükenthal Continued by M. Beier, M. Fischer, J.-G. Helmcke, D. Starck, H. Wermuth

Arthropoda: Insecta Editors Niels P. Kristensen & Rolf G. Beutel

DE GRUYTER

Richard Α. Β. Leschen Rolf G. Beutel John F. Lawrence (Volume Editors)

Coleoptera, Beetles Volume 2: Morphology and Systematics (Elateroidea, Bostrichiformia, Cucujiformia partim)

Adam Slipiriski (Associate Editor)

DE GRUYTER

Scientific Editors Richard Α. B. Leschen New Zealand Arthropod Collection Private Bag 9 2 1 7 0 Auckland, New Zealand

John F. Lawrence 130 Hartwig Road Gympie, Queensland, 4 5 7 0 , Australia

Rolf G.Beutel Institut für Spezielle Zoologie und Evolutionsbiologie Friedrich-Schiller-Universität Jena 0 7 7 4 3 Jena, Germany

Adam Slipiriski Research Entomologist CSIRO Entomology GPO Box 1 7 0 0 Canberra ACT 2 6 0 1 , Australia

ISBN 978-3-11-019075-5

© 2010 by Walter de Gruyter GmbH & Co. KG, Berlin/ New York.

Library of Congress Cataloging-in-Publication

Data

Handbook of zoology. Vol. 2, Coleoptera / (edited) by John F. Lawrence... [et al]. p. cm. Includes index. ISBN 978-3-11-019075-5 (alk. paper) 1. Zoology. 2. Beetles. I. Lawrence, John F. (John Francis), 1 9 3 4 QL573.H36 2009 595.76-dc22

2009043741

Bibliographic information published by the Deutsche Nationalbibliothek

The Deutsche Nationalbibliothek lists this publication in the Deutsche Nationalbibliografie; detailed bibliographic data are available in the Internet at http://dnb.d-nb.de.

Typesetting: Compuscript Ltd. Printing and binding: Druckhaus „Thomas Müntzer" GmbH, Bad Langensalza. © Printed on acid-free paper. Printed in Germany www.degruyter.com

Editors7 preface

Much of what had to be said in a preface was done so in the Editors' Preface of volume 1 and will not be repeated here. What follows is an annotation. There are two more installments to come, with volume 3 (Phytophaga) in progress and volume 4 which will cover special topics of morphology, natural history, and evolution. The detailed family by family review of the groups of beetles covered in this volume turned out to be a challenging scientific project. The tremendous difficulty to find suitable authors for many chapters highlights a dramatic loss of expertise, which is in stark contrast with what gets published and discussed openly in the context of the ongoing biodiversity crisis, especially as it relates to systematics and taxonomy. Without the encyclopedic knowledge of an increasingly smaller circle of outstanding coleopterists, the completion of the 116 chapters of this volume would definitely not have been possible. The groups included are two recently described families of Adephaga (chapters 2-3) and the polyphagan superfamilies Elateroidea, Derodontoidea, Bostrichoidea, Lymexyloidea, Cleroidea, Cucujoidea and Tenebrionoidea. Two introductory chapters are also included, one of which is an update on beetle classification, and the other a much-needed glossary of morphological terms. While classifications will change almost on a yearly basis due to new phylogenetic data, morphological terminology is an old calloused dog that is entrenched by tradition with many beetle specialists having their own sets of terms that are not used in other groups (the genitalic terminology of Coccinellidae is one example). During the course of editing and writing of the chapters the mix of terminology was frustratingly obvious, and not the fault of the authors themselves. Because homology is the ultimate scientific concept that underpins phylogeny and classification, the editors decided that a glossary would help to improve the communication among coleopterists and readers of the Handbook. You will note that the editorship has changed since volume 1, a combination of editors that suit our expertise. In vol. 1, Rolf G. Beutel was lead editor because of his focus on Adephaga, and here, Richard A. B. Leschen takes lead as his work has been mainly on Cucujoidea. Then, it was decided to add John F. Lawrence for his broad expertise

on much of the Polyphaga, a knowledge asset that pushed volume 2 towards its logical and most successful limit. Meanwhile, Adam Slipiiiski was also brought to the fold to assist with the management of Cucujoidea and as associate editor of Tenebrionoidea. Apart from chapter author and co-author, Adam, with the help of Anne Hastings, had managed hundreds of illustrations for many of the family chapters. Since we had begun work on volume 2, several key coleopterists have passed away, among them Alistair S. Ramsdale (Montana State University) who had contributed elateroid chapters to this volume. Another great loss was the untimely death of the outstanding staphylinid specialist Dr. Steve Ashe (University of Kansas) to whom we dedicate this volume. He was advisor to Richard A. B. Leschen and supervisor of Marc A. Branham, who is one of our vol. 2 authors. Shortly before this volume was completed, Robert E. Roughley passed away; he was a specialist on Adephaga and was coauthor on the Gyrinidae chapter in Volume 1. Lastly we remember Hiroyuki Sasaji, whose main works concerned Cucujoidea and Tenebrionoidea, especially Coccinellidae, which are covered in this volume. First of all, our greatest thanks go to the authors who have contributed to volume 2, without you coleopteran systematics would not be such a vibrant field of study and this project would have not been completed. Second, we would like to thank our institutions and funding agencies and naturally we would like to express our gratitude to our friends, colleagues, and families who supported us during the writing and editing of volume 2. We lastly thank Stephanie Dawson at De Gruyter by assisting with the overall editorial process and especially the final stages of proofing. Though Stephanie politely kept asking us about deadlines which were off by almost two years, she had to succumb to the nuances of coleopterists with patience and respect and we were happy to work alongside her in this demanding and large project. Richard A. B. Leschen Rolf G. Beutel John F. Lawrence Adam Slipiiiski (associate editor of Tenebrionoidea)

Contents

Editors' preface Contributors 1.

2.

3.

3.1.

3.2.

4.

Changes in classification and list of families and subfamilies John F. Lawrence, RolfG. Beutel, Richard A. B. Leschen and Adam Slipifiski Glossary of Morphological Terms John F. Lawrence, RolfG. Beutel, Richard A. B. Leschen and Adam Slipifiski Adephaga (Addendum) Introduction and Phylogeny RolfG. Beutel, Ignacio Ribera and Michael Balke Aspidytidae Ribera, Beutel, Balke and Vogler, 2002 RolfG. Beutel, Michael Balke and Ignacio Ribera Meruidae Spangler and Steiner 2005 RolfG. Beutel, Michael Balke and Warren E. Steiner

V XI

1

9

21

28

35

4.1.

Rhinorhipidae Lawrence, 1988 John F.Lawrence

38

4.2.

Artematopodidae Lacordaire, 1857 John F. Lawrence

42

4.3.

Brachypsectridae Horn, 1881 Cleide Costa, Sergio A. Vanin, John F. Lawrence, Sergio Ide and MarcA.Branham

54

4.5.

Eucnemidae Eschscholtz, 1829 JyrkiMuona

61

4.6.

Throscidae Laporte, 1840 JyrkiMuona, John F. Lawrence and Adam Slipifiski Elateridae Leach, 1815 Cleide Costa, John F. Lawrence and Simone Policena Rosa

103

4.9.

Family Drilidae Blanchard, 1845 Ladislav Bocak, Marc A. Branham and Robin Kundrata

104

4.10.

Family Omalisidae Lacordaire, 1857 Ladislav Bocak and Milada Bocakova . . . . 1 1 0

4.11.

Lycidae Laporte, 1836 Ladislav Bocak and Milada Bocakova

114

4.12.

Telegeusidae Leng, 1920 John F. Lawrence

123

4.13.

Phengodidae LeConte, 1861 Cleide Costa and Santiago Zaragoza-Caballero

126

Rhagophthalmidae Olivier, 1907 Itsuro Kawashimajohn F. Lawrence and Marc A. Branham

135

4.15.

Lampy ridae Latreille, 1817 Marc A. Branham

141

4.16.

Omethidae LeConte, 1861 AlistairS.Ramsdale

149

4.17.

Cantharidae Imhoff, 1856 AlistairS.Ramsdale

153

4.18.

Elateriformia Incertae Sedis John F. Lawrence, Itsuro Kawashima and Marc A. Branham

162

Derodontiformia Introduction, Phylogeny John F. Lawrence, RolfG. Beutel and Richard A.B. Leschen

179

Derodontidae LeConte, 1861 Richard A. B. Leschen and RolfG. Beutel

180

4.14.

5.

5.1.

5.2.

Nosodendridae Erichson, 1846 Richard A.B. Leschen and Rolf G. Beutel. . .185

5.3.

Jacobsoniidae Heller, 1926 John F. Lawrence and Richard A.B. Leschen

190

Bostrichiformia Latreille 1802 John F. Lawrence, Richard A. B. Leschen and RolfG. Beutel

197

Dermestidae Latreille, 1804 John F. Lawrence and Adam Slipifiski

198

Endecatomidae LeConte, 1861 John F. Lawrence

206

47

Cerophytidae Latreille, 1834 Cleide Costa, Sergio A. Vanin, John F. Lawrence and Sergio Ide

4.7.

Plastoceridae Crowson, 1972 MarcA.Branham

21

Elateroidea Introduction, Phylogeny John F. Lawrence, Ladislav Bocak, Milada Bocakova, RolfG. Beutel and JyrkiMuona

4.4.

4.8.

69

75

6.

6.1.

6.2.

VIII 6.3. 6.4.

7.

Contents Bostrichidae Latreille, 1802 ]ohnF. Lawrence

209

Ptinidae Latreille, 1802 T. Keith Philips and Karen LBell

217

CucujiformiaLameere, 1938 Introduction, Phylogeny Richard Α. B. Leschen and Adam Slipinski

10.3.

Helotidae Reitter, 1876/Chapuis, 1876 John F. Lawrence, Adam Slipinski and Chi-FengLee 292

10.4.

Protocucujidae Crowson, 1954 Adam Slipmki and RolfG. Beutel

296

SphindidaeJacquelinduVal, 1861 Juanita A. Forrester and Joseph V.McHugh

300

10.5. 227 10.6.

8.

9.

9.1.

Lymexloidae Fleming, 1821 John F. Lawrence Cleroidea Latreille, 1802 Introduction and Phylogeny Richard Α. B. Leschen Phloiophilidae Kiesenwetter, 1863 John F. Lawrence and Richard Α. B. Leschen

229

237

Trogossitidae Fabricius, 1801 JiriKolibdc and Richard A.B. Leschen . . . . 2 4 1

9.3.

Chaetosomatidae Crowson, 1952 Richard Α. B. Leschen

247

9.4.

Metaxinidae Kolibäc, 2004 JiriKolibdc and Richard A.B. Leschen . . . . 2 5 0

9.5.

Thanerocleridae Chapin, 1924 JiriKolibdc and Richard A.B. Leschen . . . . 2 5 3

9.6.

Cleridae Latreille, 1802 JiriKolibdc

9.8.

9.9.

9.10. 9.11.

10.

10.1.

10.2.

Acanthocnemidae Crowson, 1964 John F. Lawrence and Richard Α. B. Leschen Phycosecidae Crowson, 1952 Richard Α. B. Leschen and Rolf G.Beutel

10.8. 10.9.

239

9.2.

9.7.

10.7.

257

262

265

Prionoceridae Lacordaire, 1857 John F. Lawrence and Richard Α. B. Leschen

268

MauroniscidaeMajer, 1994 John F. Lawrence

271

Melyridae Leach, 1815 John F. Lawrence and Richard Α. B. Leschen

273

Biphyllidae LeConte, 1861 Andrew R. Cline and FloydW.Shockley

. . .306

Erotylidae Leach, 1815 Richard Α. B. Leschen, PaulE. Skelley and Joseph V. McHugh

311

Monotomidae Laporte, 1840 Yves Bousquet

319

Hobartiidae Sen Gupta and Crowson, 1966 Wioletta Tomaszewska and Adam Slipifiski 324

10.10. Cryptophagidae Kirby, 1837 Richard Α. B. Leschen

327

10.11. Agapythidae Sen Gupta and Crowson, 1969 Richard Α. Β. Leschen and John F. Lawrence 334 10.12. Priasilphidae Crowson, 1973 Richard A. B. Leschen and John F. Lawrence

336

10.13. Phloeostichidae Reitter, 1911 John F. Lawrence and Adam Slipinski

340

10.14. Silvanidae Kirby, 1837 Michael C. Thomas and Richard A. B. Leschen

346

10.15. Cucujidae Latreille, 1802 Michael C. Thomas and Richard A. B. Leschen

350

10.16. Myraboliidae Lawrence and Britton, 1991 Adam Slipinski, Wioletta Tomazsewska and John F. Lawrence 354 10.17. Cavognathidae Sen Gupta and Crowson, 1966 Adam Slipinski and Wioletta Tomaszewska

357

10.18. Lamingtoniidae Sen Gupta and Crowson, 1969 John F. Lawrence and Richard A. B. Leschen

360

Boganiidae Sen Gupta and Crowson, 1966 John F. Lawrence and Adam Slipinski 282

10.19. Passandridae Blanchard, 1845/ Erichson, 1845 Daniel Burckhardt and Adam Slipinski

363

ByturidaeJacquelinduVal, 1858 Andrew R. Cline, Michael A. Goodrich and Richard Α. B. Leschen

10.20. Phalacridae Leach, 1815 John F. Lawrence, Matthew L. Gimmel and Warren Ε. Steiner, Jr.

368

Cucujoidea, Latreille, 1802 Introduction, Phylogeny Richard Α. B. Leschen and Adam Slipinski

281

286

Contents

IX

10.21. Propalticidae Crowson, 1952

11.3. Pterogeniidae Crowson, 1953

John F. Lawrence

374

11.4.

10.22. Laemophloeidae Ganglbauer, 1899 Michael C. Thomas and Richard A. B. Leschen

376

381

Richard A. B. Leschen, Matthew L. Gimmel and Adam Slipinski

383

386

. .390

10.27. Smicripidae Horn, 1879 407

411

422

432

435

10.32. Endomychidae Leach, 1815 442

454

10.34. Corylophidae LeConte, 1852

11.

472

481

Tenebrionoidea Introduction, Phylogeny John F. Lawrence, Darren A. Pollock and Adam Slipinski

487

Richard A. B. Leschen and Adam Slipinski

559

John F. Lawrence, Adam and Mario Elgueta

Slipinski 563

John F. Lawrence and Richard A. B. Leschen

567

John F. Lawrence and Adam Slipinski

571

11.14. Tenebrionidae Latreille, 1802

574

11.15. Prostomidae G. G. Thomson, 1859 659

11.16. Synchroidae Lacordaire, 1859 Adam Slipinski and John F. Lawrence

667

11.17. Stenotrachelidae C. G. Thomson, 1859 John F. Lawrence and Adam Slipinski

670

John F. Lawrence and Adam Slipinski

674

11.19. Meloidae Gyllenhal, 1810 491

11.2. Archeocrypticidae Kaszab, 1964 John F. Lawrence

548

11.18. Oedemeridae Latreille, 1810

11.1. Mycetophagidae Leach, 1815 John F. Lawrence and Richard A. B. Leschen

Adam Slipinski and John F. Lawrence

Ainsley Seago andRolfG.Beutel

10.35. Latridiidae Erichson, 184 Christophers. Hartley and JosephV.McHugh

538

11.9. Zopheridae Solier, 1834

Eric G. Matthews,John F. Lawrence, Patrice Bouchard, Warren E. Steiner, Jr. and Adam Slipmki

10.33. Coccinellidae Latreille, 1802

Adam Slipinski, John F. Lawrence and Andrew R. Cline

11.8. Ripiphoridae Gemminger and Harold, 1870 (Gerstaecker, 1855)

11.13. Trachelostenidae Lacordaire, 1859

10.31. Discolomatidae Horn, 1878

Adam Slipinski and Wioletta Tomaszewska

533

11.12. Chalcodryidae Watt, 1974

10.30. Alexiidae Imhoff, 1856

Wioletta Tomaszewska

JohnF. Lawrence and Adam Slipinski

11.11. Promecheilidae Lacordaire, 1859

10.29. Cerylonidae Billberg, 1820

Andrew R. Cline and Adam Slipinski

520

11.10. Ulodidae Pascoe, 1869

10.28. Bothrideridae Erichson, 1845

Adam Slipinski and Wioletta Tomaszewska

and

John F. Lawrence, Zachary H. Falin and Adam Slipinski

10.26. Nitidulidae Latreille, 1802

Adam Slipinski and JohnF. Lawrence

514

11.7. Mordellidae Latreille, 1802

10.25. Kateretidae Erichson in Agassiz, 1846

Adam Slipinski, Nathan Lord and John F.Lawrence

504

11.6. Melandryidae Leach, 1815 Nikolai B. Nikitsky Darren A. Pollock

10.24. Cyclaxyridae Gimmel, Leschen & Slipinski, 2009

Andrew R. Cline

Ciidae Leach in Samouelle, 1819 John F. Lawrence and Cristiano Lopes-Andrade

John F. Lawrence and Richard A.B. Leschen

John F. Lawrence and Adam Slipinski

Josef Jelinek, Chris Carlton, AndrewR.ClineandRichardA.B.Leschen.

501

11.5. Tetratomidae Billberg, 1820

10.23. Tasmosalpingidae Lawrence and Britton, 1991

Josef Jelinek and Andrew R. Cline

John F. Lawrence

Marco A. Bologna, Federica Turco and John D.Pinto

681

11.20. Mycteridae Blanchard, 1845 496

Darren A. Pollock

693

χ

Contents

11.21. Boridae C. G. Thomson, 1859 Darren A. Pollock

11.22. Trictenotomidae Blanchard, 1845 Darren A. Pollock and Dmitry Telnov . .

11.23. Pythidae Solier, 1834 Darren A. Pollock

11.24. Pyrochroidae Latreille, 1807 DanielK. Young and Darren A.Pollock. .

.699

Donalds. Chandler

729

11.27. Aderidae Winkler, 1927 .704

John F. Lawrence and Adam Slipifiski

.708

11.28. Scraptiidae Mulsant, 1856

.715

11.29. Tenebrionoidea Incertae sedis

11.25. Salpingidae Leach, 1815 John F. Lawrence, Adam Slipinski, Darren A. Pollock and Hermes Escalona .

11.26. Anthicidae Latreille, 1819

.722

John F. Lawrence and Adam Slipmski John F. Lawrence, Hermes Escalona and Richard Α. B. Leschen

Taxonomic Index

741 746

750

761

Contributors

Michael Balke

Department of Entomology The Natural History Museum Cromwell Rd., London SW7 5BD, England e-mail: [email protected]

Rolf G. Beutel

Institut für Spezielle Zoologie und Evolutionsbiologie Friedrich-Schiller-Universität Jena 07743 Jena, Germany e-mail: [email protected]

Ladislav Bocak

Department of Zoology Faculty of Science Tr. Svobody 26 7 7 1 4 6 Olomouc, Czech Republic e-mail: [email protected]

Milada Bocakova

Department of Biology Faculty of Education Purkrabska 2 7 7 1 4 0 Olomouc, Czech Republic e-mail: [email protected]

Marco A. Bologna

Dipartimento di Biologia ambientale Universitä Roma Tre Viale Marconi 446 00146 Roma, Italy e-mail: [email protected]

Patrice Bouchard

Canadian National Collection of Insects, Arachnids and Nematodes Agriculture and Agri-Food Canada K. W. Neatby Building 960 Carling Avenue Ottawa, Ontario K1A 0C6, Canada e-mail: [email protected]

Yves Bousquet

Agriculture and Agri-Food Canada Central Experimental Farm Ottawa, Ontario K1A 0C6, Canada e-mail: [email protected]

Marc A. Branham

University of Florida Department of Entomology and Nematology Natural Area Drive P.O. Box 110620 Gainesville, Florida 32611-0620, USA e-mail: [email protected]

Daniel Burckhardt

Naturhistorisches Museum Augustinergasse 2 CH-4001 Basel, Switzerland e-mail: [email protected]

Chris Carlton

Hoi ton Professor of Agriculture Director, Louisiana State Arthropod Museum Department of Entomology, LSU Baton Rouge, Louisiana 70803-1710, USA e-mail: [email protected]

Donald S. Chandler

Department of Biological Sciences Spaulding Hall, 38 Academic Way University of New Hampshire Durham, New Hampshire 03824, USA e-mail: [email protected]

Andrew R. Cline

Senior Insect Biosystematist - Supervisor Plant Pest Diagnostics Center California Dept. of Food & Agriculture 3294 Meadowview Road Sacramento, California 95832-1448, USA e-mail: [email protected]

Cleide Costa

Laboratörio de Sistemätica, Evolugäo e Bionomia de Coleoptera Divisäo Cientifica - Setor de Invertebrados Museu de Zoologia da Universidade de Säo Paulo (MZUSP) Avenida Nazare, 481 - Ipiranga CEP 04263-000 Caixa Postal 42494 - CEP 04218-970 Säo Paulo, SP, Brazil e-mail: [email protected]

Mario Elgueta

Seccion Entomologia Museo Nacional de Historia Natural Casilla 787, Santiago, Chile e-mail: [email protected]

Hermes Escalona

Museo del Instituto de Zoologia Agricola Francisco Fernandez Yepez (ΜΙΖΑ) Facultad de Agronomia UCV-Maracay 2101-A, Apdo. 4579 Estado Aragua, Venezuela e-mail: [email protected]

XII

Zachary Η. Falin Division of Entomology, KUNHM/BRC Snow Entomological Collection 1501 Crestline Dr. Suite 140 University of Kansas Lawrence, Kansas 66049, USA e-mail: [email protected] Juanita A. Forrester Department of Entomology University of Georgia Athens, Georgia 30602-2603, USA e-mail: [email protected] Matthew L. Gimmel Louisiana State Arthropod Museum Department of Entomology, LSU 402 Life Sciences Building Baton Rouge, Louisiana 70803, USA e-mail: [email protected] Michael A. Goodrich Eastern Illinois University Biological Sciences Department Life Sciences Bldg. 2070 600 Lincoln Ave. Charleston, Illinois 61920, USA e-mail: [email protected] Christopher S. Hartley Missouri Botanical Garden 4344 Shaw Boulevard St. Louis, Missouri 63110, USA e-mail: [email protected] Sergio Ide Centro de Pesquisa e Desenvolvimento de Sanidade Vegetal, Instituto Biologico Av. Conselheiro Rodrigues Alves 1252 CEP 04014-900 Säo Paulo, SP, Brazil e-mail: [email protected] Josef Jelinek Department of Entomology National Museum 148 00 Praha 4 - Kunratice I, Czech Republic e-mail: jj [email protected] Itsuro Kawashima Nagasawa 1-80-9, Tokosuka Kanagawa 239-0842, Japan e-mail: [email protected] Jiri Kolibäc Moravian Museum, Entomology Hviezdoslavova 29a CZ - 627 00 Brno, Czech Republic e-mail: [email protected] Robin Kundrata Department of Zoology Faculty of Science

Contributors

tr. Svobody 26 7 7 1 4 6 Olomouc, Czech Republic e-mail: [email protected] John F. Lawrence 130 Hartwig Road Gympie, QLD 4570, Australia e-mail: [email protected] Chi-Feng Lee Applied Zoology Division, Taiwan Agricultural Research Institute 189 Chung-Cheng Road Taichung 413, Wufeng. Taiwan e-mail: [email protected] Richard A. B. Leschen New Zealand Arthropod Collection, Private Bag 92170 Auckland, New Zealand e-mail: [email protected] Cristiano Lopes-Andrade Departamento de Biologica Animal Universidade Federal de Vicosa 36570-000 Vicosa - MG, Brazil e-mail: [email protected] Nathan Lord University of Georgia Department of Entomology 413 Biological Sciences Building 30602 Athens, Georgia, USA e-mail: [email protected] Eric G. Matthews South Australian Museum North Terrace Adelaide, SA 5000, Australia e-mail: [email protected] Joseph V. McHugh Department of Entomology University of Georgia Athens, Georgia 30602-2603, USA e-mail: [email protected] Jyrki Muona Zoological Museum P.O. Box 17 FI-00014 University of Helsinki, Finland e-mail: [email protected] Nikolai B. Nikitsky Zoological Museum of Moscow Lomonosov State University Bolshaya nikitskaya str. 6 125009 Moscow, Russia e-mail: [email protected]

XIII

Contributors

John D. Pinto Department of Entomology University of California Riverside, California, USA e-mail: [email protected] T. Keith Philips Systematics and Evolution Laboratory Department of Biology Western Kentucky University 1906 College Heights Blvd. Bowling Green, Kentucky 42101-3576, USA e-mail: [email protected] Darren A. Pollock Natural History Museum Department of Biology Eastern New Mexico University Portales, New Mexico 88130, USA e-mail: [email protected] Alistair Ramsdalet Montana Entomology Collection Montana State University P.O. Box 173020 Bozeman, Montana 59717-3020, USA Ignacio Ribera Departamento de Biodiversidad y Biologia Evolutiva Museo Nacional de Ciencias Naturales Jose Gutierrez Abascal 2 28006 Madrid, Spain e-mail: [email protected] Simone Policena Rosa Museu de Zoologia, Universidade de Säo Paulo Avenida Nazare, 481, - Ipiranga - CEP 04263-000 Säo Paulo, SP, Brazil e-mail: [email protected] Ainsley Seago University of California, ESPM 137 Mulford Hall, c/o Will Lab 94720 Berkeley, California, USA e-mail: [email protected] Floyd W. Shockley University of Georgia 413 Biological Sciences Building 30602 Athens, Georgia, USA e-mail: [email protected] Paul E. Skelley Florida State Collection of Arthropods 1911 SW 34th St Gainesville, Florida 32614-7100, USA e-mail: [email protected] Adam Slipiiiski CSIRO Entomology GPO Box 1700

Canberra ACT 2601, Australia e-mail: [email protected] Warren E. Steiner, Jr. National Museum of Natural History Department of Entomology PO Box 37012 MRC 187 Washington DC 20013-7012, USA e-mail: [email protected] Dmitry Telnov Rigas rajons, Stopinu novads, Dzidrinas, Darza iela 10 LV-2130, Latvia e-mail: [email protected] Michael C. Thomas Florida State Collection of Arthropods Florida Department of Agriculture & Consumer Services P.O. Box 147100 Gainesville, Florida 32614-7100, USA e-mail: [email protected] Wioletta Tomaszewska Museum and Institute of Zoology Polish Academy of Sciences Wilcza 64 PL 00-679 Warsaw, Poland e-mail: [email protected] Federica Turco Biodiversity, Entomology, Queensland Museum, South Bank P.O. Box 3300 South Brisbane, QLD 4101, Australia e-mail: [email protected] Sergio A. Vanin Departamento de Zoologia, Instituto de Biociencias, Universidade de Säo Paulo, Rua do Matäo, Travessa 14,101, CEP 05508-900 Säo Paulo, SP, Brazil e-mail: [email protected] Daniel K. Young UW Insect Research Collection (WIRC) 445 Russell Laboratories Department of Entomology 1630 Linden Drive University of Wisconsin Madison, Wisconsin 53706, USA e-mail: [email protected] Santiago Zaragoza-Caballero Colecciön Nacional de Insectos, Departamento de Zoologia, Instituto de Biologia Universidad Nacional Autönoma de Mexico. Apartado postal 70-153, 04510 Mexico, D. F. Mexico e-mail: [email protected]

1. Changes in classification and list of families and subfamilies John F. Lawrence, Rolf G. Beutel, Richard A. B. Leschen and Adam Slipmski

The classification used in Handbook of Zoology, part 38.1 (Coleoptera, Beetles, Vol. 1) was based primarily on that of Lawrence & Newton (1995) and Lawrence et al. (1999), with a few additions or modifications based on subsequent work. Further changes and additions to that family list based on more recent publications are listed below: Artematopodidae. The tribe Allopogonini Crowson was elevated to subfamily level by Lawrence (2005a). Eucnemidae. The subfamily Anischiinae Fleutiaux, recognized at the family level in 38.1, was moved into the family Eucnemidae by Lawrence et al. (2007) and considered to be a subfamily well within the major eucnemid clade (excluding Perothopinae, Phyllocerinae, Palaeoxeninae, Pseudomeninae and Phlegoninae). Elateridae. The subfamily Tetralobinae was reduced to a tribe within Agrypninae by Costa et al. (1992, 1994). Campyloxeninae Costa (1975) and Morostominae Dolin (2000) were added as subfamilies. Lycidae. The new subfamily and tribal classification proposed by Bocak & Bocakova (2008) and based in part on Bocakova et al. (2007) was used in Chapter 4.11. Phengodidae. Mastinocerinae LeConte and Penicillophorinae Paulus were recognized at the subfamily level. Rhagophthalmidae. Although treated as a separate family here, as in 38.1, some molecular evidence (Bocakova et al. 2007) suggests that this group does belong within Phengodidae, as suggested by Crowson (1972). Lampyridae. The constitution of this family used in Chapter 4.15 is based on cladograms produced by Branham & Wenzel (2001, 2003); thus several formerly included taxa within Pterotinae, Ototretinae and Ototretadrilinae have been discussed in a separate Chapter (4.18) dealing with Elateriformia incertae sedis. Subfamilies remaining in Lampyridae are the Psilocladinae (= Cyphonocerinae; Jeng et al. 1998), Amydetinae, Lampyrinae, Photurinae and Luciolinae.

Elateriformia incertae sedis. Chapter 4.18 includes elateriform taxa whose family-group relationships are in doubt. All of these taxa belong within Elateroidea with the exception of Podabrocephalidae, which appears to be related to Ptilodactylidae. All of the taxa excluded from Lampyridae by Branham & Wenzel (2001, 2003) are also included here, as well as the doubtful throscid genus Νeocrowsonium Kistner & Abdel-Galil and the subfamily Cydistinae Paulus (1972), formerly included in Karumiidae. Derodontiformia. The placing of Derodontidae, Nosodendridae and Jacobsoniidae in a separate superfamily increases the likelihood that Bostrichoidea is a monophyletic group. It is not at all certain, however, that the three families placed here form a single clade. This is discussed in Chapters 5 and 6. Dermestidae. New data on the phylogeny of Dermestidae have been published by Lawrence & Slipiiiski (2005) and Kiselyova & McHugh (2006). In the former work, Dermestinae and Trinodinae were redefined to include Marioutinae and Thylodriinae, respectively. Ptinidae. The priority of Ptinidae Latreille 1802 over Anobiidae Fleming 1821 was pointed out by Lawrence & Newton (1995); the family name has been changed in this volume. Lymexylinae. Atractocerinae is recognized a subfamily separate from Lymexylinae and containing several genera following Paulus (2004). Trogossitidae. A new family-group classification involving only two subfamilies was proposed by Kolibäc (2005,2006) and is followed in Chapter 9.2. Thanerocleridae. The clerid subfamily Thaneroclerinae was elevated to family rank by Kolibäc (1992,1998) and a new subfamily Zenodosinae was proposed. Metaxinidae. The genus Metaxina Broun, tentatively included in Thaneroclerinae by Crowson (1964) was made the type of this new family by Kolibäc (2004). Mauroniscidae. This family was proposed by Majer (1994 b, 1995) and recognized in Lawrence etal. (1999).

2

John F. Lawrence, Rolf G. Beutel, Richard Α. Β. Leschen and Adam Slipinski

Melyridae. The subfamily Gietellinae, recognized by Lawrence & Newton (1995), was reduced to tribal level within Dasytinae. Cucuj oidea. A new arrangement of cucuj oid families was proposed by Leschen etal. (2005). These authors also noted that Cucujoidea was probably paraphyletic if the cleroids are excluded, and similar statements have been made in other recent works (Beutel & Pollock 2000; Hunt etal. 2007). Cryptophagidae. The subfamily Hypocoprinae was considered to be a tribe within Atomariinae by Leschen (1996). Phloeostichidae. The subfamilies Agapythinae Sen Gupta & Crowson, Myraboliinae Lawrence & Britton, Priasilphinae Crowson and Tasmosalpinginae Lawrence & Britton were all raised to family rank by Leschen et al. (2005), leaving only Phloeostichinae and Hymaeinae within this family. See Chapters 10.11, 10.12, 10.13, 10.16 and 10.23. Nitidulidae. Three additional subfamilies were proposed by Kirejtshuk: Amphicrossinae (1986), Epuraeinae (1986) and Maynipeplinae (1998). Endomychidae. A reanalysis of the subfamilies of Endomychidae (Tomaszewska 2005) confirmed the classification presented in 38.1. Coccinellidae. A new classification involving only two subfamilies, with almost all currently recognized subfamilies considered to be tribes within Coccinellinae, was proposed by Slipiriski (2007) and is used in Chapter 10.33. Corylophidae. A new subfamily and tribal classification was proposed by Slipinski et al. (2009). Only two subfamilies, Periptyctinae and Corylophinae are recognized, with all other supergeneric groups included within the latter. Tetratomidae. The melandryid subfamilies Hallomeninae and Eustrophinae were transferred to this family by Nikitsky (1998,2004 a, b). Melandryidae. A new classification based in part on Nikitsky (2002) is included in Chapter 11.6. Ripiphoridae. A new classification with the subfamilies Pelecotominae and Ptilophorinae redefined was proposed by Falin (2004). See Chapter 11.8. Zopheridae. In the classification proposed by Slipinski & Lawrence (1999) only two subfamilies are recognized, with both Usechinae and Monommatinae being downgraded to tribes within Zopherinae.

Promecheilidae. With the addition of the genus Promecheilus Solier to this family, the name Perimylopidae St. George (1939) has been replaced by the older name Promecheilidae Lacordaire (1959). Tenebrionidae. The major changes at the subfamily level in in Tenebrionidae are the recognition of the Nilioninae (including only Nilio Latreille) and the use of Stenochiinae as a senior synonym of Coelometopinae (Bouchard et al. 2005). See Chapter 11.14. Oedemeridae. A new subfamily Polypriinae was proposed by Lawrence (2005 b) to include Polypria Chevrolat (Tenebrionoidea incertae sedis in Lawrence & Newton 1995) and Dasytomima Lawrence. Meloidae. The subfamily Tetraonycinae Boving & Craighead (1931) was added by Bologna & Pinto (2001). Mycteridae. As pointed out by Lawrence & Newton (1995), Eurypinae Thomson has priority over Lacconotinae and is used in Chapter 11.20. Anthicidae. The subfamilies Afreminae, Lagrioidinae and Ischaliinae were removed from Anthicidae and included in the next Chapter. With their removal a much stronger case can be made for the monophyly of this family. Ischaliidae was recognized at the family level by Nikitsky & Egorov (1992) and Nikitsky (1996), and doubts have been expressed about the inclusion of Afremus Levey and Lagrioida Fairmaire & Germain in Anthicidae (Lawrence & Britton 1991; Costa etal. 1995; Werner & Chandler 1995; Chandler 2002). Tenebrionoidea incertae sedis. Chapter 11.29 includes three family-group taxa listed in the previous paragraph plus the genera Rhizonium Sharp and Aprostomis Grouvelle.

Literature Beutel, R. G. & Pollock, D. A. (2000): Larval head morphology of Phycosecis litoralis (Pascoe) (Coleoptera: Phycosecidae) with phylogenetic implications. -Invertebrate Taxonomy 14:825-835. Bocak, L. & Bocakova, M. (2008): Phylogeny and classification of the family Lycidae (Insecta): Coleoptera. - Annales Zoologici Warszawa 58 (4): 695-720. Bocakova, M., Bocak, L., Hunt, T. & Vogler, A. P. (2007): Molecular phylogenetics of Elateriformia (Coleoptera): evolution of bioluminescence and neoteny. - Cladistics 23:477-496. Boving, A. G. & Craighead, F. C. (1931): An illustrated synopsis of the principal larval forms of the order Coleoptera. - Εntomologica Americana (N.S.) 11(1930): 1-351. Bologna, M. A. & Pinto, J. D. (2001): Phylogenetic studies of Meloidae (Coleoptera), with emphasis on the evolution of phoresy. - SystematicEntomology 26:33-72.

Changes in classification and list of families and subfamilies

Bouchard, P., Lawrence, J. F., Davies, A. & Newton, A. F. (2005): Synoptic classification of the world Tenebrionidae (Insecta: Coleoptera) with a review of family-group names. - Annales Zoologici, Warszawa 55 (4): 499-530. Branham, M. & Wenzel, J. W. (2001): The evolution of bioluminescence in cantharoids (Coleoptera: Elateroidea). - Florida Entomologist 84:565-586. - (2003): The origin of photic behavior and the evolution of sexual communication in fireflies (Coleoptera: Lampyridae). - Cladistics 19:1-22. Chandler,D. S. (2002): 117. Anthicidae Latreille 1819. Pp. 549-558 in Arnett, R. H., Jr., Thomas, M. C., Skelley, P. E. & Frank, J. H. (eds.) American Beetles. Volume

2. polyphaga:

Scarabaeoidea

through

Curcu-

lionoidea. CRC Press, Gainesville, Florida. Costa, C. (1975): Systematics and evolution of the tribes Pyrophorini and Heligmini with description of Campyloxeninae, new subfamily (Coleoptera, Elateridae). -Arquivos deZoologia 26 (2): 49-191. Costa, C., Casari-Chen, S. A. & Vanin, S. A. (1992): On the larvae of Tetralobini (Coleoptera, Elateridae). - Revista Brasileira de Entomologia

3 6 (4): 8 7 9 - 8 8 8 .

(1994): Cladistic Analysis and Systematics of the Tetralobini sensu Stibick, 1979 (Coleoptera, Elateridae, Pyrophorinae). - Arquivos de Zoologia 32 (3): 111-157. Costa, C., Vanin, S. A. & Ide, S. (1995): Larvae of Neotropical Coleoptera XXII. Description of adults and immatures of Lagrioida nortoni sp. n., and bionomics (Coleoptera, Tenebrionoidea,

3 Kirejtshuk, A. G. (1986): Analysis of structure of genitalia for reconstruction of phylogeny and substantiation of the system of the family of sap beetles (Nitidulidae, Coleoptera). - Trudy Vsesoyuznogo Entomologicheskogo Obtchestva (Proceedings of the All-Union Entomological Society) 6 8 : 2 2 - 2 8

(in Russian). (1998): The position of the subfamily Maynipeplinae subfam. n. (Coleoptera, Nitidulidae) in the classification and notes on the evolution and structural modifications among sap beetles. Entomologicheskoye Obozreniye 77: 540-554 (in Russian, translation in Entomological Review 78:793-807). Kiselyova, T. & McHugh, J. (2006): A phylogenetic study of Dermestidae (Coleoptera) based on larval morphology. -Systematic Entomology 31 (3): 469-507. Kolibäc, J. (1992): Revision of Thanerocleridae n. stat. (Coleoptera, Cleroidea). - Mitteilungen der Schweize-

-

rischen Entomologischen

-

-

-

A n t h i c i d a e ) . - Iheringia,

-

-

Porto Alegre, Serie Zoologia

78:113-126. Crowson, R. A. (1964): A review of the classification of Cleroidea (Coleoptera), with descriptions of two genera of Peltidae and of several new larval types. - Transactions

-

of the Royal Entomological

Society of

London 116:275-327. (1972): A review of the classification of Cantharoidea (Coleoptera), with the definition of two new families, Cneoglossidae and Omethidae. - Revista de la Universidad de Madrid 2 1 (82): 3 5 - 7 7 .

Dolin, V. G. (2000): The role of larval and wing venation characters in the systematics of Elateroidea (Coleoptera). Meetings in Memory kovsky. Lecture at the 52nd Annual

of N. A. CholodMeeting. 1 Parti

1999. St. Petersburg (in Russain). 50 pp. Rossiiskaya Akademiya Nauk & Russoye Entomologicheskoye Obshchestvo, St. Petersburg. F a l i n , Z . ( 2 0 0 4 ) : Phylogenetic analysis and revision of the genera and subfamilies of the Ripiphoridae (Coleop-

tera). PhD Thesis, University of Kansas. 560 pp. University Microfilms International, Ann Arbor, Michigan. Hunt, T., Bergsten, J., Levkanicova, Z., Papadopoulou, Α., St. John, O., Wild, R., Hammond, P.M., Ahrens, D., Balke, M., Caterino, M. S., Gömez-Zurita, J., Ribera, I., Barraclough, T. G., Bocakova, M., Bocak, L. & Vogler, A. P. (2007): A comprehensive phylogeny of beetles reveals the evolutionary origins of a superradiation. - Science 318(5858):1913-1916. Jeng, M.-L., Yang, P.-S. & Satö, M. (1998): The genus Cyphonocerus (Coleoptera, Lampyridae) from Taiwan and Japan, with notes on the subfamily Cyphonocerinae. -Elytra, Tokyo 26 (2): 379-398.

Gesellschaft

mologica Basiliensia et Collections

-

65:303-340.

(1997): Classification of the subfamilies of Cleridae (Coleoptera: Cleroidea). -ActaMuseiMoraviae, ScientiaeBiologicae, Brno 81 (1996): 307-361. (1998): New Australian Thanerocleridae, with notes on the biogeography of the subtribe Isoclerina Kolibäc (Coleoptera: Cleroidea). - Invertebrate Taxonomy 12:951-975. (2004): Metaxinidae fam. nov., a new family of Cleroidea (Coleoptera). - Entomologica Basiliensia 26: 239-268. (2005): A review of the Trogositidae. Part 1: Morphology of the genera (Coleoptera, Cleroidea). - EntoFrey 2 7 : 3 9 - 1 5 9 .

(2006): A review of the Trogositidae. Part 2: Larval morphology, phylogeny and taxonomy (Coleopt e r a , Cleroidea). - Entomologica Basiliensia tions Frey 2 8 : 1 0 5 - 1 5 3 .

et Collec-

Lawrence, J. F. (2005 a): Brevipogon, a new genus of North American Artematopodidae (Coleoptera). - The ColeopteristsBulletin 59 (2): 223-236. - (2005 b): Dasytomima, a new genus of Australian Oedemeridae and its relationship to Polypria Chevrolat (Coleoptera: Tenebrionoidea). - Annales Zoologici, Warszawa 5 5 (4): 6 6 3 - 6 7 6 .

Lawrence, J. F. & Britton, Ε. B. (1991): Coleoptera (Beetles). Pp. 543-683 in CSIRO Division of E n t o m o l o g y (ed.) Insects of Australia: a Textbook for Students and Research Workers, Second Edition. Vol. 2.

Melbourne University Press, Carlton, Victoria. Lawrence, J. F. & Newton, A. F., Jr. (1995): Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names). Pp. 779-1006 in Pakaluk, J. & Slipinski, S. A. (eds.) Biology, Phylogeny, and Classification Coleoptera: Papers Celebrating the 80th Birthday

of of

Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warsaw. Lawrence, J. F. & Slipinski, S. A. (2005): Three new genera of Indo-Australian Dermestidae (Coleoptera) and their phylogenetic significance. - Invertebrate Systematics 19:231-261. Lawrence, J. F., Hastings, A. M., Dallwitz, Μ. J., Paine, T. A. & Zürcher, Ε. J. (1999): "Beetles of the World: A Key and Information

System for Families

and

Sub-

families." CD-ROM, Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne.

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Lawrence, J. F., Muona, J., Teräväinen, Μ., Stahls, G. & Vahtera, V. (2007): Anischia, perothops and the phylogeny of Elateroidea. - Insect Systematics and Evolution 38: 2 0 5 - 2 3 9 . {Apparently Jyrki Muona hates commas) Leschen, R. A. B. (1996): Phylogeny and revision of the genera of Cryptophagidae (Coleoptera: Cucujoidea). - The University of Kansas Science Bulletin 55 (15): 5 4 9 - 6 3 4 . Leschen, R. A. B., Lawrence, J. F. & Slipiriski, S. A. (2005): Classification of basal Cucujoidea (Coleoptera: Polyphaga): cladistic analysis, keys and review of new families. - Invertebrate Systematics 1 9 : 1 7 - 7 3 . Levey, B. (1985): Afreminae: a new subfamily of Anthicidae (Coleoptera) from southern Africa. -EntomologicaScandinavica 15 (1984): 4 1 9 - 4 2 2 . Majer, K. (1994): A review of the classification of the Melyridae and related families (Coleoptera, Cleroidea).-EntomologicaBasiliensia 1 7 : 3 1 9 - 3 9 0 . - (1995): Revision of the family Mauroniscidae (Insecta: Coleoptera: Cleroidea). - Entomologische Abhandlungen aud dem Staatliches Museum für Tierkunde in Dresden 57: 5 7 - 8 9 . Nikitsky, Ν. Β. (1996): Fam. Ischaliidae - False Firered Beetles. Pp. 4 2 7 - 4 2 8 in Ler, P. A. (ed.) Keys to the Insecta of the Ear Eastern U.S.S.R. in Six Volumes. Vol. 3. Coleoptera or Beetles, Part 3. Dal'nauka, Vladivostok (in Russian). - (1998): Generic Classification of the Beetle Family Tetratomidae (Coleoptera, Tenebrionoidea) of the World, with Description of New Taxa. 80 pp. Pensoft (Series Faunistica No. 9), Sofia, Moscow. - (2002): The larvae of the false darkling beetles (Coleoptera, Melandryidae) of Russia, with taxonomic notes. - Byulletin Moskovskogo Obshchestva Ispytateleiprirody, Moscow 107 (3): 8 - 3 0 (in Russian). - (2004 a): The beetles of the subfamily Tetratominae Billberg, 1820 (Coleoptera, Tetratomidae) of the world fauna. - Byulleten Moskovskogo Obshestva Ispytatelei Prirody, Otdel Biologicheskii 109 (2): 2 5 - 3 6 (in Russian). - (2004 b): The beetles of the subfamily Piseninae Miyatake, 1960 (Coleoptera, Tetratomidae) of the world fauna. - Byulleten Moskovskogo Obshestva IspytateleiPrirody, Otdel Biologicheskii 109 (4): 3 - 7 (in Russian). Nikitsky, Ν. B. & Egorov, A. B. (1992): Fam. Ischaliidae, stat. n. - false fire-red beetles. Pp. 4 9 7 - 4 9 8 in Ler, P. A. (ed.) Keys to the Insecta of the Far Eastern U.S.S.R. in Six Volumes. Vol. 3. Coleoptera or Beetles, Part 2. Nauka, St. Petersburg (in Russian). Paulus, Η. F. (1972): Die systematische und phylogenetische Stellung der Karumiidae, mit einer Beschreibung von Escalerina serraticornis n. sp. aus S.-Persien (Ins.: Coleoptera: Cantharoidea). - SenkkenbergianaBiologica 53 (1/2): 3 7 - 5 4 . - (2004): Urteagraeca nov. gen. et nov. spec., der erste Vertreter der tropischen Atractocerinae in Europa, sowie eine Beschreibung von Homaloxylon aspoecki nov. spec, aus Yunnan (China) (Coleoptera: Cucujiformia: Lymexylidae, Atractocerinae nov. status). -Denisia 1 3 : 2 7 7 - 2 9 0 . Slipiriski, S. A. (2007): Australian Ladybird Beetles (Coleoptera: Coccinellidae). their Biology and Classification. xviii + 286 pp. Australian Biological Resources

Study, Department of the Environment and Water Resources, Canberra. Slipiriski, S. A. & Lawrence, J. F. (1999): Phylogeny and classification of Zopheridae sensu novo (Coleoptera: Tenebrionoidea) with a review of the genera of Zopherinae (excluding Monommatini). - Annates Zoologici(Warszawa) 4 9 (1/2): 1 - 5 3 . Slipiriski, Α., Tomaszewska, W. & Lawrence, J. F. (2009): Phylogeny and classification of Corylophidae (Coleoptera: Cucujoidea) with descriptions of new genera and larvae. - Systematic Entomology (in press). Tomaszewska, K. W. (2005): Phylogeny and generic classification of the subfamily Lycoperdininae with a re-analysis of the family Endomychidae (Coleoptera: Cucujoidea). - Annates Zoologici, Warszawa 55, Supplement 1 : 1 - 1 7 2 .

List of Families and Subfamilies Adephaga

Meruidae Spangler & Steiner, 2 0 0 5 Aspidytidae Ribera etal. 2 0 0 2

Polyphaga Elateriformia Elateroidea Rhinorhipidae Lawrence, 1 9 8 8 Artematopodidae Lacordaire, 1 8 5 7 Electribiinae Crowson, 1 9 7 5 Allopogoniinae Crowson, 1 9 7 3 Artematopodinae Lacordaire, 1 8 5 7 Brachypsectridae LeConte & Horn, 1 8 8 3 Cerophytidae Latreille, 1 8 3 4 Eucnemidae Eschscholtz, 1 8 2 9 Perothopinae Lacordaire, 1 8 5 7 Phyllocerinae Reitter, 1905 Pseudomeninae Muona, 1933 Palaeoxeninae Muona, 1 9 9 3 Phlegoninae Muona, 1 9 9 3 Anischiinae Fleutiaux, 1 9 3 6 Melasinae Fleming, 1 8 2 1 Eucneminae Eschscholtz, 1 8 2 9 Macraulacinae Fleutiaux, 1 9 2 3 Throscidae Laporte, 1 8 4 0 Elateridae Leach, 1815 Cebrioninae Latreille, 1 8 0 2 Agrypninae Candeze, 1 8 5 7 (incl. Tetralobinae) Thylacosterninae Fleutiaux, 1 9 2 0 Lissominae Laporte, 1835 Semiotinae Jacobson, 1 9 1 3 Campyloxeninae Costa, 1975 Pityobiinae Hyslop, 1 9 1 7 Oxynopterinae Candeze, 1 8 5 7 Denticollinae Stein & Weise, 1 8 7 7 (1856) Negastriinae Nakane & Kishii, 1 9 5 6 Elaterinae Leach, 1 8 1 5 Cardiophorinae Candeze, 1 8 6 0 Hemiopinae Fleutiaux, 1 9 4 1 Physodactylinae Lacordaire, 1 8 5 7 Eudicronychinae Girard, 1 9 7 1

5

Changes in classification and list of families and subfamilies

Subprotelaterinae Fleutiaux, 1936 Morostominae Dolin, 2000 Plastoceridae Crowson, 1972 Drilidae Blanchard, 1845 Omalisidae Lacordaire, 1857 Lycidae Laporte, 1836 Libnetinae Bocak & Bocakova, 1990 Dictyopterinae Kleine, 1928 Lypropaeinae Bocak & Bocakova, 1989 Ateliinae Kleine, 1928 Lycinae Laporte, 1836 Dexorinae Bocak & Bocakova, 1989 Telegeusidae Leng, 1920 Phengodidae Leconte, 1861 Phengodinae LeConte, 1861 Mastinocerinae LeConte, 1881 Penicillophorinae Paulus, 1974 Rhagophthalmidae Olivier, 1907 Lampyridae Latreille, 1817 Psilocladinae McDermott, 1964 Lampyrinae Latreille, 1817 Luciolinae Lacordaire, 1857 Photurinae Lacordaire, 1857 Omethidae Leconte, 1861 Omethinae LeConte, 1861 Matheteinae LeConte, 1881 Driloniinae Crowson, 1972 Cantharidae Imhoff, 1856 (1815) Cantharinae Imhoff, 1856 (1815) Silinae Mulsant, 1862 Dysmorphocerinae Brancucci, 1980 Malthininae Kiesenwetter, 1852 Chauliognathinae Leconte, 1861

Elateriformia Incertae Sedis

Podabrocephalidae Pic, 1930 Neocrowsonia Kistner & Abdel-Galil, 1986 Cydistinae Paulus, 1972 Pterotinae LeConte, 1861 Harmatelia Walker, 1858 Stenocladius Fairmaire, 1878 Ototretinae McDermott, 1964

Derodontiformia Derodontoidea

Derodontidae LeConte, 1861 Peltasticinae Leconte, 1861 Derodontinae LeConte, 1861 Laricobiinae Mulsant & Rey, 1 8 6 3 - 4 Nosodendridae Erichson, 1846 Jacobsoniidae Heller, 1926

Bostrichiformia Bostrichoidea

Dermestidae Latreille, 1804 Dermestinae Latreille, 1804 (incl. Marioutinae) Thorictinae Agassiz, 1846 Orphilinae LeConte, 1861 Trinodinae Casey, 1900 (incl. Thylodriinae) Attageninae Laporte, 1840 Megatominae Leach, 1815 Endecatomidae Leconte, 1861

Bostrichidae Latreille, 1802 Dysidinae Lesne, 1921 Polycaoninae Lesne, 1936 Bostrichinae Latreille, 1802 Psoinae Blanchard, 1851 Dinoderinae C. G. Thomson, 1863 Lyctinae Billberg, 1820 Euderiinae Lesne, 1934 Ptinidae Latreille, 1802 Eucradinae LeConte, 1861 Ptininae Latreille, 1802 Dryophilinae LeConte, 1861 Ernobiinae Pic, 1912 Anobiinae Fleming, 1821 Ptilininae Shuckard, 1840 Alvarenganiellinae Viana & Martinez, 1971 Xyletininae Gistel, 1956 Dorcatominae Thomson, 1859 Mesocoelopodinae Mulsant & Rey, 1864

Cucujiformia Lymexyloidea

Lymexylidae Fleming, 1821 Hylecoetinae Gistel, 1856 Lymexylinae Fleming, 1821 Atractocerinae Laporte, 1840 Melittommatinae Wheeler, 1986

Cleroidea

Phloiophilidae Kiesenwetter, 1863 Trogossitidae Latreille, 1802 Peltinae Kirby, 1837 Trogossitinae Latreille, 1802 Chaetosomatidae Crowson, 1952 Metaxinidae Kolibäc, 2004 (from Cleridae) Thanerocleridae Chapin, 1924 (from Cleridae) Zenodosinae Kolibäc, 1992 Thaneroclerinae Chapin, 1924 Cleridae Latreille, 1802 Tillinae Leach, 1815 Hydnocerinae Spinola, 1844 Clerinae Latreille, 1802 Korynetinae Laporte, 1836 (incl. Epiphloeinae, Enopliinae, Tarsosteninae) Acanthocnemidae Crowson, 1964 Phycosecidae Crowson, 1952 Prionoceridae Lacordaire, 1857 Mauroniscidae Majer, 1994 Melyridae Leach, 1815 Rhadalinae LeConte, 1861 (incl. Aplocneminae) Melyrinae Leach, 1815 Dasytinae Laporte, 1840 (incl. Gietellinae) Malachiinae Fleming, 1821 (incl. Attalomiminae)

Cucujoidea (arrangement is new)

Boganiidae Sen Gupta & Crowson, 1966 Paracucujinae Endrödy-Younga & Crowson, 1986 Boganiinae Sen Gupta & Crowson, 1966 Byturidaejacquelin DuVal, 1858 Platydascillinae Pic, 1914 Byturinaejacquelin DuVal, 1858

6

John F. Lawrence, Rolf G. Beutel, Richard Α. Β. Leschen and Adam Slipinski

Helotidae Reitter, 1876/Chapuis, 1876 ProtocucujidaeCrowson, 1954 Sphindidaejacquelin DuVal, 1860 Protosphindinae Sen Gupta & Crowson, 1979 Odontosphindinae Sen Gupta & Crowson, 1979 Sphindiphorinae McHugh, 1993 Sphindinaejacquelin DuVal, 1860 Biphyllidae LeConte, 1861 Erotylidae Latreille, 1802 Xenoscelinae Ganglbauer, 1899 Pharaxonothinae Crowson, 1952 (incl. Setariolinae) Loberinae Bruce, 1951 Languriinae Crotch, 1873 Cryptophilinae Casey, 1900 (incl. Toraminae) Erotylinae Latreille, 1802 (incl. Tritominae, Dacninae, Megalodacninae, Encaustinae) Monotomidae Laporte, 1840 Rhizophaginae Redtenbacher, 1845 Monotominae Laporte, 1840 Hobartiidae Sen Gupta & Crowson, 1966 Cryptophagidae Kirby, 1837 Cryptophaginae Kirby, 1837 Atomariinae LeConte, 1861 (incl. Hypocoprinae, Alfieriellinae) Agapythidae Sen Gupta & Crowson, 1969 Priasilphidae Crowson, 1973 Phloeostichidae Reitter, 1911 Silvanidae Kirby, 1837 Brontinae Erichson, 1845/Blanchard, 1845 Silvaninae Kirby, 1837 Cucujidae Latreille, 1802 Myraboliidae Lawrence & Britton, 1991 Cavognathidae Sen Gupta & Crowson, 1966 Lamingtoniidae Sen Gupta & Crowson, 1966 Passandridae Erichson 1845/Blanchard, 1845 Phalacridae Leach, 1815 Phaenocephalinae Matthews, 1899 Phalacrinae Leach, 1815 Propalticidae Crowson, 1952 Laemophloeidae Ganglbauer, 1899 Tasmosalpingidae Lawrence & Britton, 1991 Cyclaxyridae Gimmel etal, 2009 Kateretidae Erichson, 1846 Nitidulidae Latreille, 1802 CalonecrinaeKirejtshuk, 1982 MaynipeplinaeKirejtshuk, 1998 Epuraeinae Erichson, 1843 Carpophilinae Erichson, 1842 AmphicrossinaeKirejtshuk, 1986 Meligethinae C. G. Thomson, 1859 Nitidulinae Latreille, 1802 Cillaeinae Kirejtshuk & Audisio in Kirejtshuk, 1986 Cryptarchinae C. G. Thomson, 1859 Cybocephalinae Jacquelin Du Val, 1858 Smicripidae Horn, 1879 Bothrideridae Erichson, 1845 Teredinae Seidlitz, 1888 Xylariophilinae Pal & Lawrence, 1986 Anommatinae Ganglbauer, 1899 Bothriderinae Erichson, 1845

Cerylonidae Bilberg, 1820 Euxestinae Grouvelle, 1908 Loeblioryloninae Slipinski, 1990 Ostomopsinae Sen Gupta & Crowson, 1973 Murmidiinae Jacquelin DuVal, 1858 Ceryloninae Bilberg, 1820 Alexiidae Imhoff, 1856 Discolomatidae Horn, 1878 Notiophyginae Jacobson, 1915 Discolomatinae Horn, 1878 Aphanocephalinae Grouvelle, 1912 Cephalophaninaejohn, 1954 Pondonatinaejohn, 1954 Endomychidae Leach, 1815 Merophysiinae Seidlitz, 1872 Pleganophorinae Jacquelin Du Val, 1885 Anamorphinae Strohecker, 1953 Leiestinae C. G. Thomson, 1863 Mycetaeinae Jacquelin DuVal, 1857 Eupsilobiinae Casey, 1895 Xenomycetinae Strohecker in Arnett, 1962 Danascelinae Tomaszewska, 2000 Endomychinae Leach, 1815 Epipocinae Gorham, 1873 Stenotarsinae Chapuis, 1876 Lycoperdininae Redtenbacher, 1844 Coccinellidae Latreille, 1807 Microweiseinae Leng, 1920 (= Sticholotidinae part) Coccinellinae Latreille, 1807 (incl. Sticholotidinae part, Chilocorinae, Coccidulinae, Scymninae, Ortaliinae, Epilachninae) Corylophidae Leconte, 1852 Periptyctinae Slipinski etal, 2001 Corylophinae LeConte, 1852 Latridiidae Erichson, 1842 Latridiinae Erichson, 1852 Corticariinae Curtis, 1829

Tenebrionoidea Mycetophagidae Leach, 1815 Esarcinae Reitter, 1882 Mycetophaginae Leach, 1815 Bergininae Leng, 1920 Archeocrypticidae Kaszab, 1964 Pterogeniidae Crowson, 1953 Ciidae Leach in Samouelle, 1819 Sphindociinae Lawrence, 1974 Ciinae Leach in Samouelle, 1819 Tetratomidae Billberg, 1820 Tetratominae Billberg, 1820 Piseninae Miyatake, 1960 Penthinae Lacordaire, 1859 Hallomeninae Mulsant 1856/Gistel, 1856 (from Melandryidae) Eustrophinae Gistel, 1856 (from Melandryidae) Melandryidae Leach, 1815 Melandryinae Leach, 1815 Osphyinae Mulsant, 1856 (1840)

7

Changes in classification and list of families and subfamilies

Mordellidae Latreille, 1802 Ctenidiinae Franciscolo, 1951 Mordellinae Latreille, 1802 Ripiphoridae Gemminger & Harold, 1870(1853) Ptilophorinae Gerstaecker, 1855 Pelecotominae Seidlitz, 1875 (incl. Micholaeminae) Hemirhipidiinae Heller, 1921 Ripidiinae Gerstaecker, 1855 Ripiphorinae Gemminger & Harold, 1870(1853) Zopheridae Solier, 1834 Colydiinae Erichson, 1842 Zopherinae Solier, 1834 (incl. Monommatinae, Usechinae, Pycnomerinae) Ulodidae Pascoe, 1869 Promecheilidae Lacordaire, 1859 (= Perimylopidae) Chalcodryidae Watt, 1974 Trachelostenidae Lacordaire, 1859 Tenebrionidae Latreille, 1802 Lagriinae Latreille, 1825 (1820) Nilioninae Lacordaire, 1859 Cossyphodinae Wasmann, 1899 Phrenapatinae Solier, 1834 Zolodininae Watt, 1974 Pimeliinae Latreille, 1802 Tenebrioninae Latreille, 1802 Alleculinae Laporte, 1840 Diaperinae Latreille, 1802 Stenochiinae Kirby, 1837 (= Coelometopinae) Prostomidae C. G. Thomson, 1859 Synchroidae Lacordaire, 1859 Stenotrachelidae C. G. Thomson, 1859 Stenotrachelinae C. G. Thomson, 1859 Cephaloinae LeConte, 1862 Nematoplinae Leconte, 1862 Stoliinae Nikitsky, 1985 Oedemeridae Latreille, 1810 Polypriinae Lawrence, 2005 Calopodinae Costa, 1852 Oedemerinae Latreille, 1810 (incl. Nacerdinae) Meloidae Gyllenhal, 1810 EleticinaeWellman, 1910 Meloinae Gyllenhal, 1810

Tetraonycinae Boving & Craighead, 1931 Nemognathinae Laporte, 1840 Mycteridae Blanchard, 1845 Mycterinae Blanchard, 1845 Eurypinae J. Thomson, 1860 (= Lacconotinae) Hemipeplinae Lacordaire, 1854 Boridae C. G. Thomson, 1859 Borinae C. G. Thomson, 1859 Synercticinae Lawrence & Pollock, 1994 Trictenotomidae Blanchard, 1845 Pythidae Solier, 1834 Pyrochroidae Latreille, 1807 Tydessinae Nikitsky, 1986 Pilipalpinae Abdullah, 1964 Pedilinae Lacordaire, 1859 Pyrochroinae Latreille, 1807 Agnathinae Lacordaire, 1859 Salpingidae Leach, 1815 Othniinae LeConte, 1861 Prostominiinae Grouvelle, 1914 Agleninae Horn, 1878 Inopeplinae Grouvelle, 1908 Dacoderinae LeConte, 1862 Aegialitinae LeConte, 1862 Salpinginae Leach, 1815 An thicidae Latreille, 1819 Eurygeniinae LeConte, 1862 Macratriinae LeConte, 1862 Steropinae Jacquelin DuVal, 1863 Copobaeninae Abdullah, 1969 Lemodinae Lawrence & Britton, 1991 Tomoderinae Bonadona, 1961 Anthicinae Latreille, 1819 Aderidae Winkler, 1927 Scraptiidae Mulsant 1856/Gistel, 1856 Scraptiinae Mulsant, 1856/Gistel, 1856 Anaspidinae Mulsant, 1856 Tenebrionoidea Incertae Sedis Lagrioidinae Abdullah & Abdullah, 1968 (from Anthicidae) Afreminae Levey, 1985 (from Anthicidae) Ischaliinae Blair, 1920 (from Anthicidae) Rhizonium Sharp, 1876 Aprostomis Grouvelle, 1912

2. Glossary of Morphological Terms John F. Lawrence, Rolf G. Beutel, Richard A. B. Leschen and Adam Slipmski

Although a brief overview of beetle morphology may be found in Crowson (1955, 1981), Lawrence (1991), Lawrence & Britton (1991), Lawrence et al. (1999 a, b) and Beutel & Lawrence (2005), the following set of definitions and discussions represents a preliminary attempt to standardize nomenclature within the order, especially with respect to terminology introduced in relatively recent publications and not included in general textbooks or dictionaries. This is not meant to be a complete glossary, and a detailed treatment of beetle morphology will be included in a later volume of this series. The terms discussed below are, in general, those which turn up regularly in adult and larval descriptions of the beetle families included in this volume, and commonly used words found in general insect morphology texts are included only when their use within the order Coleoptera requires some explanation or elaboration. Terms or topics are arranged alphabetically under each of eight subheadings: head, prothorax, pterothorax, hind wing and abdomen for adults and head, thorax and abdomen for larvae.

Adult Head Antennae: Clavate. Antennomeres gradually broader towards the apex; synonymous with incrassate.

often arched and occasionally bearing an anteriorly projecting median process (Stickney 1923). Corporotentorium (see Corpotentorium). Epistomal Suture (or Sulcus) (see Frontoclypeal Suture). Frontoclypeal Suture. Transverse suture representing an invagination between the frons and clypeus to form the epistomal ridge beneath with the anterior tentorial pits at either end. Laminatentoria. Mesal expansions of the anterior tentorial arms, which may meet at the midline to form a secondary bridge anterior to the corpotentorium, if present (Stickney 1923). Median Occipital Endocarina. Internal longitudinal ridge extending anteriorly from the dorsal edge of the occipital foramen. Subantennal Groove. Groove or concavity lying below the antennal insertion and housing the base of the antenna. Placed between the eye (if present) and the mandibular articulation, and sometimes extends below or behind the eye.

Antennae: Capitate. One or more apical antennomeres are abruptly broader than and/or distinctly longer than those preceding them.

Transverse Occipital Ridge. Any transverse line, carina or elevated ridge extending across the occipital region between the eyes and the occipital foramen. Usually indicates the extent to which the head is retracted into the prothorax; an elevated ridge may abut the anterior edge of the pronotum.

Antennal Fossa. Saucer-like concavity surrounding the countersunk antennal insertion (not to be confused with the antennal insertion or socket).

Vertexal Line (see Transverse Occipital Ridge).

Antennal Insertion. Point of attachment of the antenna. Antennal insertions are considered to be exposed when at least some portion of each antennal socket is visible from above the long axis of the head (without regard to head orientation with respect to the body). Cervical Sclerite. Compound sclerite in Polyphaga joining the ventrolateral edge of the occipital foramen with the lateral portion of the presternum on each side. Each sclerite is usually divided into an anterior and posterior section separated by membrane. Corpotentorium. Transverse bridge (tentorial bridge) connecting the posterior tentorial arms,

Adult Prothorax Cryptopleuron (see Endopleuron). Endopleuron. Portion of the propleuron beneath the pronotum; not visible externally. In Archostemata, Adephaga and some Myxophaga this is a relatively minor part of the pleuron, but in almost all Polyphaga the pleuron is entirely internalized. Hypomeron. Ventral portion of pronotum below the lateral pronotal carinae when these are present. In many Polyphaga, the two hypomera extend mesally behind the coxae partly or completely closing the procoxal cavities externally (see Postcoxal Process).

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John F. Lawrence, Rolf G. Beutel, Richard Α. B. Leschen and A d a m Slipiriski

A

1. 2. 3. 4. 5. 6.

Mesoventrite Mesoventral Process Procoxal Rest Mesoventral Cavity Mesothoracic Discrimen Mesanepisternum

7. 8. 9. 10. 11. 12.

Mesepimeron Mesocoxa Mesocoxal Cavity Mesotrochantin Metanepisternum Metaventrite

13. 14. 15. 16. 17. 18.

Metaventral Process Metathoracic Discrimen Metakatepisternal Suture Metepimeron Metacoxa Metatrochantin

Fig. 2.1. Coleopteran pterothoraces, ventral: A, Distocupes varians (Lea) (Archostemata: Cupedidae); B, Episcaphula australis (Boisduval) (Polyphaga: Cucujoidea: Erotylidae); C, Sclerocyphon sp. (Polyphaga: Byrrhoidea: Psephenidae). (© CSIRO Australia).

Lateral P r o n o t a l Carina. Distinct edge separating t h e p r o n o t a l disc f r o m t h e p r o n o t a l h y p o m e r o n o n each side. Equivalent to t h e lateral edge of t h e prot h o r a x (but n o t t h e p r o n o t u m , w h i c h t e r m i n a t e s only at t h e n o t o p l e u r a l or notosternal suture). T h e lateral carina may be raised to f o r m a m a r g i n or bead, b u t t h a t is n o t always t h e case. P o s t c o x a l Process. Mesal e x t e n s i o n of t h e posterior p a r t of t h e p r o p l e u r o n (Archostemata a n d Adephaga) or h y p o m e r o n ( M y x o p h a g a a n d Polyphaga) b e h i n d t h e procoxa, w h i c h may m e e t t h e p r o s t e r n a l process or t h e o p p o s i n g postcoxsal process, t h u s closing t h e procoxal cavitiy externally. P r o n o t a l Disc. Dorsal p o r t i o n of t h e p r o n o t u m , lying above t h e lateral pronotal carinae w h e n these are present. P r o c o x a l Cavity. C o u n t e r s u n k prothoracic housi n g into which t h e procoxa fits. F o r m e d in part by t h e p r o s t e r n u m (but s e e Prosternum) a n d in part by t h e p r o p l e u r o n (Archostemata, Adephaga a n d Myxophaga) or pronotal h y p o m e r o n (Polyphaga). This is an a u t a p o m o r p h y of t h e order Coleoptera b u t is secondarily reduced in a n u m b e r of softbodied beetles.

P r o c o x a l Cavities: E x t e r n a l Closure. Externally closed w h e n t h e postcoxal processes of t h e h y p o m e r a m e e t the prosternal process or m e e t one another. P r o c o x a l Cavities: I n t e r n a l Closure. Internally closed w h e n a n a r r o w or b r o a d b r i d g e connects a p o r t i o n of t h e internal l i n i n g of t h e cavity (usually in t h e vicinity of the sternal apophysis) w i t h t h e internal wall of t h e postcoxal process. Prosternal Process. Posterior projection of t h e mesal p o r t i o n of t h e p r o s t e r n u m w h i c h extends between t h e procoxae a n d may overlap t h e mesoventrite or fit i n t o t h e mesoventral cavity. T h e t e r m " s p i n a s t e r n u m " has b e e n used by those s t u d y i n g Curculionoidea for t h e posterior e n d of t h e prosternal process, w h e n t h e central p o r t i o n of t h a t process has been obliterated to a c c o m m o d a t e t h e enlarged a n d contiguous procoxae. P r o s t e r n u m . Used for t h e e n t i r e ventral plate lying in f r o n t of a n d b e t w e e n t h e procoxae a n d b e t w e e n t h e n o t o s t e r n a l or p l e u r o s t e r n a l s u t u r e s , a l t h o u g h it is likely t h a t this sclerite has a complex origin, like c o m p a r a b l e s t r u c t u r e s in t h e m e s o t h o r a x a n d m e t a t h o r a x (see M e s o v e n t r i t e a n d Metaventrite).

Glossary of morphological terms

11

Spinasternum (see Prosternal Process).

Mesosternal Cavity (see Mesoventral cavity).

Sublateral Pronotal Carina. Applied to various longitudinal carinae lying mesad of the lateral carinae. These may extend the length of the pronotal disc, as in Laemophloeidae, or be restricted to the posterior angles, as in many Elateridae.

Mesosternum (see Mesoventrite).

Adult Pterothorax Elytral Sutural Flange: Deflection. Deflected when the apical portion of the flange is expanded and thus visible when the elytra are in the closed position. Characteristic of certain families of Cucujoidea (Leschen et al. 2005) b u t may occur elsewhere. Lateral Closure of Mesocoxal Cavity. Laterally open when the lateral wall is formed partly by one or more pleural sclerites and laterally closed when the lateral wall is formed entirely by the meeting of the mesoventrite and metaventrite. Mesanepisternum. Anterior pleural sclerite of the mesothorax. Mesendosternite. A pair of internal apodemes formed by invaginations within the mesocoxal cavities and representing the original furca. Mesepimeron. Posterior pleural sclerite of the mesothorax. Mesocoxal Cavity. Countersunk pterothoracic housing into which the mesocoxa fits. Formed by portions of the mesoventrite and metaventrite, often with the addition of mesopleural sclerites and less commonly the metanepisternum. This autapomorphy of the order Coleoptera is secondarily reduced in a number of soft-bodied beetles. Mesofurca (see Mesendosternite). Mesometathoracic Joint. Joint between mesothorax and metathorax which is visible only within the mesocoxal cavities; membranous in Archostemata and Adephaga and some families of Polyphaga, b u t either consists of a solid joint or is completely absent in Myxophaga and most Polyphaga.

Mesothoracic Discrimen. Median line on the mesoventrite representing the invagination of the true mesosternum. This line is rarely complete and often absent in the mesothorax, where the single invagination or furca is replaced by well separated endosternal apodemes. Mesoventral Cavity. Cavity on the mesoventrite into which the prosternal process fits. Mesoventral Process. Mesal lobe at the posterior edge of the mesoventrite which usually extends between the mesocoxal cavities and meets the metaventral process (see Mesometaventral Junction). Mesoventrite. Applies to the ventral plate lying in front of and between the mesocoxal cavities; delimited laterally by the mesothoracic pleurosternal sutures. Although called the mesosternum in most earlier works on Coleoptera, this sclerite is equivalent to the paired mesothoracic preepisterna and paired mesolcatepisterna, the true mesosternum having been largely invaginated and represented only by the area in the immediately vicinity of the bases of the mesendosternites). The transverse suture separating the katepisterna from the preepisterna is never complete in the mesothorax of beetles (indicated by an internal transverse ridge in a few Archostemata) and the discrimen, representing the invagination of the original mesosternum, is present in most Archostemata, Gyrinidae, most Scirtoidea, most Buprestidae, a number of byrrhoid families and the elateroid family Artematopodidae. Metacoxal Cavity. Countersunk abdominal housing into which the metacoxa fits. Usually formed by abdominal sternites II and III combined with the posterior wall of the metaventrite. This autapomorphy of the order Coleoptera is secondarily reduced in a number of soft-bodied beetles. Metafurca (see Metendosternite).

Mesometaventral Joint (see Mesometaventral Junction).

Metakatepisternal Suture. Transverse suture on the metaventrite which separates the paired metathoracic preepisterna from the paired metakatepisterna. Although a complete suture, extending from the discrimen to the lateral edges of the mesoventrite on each side, is part of the groundplan of Coleoptera, it is often shortened so that it extends for only a short distance on either side of the discrimen.

Mesometaventral Junction. Meeting of the mesoventrite and metaventrite between the mesocoxal cavities. The junction may involve the abutment of the two sclerites, the overlapping of one by the other, or a complex, monocondylic or dicondylic joint. In some instances the two sclerites are fused together with or without a visible joint.

Metanepisternum. Anterior pleural sclerite of the metathorax, which in Coleoptera is laterad of the metaventrite and mesoventrad of the metepimeron. Because the lateral (dorsal) portion of the metanepisternum is often concealed beneath the elytral epipleura, its shape in descriptions is based on the visible portion only.

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John F. Lawrence, Rolf G. Beutel, Richard Α. Β. Leschen and Adam Slipiriski

Metathoracic Discrimen. Median line extending forward from the posterior edge of the metaventrite, internally corresponding with a more or less high median ridge representing the invagination of the true metasternum. The ridge is usually connected with the metendosternite. The discrimen is often relatively long and may extend anteriorly to or beyond the base of the metaventral process, but it is reduced or absent in a number of groups. Metaventral Process. Mesal lobe at the anterior end of the metaventrite which often extends forward between the mesocoxae and meets the mesoventral process (see Mesometaventral junction). Metaventrite. Ventral plate lying behind and between the mesocoxal cavities and delimited laterally by the metanepisterna. Although called the metasternum in most earlier works on Coleoptera, this sclerite is equivalent to the paired metapreepisterna and paired metakatepisterna, the true metasternum havingbeen invaginated along the midline. The transverse suture separating the katepisterna from the preepisterna is often complete, but may be shortened (extending for a short distance on either side of the discrimen) or absent, and the discrimen, representing the invagination of the original metasternum, is often very long, sometimes completely dividing the metaventrite into halves, but may be shortened or absent in various taxa. [Ferris 1940; Campau 1940; Matsuda 1960; Lawrence 1999; Lawrence etal. 1999 b; Beutel & Haas 2000.] Metendosternite. Complex internal apodeme equivalent to the metafurca. Arises at or near the posterior edge of the metaventrite usually at the junction of the discrimen and the metakatepisternal suture and projecting anterodorsally. It usually consists of a median stalk, two short to long lateral arms and an anterior process from which a pair of tendons arise; however a lamina may be associated with each of the lateral arms and a pair of anteroventral processes may arise from the point where the lateral arms meet the stalk. In some taxa the stalk may be short or absent and the anterior tendons often arise on the arms. [Crowson 1938,1944, 1955.] Metepimeron. Posterior pleural sclerite of the metathorax, which in Coleoptera is located laterad of and above the metanepisternum and mostly concealed by the elytral epimeron. In most beetle groups a small portion of this sclerite is visible near the lateral edge of the metacoxa. Prepectus. Anterior portion of mesoventrite and combined mesanepisterna, especially when set off from remainder of ventrite. Pretarsus. The terminal segment of the leg, consisting of a pair of claws in most adults or rarely a single claw.

Postcoxal Lines of Metaventrite. Slightly raised ridges or abrupt edges of impressions at the anterior edge of the metaventrite. Sometimes these lines delimit crural impressions, but this is not always the case. Procoxal Rests of Mesoventrite. Pair of impressions at the anterior edge of the mesoventrite into which the procoxae fit; these impressions may be almost horizontal in lateral view, but more often they are distinctly inclined or occasionally almost vertical. The procoxal rests sometimes extend laterally beyond the mesoventrite and onto the mesanepisterna. Prosternal Rest of Mesoventrite. Median elevation at the anterior edge of the mesoventrite which fits against the underside of the prosternal process; in those elateroid taxa utilizing the clicking maneuver this process blocks the prosternal process from sliding into the mesoventral cavity until sufficient pressure is built up to overcome the inertia. In some other taxa the process may be flattened and abut the underside of the prosternal process. Prothoracic Rest of Mesoventrite. Refers to a condition in which the entire anterior edge of the mesoventrite and flanking mesanepisterna form a peduncle over which the prothorax fits; this is usually correlated with a broad external closure of the procoxal cavities. Scutellar Shield. Exposed portion of the mesoscutellum which lies between the bases of the elytra. Scutellum. Posterior portion of mesotergum. Often referring only to that portion of the scutellum which is visible between the bases of the elytra (see Scutellar Shield). Transverse Metasternal Suture (see Metakatepisternal Suture). Transverse Metaventral Suture (see Metakatepisternal Suture). Trochantin. Precoxal remnant articulating with the coxa, sternum and pleuron. Trochantin (Prothorax). Used in Myxophaga and Polyphaga for the exposed portion of the trochantinopleuron. Trochantinopleuron. Fusion product of the prothoracic trochantin with the propleuron.

Hind Wing The terms for wing veins and related structures used in this volume are based primarily on that of Kukalovä-Peck & Lawrence (1993, 2004). Some changes in wing vein terminology listed on p. 144

Glossary of morphological terms

13

^ Medial Embayment

Fig. 2.2. Coleopteran hind wings: A, Omma stanleyi Newman (Archostemata: Ommatidae); B, Notodascillus sublineatus Carter (Polyphaga: Dascilloidea: Dascillidae). (Modified from Lawrence & Britton 1991; © CSIRO Australia.) RC = Radial cell; OC = Oblongum cell; WC = Wedge cell.

of the latter w o r k a n d based in part on t h e s t u d y by Haas & Kukalovä-Peck (2001) have been overlooked in s o m e s u b s e q u e n t studies. T h e m o s t i m p o r t a n t of these deal w i t h venation in t h e medial field a n d will be repeated here: CuA = Cu, CuA 1 + 2 = CuA, CuA 3 + 4 = CuP, C U A 2 = CUA 3 + 4 , AA = AA 3+4 , AA 1+2 = AA 3 , AA 3+4 = AA 4 .

reduced in beetle wings) usually contains AP 3 + 4 (which may be forked to f o r m AP 3 a n d AP 4 ) a n d occasionally a small AP 1+2 may also be present. In Lawrence et al. (1999 b) t h e t e r m "anal lobe" was used in narrower sense for those instances w h e r e the anal field was separated f r o m t h e r e m a i n d e r of the w i n g by a deep notch.

Anal Area (see Anal Lobe).

Anal N o t c h . embayment.

Deep a n d

usually n a r r o w

anal

Anal Embayment. Embayment in the wing margin at the e n d of t h e anal fold w h i c h may be weak a n d g r a d u a l or consist of a deep notch. Anal Field (see Anal Lobe).

Ano-Jugal Lobe (see Anal Lobe).

Anal Fold. Fold b e t w e e n AA 3+4 a n d AP; d e l i m i t i n g t h e anal lobe.

Apical Area (see Apical Field),

Anterior Wing Strut (see Radial Bar).

Apical Extensions of RP. Branches of RP (usuAnal Lobe. Portion of the w i n g basad of t h e anal fold. T h e anal lobe (actually h o m o l o g o u s w i t h t h e ano-jugal lobe b u t t h e jugal c o m p o n e n t is highly

ally RP1 a n d RP2) e x t e n d i n g apicad of t h e R-M loop. In some taxa t h e r e may be two or three branches, in others only RP2 remains a n d branches

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John F. Lawrence, Rolf G. Beutel, Richard Α. Β. Leschen and Adam Slipiriski

may be vaguely indicated or absent. Linear, often oblique sclerotizations in the apical field of various elateroid wings (Crowson 1961; Dolin 1975) may represent remnants of these radial extensions, b u t this is uncertain. Apical Field. Portion of the wing m e m b r a n e lying apicad of the radial cell (Polyphaga), cross-vein r4, the R-M loop (Polyphaga), the oblongum cell (most Archostemata, Myxophaga and Adephaga) and medial spur, and often containing terminal branches of RA and RP or the elateroid sclerites possibly derived f r o m them. Basal Portion of RP. Portion of RP between its origin (usually obliterated) and its junction with an R-M cross-vein forming the R-M loop or the base of the oblongum cell. Rarely complete and very short or even absent in some taxa. B e n d i n g Zone. Area of weakness in the radial and medial bars which allows t h e m to be curved towards one another when the bar is rotated in a certain way (by movements of the axillary sclerites). The weakness may be caused by a series of transverse impressions or by a flattening of the strut. Bending zones may be relatively broad so that bending is gradual or may be shortened to the point where they are more like a sharp hinge. Bending zones are found in only a few of the polyphagan families included in this volume. Cross-vein Subapical cross-vein between RA 3+4 and RP; transverse (perpendicular to long axis of wing) or strongly oblique in Archostemata, Myxophaga and Adephaga and often slightly oblique to longitudinal (parallel to long axis of wing) in Polyphaga; rarely complete and often absent. Termed the "radial recurrent vein" in some older systems of nomenclature. Cross-vein r4. Apical cross-vein between RA 3+4 and RP; always more or less transverse and usually complete. Cubito-Anal R e m n a n t . Occasionally used when a single c o m p o u n d vein consisting of Cu, CuP and AA3 remains in the medial field. D e f l e x i o n Zone (see Bending Zone). Free Veins. Veins not joined apically to another vein and often extending to the wing margin; in the medial field usually consisting of MP 3 + 4 (or MP 3 a n d MP4), CUA2, AA3 a n d AA4.

Medial Area (see Medial Field). Medial Bar. Posterior wing strut, entirely of MP 1+2 .

consisting

Medial B i n d i n g Patch (see Medial Fleck).

Medial Bridge. Bridge at the base of the wing connecting RA with MP 1+2 ; important in the folding and unfolding of the wing. Medial Embayment. Emargination of the wing membrane at or beyond the end of the medial spur. Medial Field. Portion of the wing m e m b r a n e between medial bar (MP 1+2 ) and the anal fold and usually containing branches of MP 3 + 4 , Cu and AA 3+4 . Veins in the medial field, referred to by Crowson (1955, p. 91) as "anal veins in the main group", tend to anastomose in various ways and reduction of free veins f r o m five to one or none is relatively common, especially in small species. Medial Fleck. Binding patch located in the medial field. In Polyphaga this is located in f r o n t of MP 3 + 4 . In some Adephaga a non-homologous fleck is located behind MP 3 + 4 . and usually containing MP 3 + 4 , Cu and AA 3+4 and their branches. Medial Spur. Apical extension of MP 1 + 2 beyond the R-M loop or oblongum cell. Very short and straight or absent in Archostemata, Myxophaga and Adephaga and usually longer and posteriorly curved in Polyphaga. Median Area. Error in Lawrence etal. (1999 b) (see Medial Field). Median Bar. Error in Lawrence etal. (1999 b) (see Medial Bar). Medio-Cubital Brace. Reinforced cross-vein joining MP 1 + 2 and Cu near their bases, b u t usually broken in the middle by a fold. M P 3+4 „ , : Cross-vein a n d Basal Spur. Sub-basal cross-vein joining MP 3 + 4 to MP 1 + 2 and often accompanied by a basal spur, which is a r e m n a n t of the true base of the vein. When the spur is absent, the cross-vein may appear to be the base of MP 3 + 4 , b u t in a more apical position. O b l o n g u m Cell. Cell in most Archostemata, Adephaga and Myxophaga formed between two R-M cross-veins. Posterior w i n g strut (see Medial Bar). Radial Bar. Anterior wing strut consisting of an alignment and fusion of ScP and RA. Radial Cell. Forking of RA into RA 1+2 and RA 3+4 and a subsequent rejoining of these two branches near the anterior wing margin in Polyphaga. Not homologous with cells formed by cross-veins between RA 1+2 and RA 3 or RA 3+4 in Archostemata, Myxophaga and Adephaga.

Glossary of morphological terms

15

Medial Spur Anal Notch J

^

AA.

CuA„ MP.

RC

AP,

R-M Loop SA o

•Medial Spur

.CuA

Β •t AA,

V AA,

-MP, CuA„

\ MP,

Fig. 2.3. Coleopteran hind wings: A, Osslimusfreyi (Cobos) (Polyphaga: Elateroidea: Elateridae); B, Stichotomus sp. (Polyphaga: Elateroidea: Elateridae). (Modified from Calder 1996; © CSIRO Australia.)

Radio-Medial Loop (see R-M Loop). R-M Loop. Apically arched R-M cross-vein joining the basal portion of RP and M P 1 + r Homologous to the apical side of the oblongum cell. Often forming a continuous curve between RP and MA, but sometimes narrowed so that an acute angle is formed.

the first being formed at the wing base between Cu and AA3+4, with CuP at its apex.

Adult Abdomen Accessory Lobes. Paired lobes articulated with the apicale in the aedeagus of various Tenebrionoidea.

Second Cubito-Anal Cell (see Wedge Cell).

Aedeagus. Combined structure consisting of phallobase, parameres and penis.

Subcubital Fleck (see Medial Fleck).

Apicale. Formed by the fusion of the two parameres.

Transverse Folds. More or less obliquely transverse folds; involved in the shortening of the wing length.

Baculum. Sclerotized bar or strut which serves to support a membranous ovipositor. Paired longitudinal baculi are often present on the proctiger and paraprocts, and a transverse baculum often occurs at the base of the coxite.

Wedge Cell. Cell formed by CuA anteriorly, CuP basally, AA3 posteriorly, and CuA3+4 apically and usually giving rise apically or subapically to one or more branches of Cu and to the terminal portion of AA3. This cell is also referred to as the 2nd cubito-anal cell,

Basal Piece (see Phallobase). Basale (see Phallobase).

16

John F. Lawrence, Rolf G. Beutel, Richard Α. Β. Leschen and Adam Slipiriski

Bursa. Enlargement of the female genital tract, which is often a blind sac at the anterior end of the tract (anterior bursa), but may be an enlarged section of the tract in the vicinity of the common oviduct (vaginal bursa). Connate Ventrites. Visible sternites which are not freely movable. This is not always easy to determine in lightly sclerotized abdomens, and there tends to be a continuum from sclerites which are capable of some limited movement to those which are solidly fused together. In rare cases the sutures between connate ventrites may be partly to almost entirely effaced. Coxite (see Gonocoxite). Cucujiform Aedeagus. That type of aedeagus in which the phallobase forms a sheath or ring partly or entirely enclosing the penis. Double Tegmen. Referring to a type of cucujiform aedeagus found in a number of Cleroidea plus the cucujoid families Biphyllidae and Byturidae in which the tegmen bears a pair of anterior struts and an opposing median anterior strut. Endophallus. Inverted membranous tube within the penis and sometimes armed with a variety of sclerotized structures.

Ovipositor. In female Coleoptera usually referring to the combined proctiger, paraprocts, gonocoxites and styli. One or more of these elements may be absent in certain groups and in a few families there is no distinct ovipositor, but just a vulva surrounded by membrane. Parameral Piece (see Apicale). Parameres. Paired structures articulated with the posterior end of the phallobase in aedeagi of the trilobate type; sometimes fused to the phallobase, fused together in various ways or fused to the anterior end of the penis. Paraproct. Referring to one of the two hemitergites or laterotergites of segment IX in most female beetles in which segment IX has become entirely divided into two parts which lie on either side of tergite X. Penis. Major intromittent organ containing the inverted endophallus and the opening of the ejaculatory duct; enclosed by the phallobase and parameres or the tegmen. Phallobase. Anterior or basal portion of the aedeagus to which the parameres are attached in aedeagi of the trilobate type. In aedeagi of the cucujiform type, the phallobase forms most of the sheath or ring enclosing the penis.

Epiproct (see Proctiger). Genital Ring. Ring-like structure surrounding the aedeagus and consisting of segments IX and X, which may be partly or entirely fused together. Gonangulum (see Gonocoxite). Gonocoxite. One of two coxites associated with segment IX in female Coleoptera. The coxite is often divided into a basal and apical lobes, the first of which may have a transverse or oblique baculum which articulates with the paraproctal baculum. The apical lobe is further subdivided in some taxa. The basal coxital lobe may be homologous to the gonangulum of Scudder (1961).

Postcoxal Lines of First Abdominal Ventrite. Slightly raised ridges of abrupt edges of impressions at the anterior edge of ventrite 1. Usually delimiting crural impressions. Proctiger. Referring to tergite X in both male and female; lies immediately above the anal opening and may become membranous in some groups. Spermatheca. Usually referring to a sac-like structure, often sclerotized; attached by a slender duct to the genital tract. Spermathecal Gland. Gland associated with the spermatheca, usually either joined directly to it or to the spermathecal duct.

Gonostylus (see Stylus). Hemitergite IX (see Laterotergite IX, Paraproct). Internal Sac (see Endophallus).

Spiculum Gastrale. Slender strut formed by an anterior extension of the subgenital plate. Spiculum Relictum. Anterior strut on sternite VIII in the male.

Lateral Lobes (see Parameres). Laterotergite IX. Paired plates formed when tergite IX is divided into two parts in either male or female (see Paraproct). Median Lobe (see Penis).

Spiculum Ventrale. Anterior strut on sternite VIII in the female. Stylus. Small lobe articulated apically or subapically to the coxite in the ovipositor of most Coleoptera.

Glossary of morphological terms

Subgenital Plate. Ventral portion of the genital ring, formed by a fusion of sternite IX and anteroventral extensions of tergite IX or laterotergites IX. Tegmen. Usually synonymous with the phallobase in aedeagi of the cucujiform type, in which the phallobase forms a sheath or ring partly or entirely surrounding the penis (see Cucujiform Aedeagus). When the parameres are fused to the end of the phallobase, the term tegmen is often used for the entire combined structure. Tegmenite. Slender sclerite articulated with sternite IX or in the connecting membrane between the sternite IX and the phallobase. Tergite X (see Proctiger). Trilobate Aedeagus. Basal type of aedeagus in Coleoptera, which consists of a penis lying dorsad of a phallobase and paired, articulated parameres. In many Coleoptera, including most Archostemata, Adephaga and Myxophaga, as well as a variety of Polyphaga, the phallobase appears to be absent, having become membranous or fused to the parameres. Vaginal Palp. Combined gonocoxite and stylus (Mikoleit 1973). Valvifer. Used inconsistently by Tanner (1927) but in general homologous with the paraproct (see Paraproct). Ventral Sclerite. Slender longitudial sclerite lying ventrally between the coxites and below the vulva. Ventrite: Abdominal sternite exposed in intact specimens; not concealed by the metacoxae or enclosed within the abdominal apex. The first ventrite in most Polyphaga and in Myxophaga and Archostemata is sternite III, but it is sternite II in Adephaga and in some groups of Polyphaga. Vulvar Sclerite (see ventral sclerite). Larval Head Coronal suture (see Epicranial Stem). Ecdysial lines. Lines of weakness on the dorsal head surface where the cuticle separates during moulting.

17 Frontal Sutures (see Frontal Arms). Gula. Sclerotised region posterad of the posterior tentorial pits. Hypopharyngeal Bracon. More or less sclerotized (but not usually pigmented) bridge j oining the ventral mandibular articulation on each side with the hypopharynx. Hypopharyngeal Sclerome. Sclerotized bar or tooth-like structure on the hypopharynx; acts in conjunction with the mandibular molae. Hypostomal Cavity. Excavation of the ventral portion of the head capsule enclosing the retracted maxillae and labium. Hypostomal Ridges. Sclerotized lateral edges of the hypostomal cavity. Hypostomal Rods. Posterior continuations of the hypostomal ridges beyond the maxillary bases and onto the ventral portion of the head capsule. Ligular Sclerome. Sclerotized wedge-like sclerite formed from the ligula. Not equivalent to the hypopharyngeal sclerome, which is located well posterad of this on the surface of the hypopharynx. Maxillolabial Complex. The combination of maxillae and labium when these are closely associated and mainly or exclusively moved in a nearly vertical direction (without an intervening maxillary articulating area). Medial Endocarina. Dorsal internal ridge on the head usually lying beneath the epicranial stem, when present, and often extending forward between the frontal arms. May also be present when ecdysial lines are absent. Mola. Usually basal and armed with asperities, tubercles or transverse ridges. In some groups somewhat reduced and sub-basal. Paired Endocarinae. Pair of dorsal internal ridges on the head, usually lying beneath the bases of the frontal arms.

Epicranial suture (see Ecdysial Lines, Epicranial Stem, Frontal Arms).

Prostheca. Located distad of the basal mola and either membranous, sclerotized but hyaline, or consisting of hair-like structures of various kinds. Similar structures in the absence of a mola are not referred to as prosthecae.

Frontal Arms. Paired ecdysial lines either arising from the epicranial stem or independently joined to the posterior edge of the head capsule.

Sensorium. Usually refers to a large sensillum located on the preapical antennomere. Although it is usually conical or palpiform, it may be dome-like

Epicranial Stem. Median ecdysial line, which forks to form the frontal arms.

18

John F. Lawrence, Rolf G. Beutel, Richard Α. Β. Leschen and Adam Slipiriski

or occasionally multiple or complex (consisting of several dome-like sections). Occasionally located on the apical palpomere or very rarely (some Phalacridae) on the basal one. Sensory Appendage (see Sensorium). Ventral Epicranial Ridges. Pair of ridges extending posteriorly from the ventral mandibular articulations; located laterad and ventrad of the hypostomal ridges and more or less supporting the maxillolabial complex. Ventral Mouthparts. Refers to the maxillae plus the labium or the maxillolabial complex w h e n these structures form a single unit. Ventral Mouthparts: Retraction. The retraction of the ventral mouthparts is the distance between the ventral mandibular condyles and the basal attachments of the cardines or the depth of the hypostomal cavity. In larvae w i t h protracted ventral mouthparts, this distance or depth is minimal or zero, and the mandibles and maxillae are attached approximately at the same level.

Tibiotarsus. Formed by the fusion of the tibia and tarsus in larvae of M y x o p h a g a and Polyphaga.

Larval Abdomen Interurogomphal Pit. Pit or one of a pair of pits lying between the urogomphi. Laterosternite. Sclerotized plate lying between the main sternal plate and the tergosternal fold. Laterotergite. Sclerotized plate lying between the inflected part of the tergum and the tergosternal fold. One of the laterotergites usually contains the spiracle. Pygopods. Paired projections on segment X which may or may not bear hooks and usually assist in locomotion. Tergosternal Fold. Fold or membrane connecting the abdominal terga and sterna.

Larval Thorax Basisternum. Anterior part of the larval sternum, anterad of the sternal pits. Cervicosternum. Small anteromesal sclerite, separate from the presternum. Laterotergite. One of the sclerotized plates lying between the inflected part of the tergum and the pleural sclerites (precoxale and postcoxale); one of the mesothoracic laterotergites may contain the spiracle. Precoxale. Pleural (episternal) sclerotised anterolaterad of the coxal articulation.

Tarsungulus. The terminal of claw s e g m e n t of the larval leg in Polyphaga as used by those w h o maintain that this s e g m e n t represents a fusion of the tarsus and pretarsus (see Pretarsus and Tibiotarsus).

area

Postcoxale. Pleural (epimeral) sclerotised area posterolaterad of the lateral part of the coxal articulation. Presternum. Separate anteromedian sclerite anterad of the basisternum; sometimes large and more or less triangular, but often vaguely defined or absent. Pretarsus. The terminal segment of the larval leg, consisting of a pair of claws in most Adephaga and a few Archostemata or a single claw in some Adephaga, most Archostemata and all M y x o p h a g a and Polyphaga.

Urogomphi. General term used for a number of non-homologous, doubtfully homologous or rarely clearly homologous paired cuticular outgrowths arising from the posterior end of tergum IX. The basally articulated u r o g o m p h i occurring in some Adephaga and Staphyliniformia were formerly h o m o l o g i z e d w i t h cerci, w h i c h occur on segment X or XI in some other orders, b u t these, like the fixed urogomphi, have clearly evolved on several occasions w i t h i n Coleoptera.

Larval Spiracles Spiracular Closing Apparatus. Apodeme attached to the trachea at or near the base of the spiracular atrium. It is involved in the closure of the trachea. Annular-biforous Spiracle. T y p e of spiracle w i t h the m a i n spiracular o p e n i n g accompanied by a pair of accessory o p e n i n g s . In some families (e.g., Histeridae, Nitidulidae), these accessory o p e n i n g s are m u c h l o n g e r than the m a i n opening, so that they resemble the b i f o r o u s type of spiracles; h o w e v e r the spiracular scar is always absent. Annular-multiforous Spiracle. Type of spiracle w i t h a number of accessory openings around the perimeter of the main opening.

Glossary of morphological terms

19

Annular-uniforous Spiracle. Type of spiracle with the main spiracular opening accompanied by a single accessory opening. Bicameral Spiracle).

Spiracle

(see

Annular-biforous

Biforus Spiracle. Spiracular opening entirely blocked by a medium septum with a narrow, slitlike opening on either side. Biforous spiracles have evolved in connection with the elateroid ecdysial process (Hinton 1947) and are always associated with a spiracular scar. Bilabiate Spiracle (see Biforous Spiracle). Cribriform Spiracle. Spiracular opening entirely or partly blocked by a sieve plate. Three or four types of cribriform spiracles have evolved in connection with the elateroid ecdysial process (Hinton 1947) and are associated with a spiracular scar, and another two or three in which normal ecdysis occurs and the spiracular scar is absent. Ecdysial Scar (see Spiracular Scar). Stigmatic Scar (see Spiracular Scar). Spiracular Scar. Collapsed opening of a tube through which the spiracular lining is removed at ecdysis. Occurs only in those larvae which have biforous or cribriform spiracles and which undergo the elateroid type of ecdysis. Unicameral Spiracles).

Spiracle

(see

Annular-uniforous

Multicameral Spiracle (see Annular-multiforous Spiracle).

Literature Beutel, R. G. & Haas, F. (2000): Phylogenetic relationships of the suborders of Coleoptera (Insecta). -Cladistics 16:103-141. Beutel, R. G. & Lawrence, J. F. (2005): 4. Coleoptera, Morphology. Pp. 23-27 in Beutel, R. G. & Leschen, R. A. B. (eds.) Handbook of Zoology. Volume IV Arthropoden Insecta. Part 38. Coleoptera. Volume 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, polyphaga partim). Walter de Gruyter, Berlin, New York. Calder, A. A. (1996): ClickBeetles. Genera of the Australian Elateridae (Coleoptera). χ + 401 pp. CSIRO Publishing (Monographs on Invertebrate Taxonomy. Vol. 2), Collingwood, Victoria. Campau, E. J. (1940): The morphology of Chauliognathus pennsylvanicus (DeGeer) (Coleoptera: Cantharidae). - Microentomlogy 5 (3): 57-85. Crowson, R. A. (1938): The metendosternite of Coleoptera: a comparative study. - Transactions of the

Royal Entomological Society of London 87 (17): 3 9 7 416,13 pis. - (1944): Further studies on the metendosternite in Coleoptera. - Transactions of the Royal Entomological Society of London 94(2): 2 7 3 - 3 1 0 , 1 0 pis. - (1955): The Natural Classification of the Families of Coleoptera. 187 pp. Nathaniel Lloyd, London. - (1961): On some new characters of classificatory importance in adults of Elateridae (Coleoptera). - The Entomologist's Monthly Magazine 96: 158-161. - (1981): The Biology of Coleoptera. xii + 802 pp. Academic Press, London. Dolin, V. G. (1975): Wing venation in click beetles and its significance in the taxonomy of the family. - Zoologicheskii Zhurnal 54 (11): 1618-1633 (in Russian). Ferris, G. F. (1940): The myth of the thoracic sternites in insects. - Microentomology 5 (3): 87-90. Haas, F. & Kukalovä-Peck, J. (2001): Dermaptera hindwing structure and folding: new evidence for familial, ordinal and superordinal relationships within Neoptera (Insecta). - European Journal of Entomology 98:445-509. Hinton, Η. E. (1947): On the reduction of functional spiracles in the aquatic larvae of the Holometabola, with notes on the moulting process of spiracles. - Transactions of the Royal Entomological Society of London 98:449-473. Kukalovä-Peck, J. & Lawrence, J. F. (1993): Evolution of the hind wing in Coleoptera. - Canadian Entomologist 125:181-258. - (2004): Use of hind wing characters in assessing relationships among coleopteran suborders and major endoneopteran lineages. - European Journal of Entomology 101 (1): 95-144. Lawrence, J. F. (1991): Order Coleoptera. Pp. 144-298 in Stehr, F. W. (ed.) Immature Insects, Vol. 2. Kendall/ Hunt Publishing Co., Dubuque, Iowa. Lawrence, J. F. (1999): The Australian Ommatidae (Coleoptera: Archostemata): new species, larva and discussion of relationships. - Invertebrate Taxonomy 13: 369-390. Lawrence, J. F. & Britton, Ε. B. (1991): Coleoptera (Beetles). Pp. 543-683 in CSIRO Division of Entomology (ed.) Insects of Australia: a Textbook for Students and Research Workers, Second Edition. Vol. 2. Melbourne University Press, Carlton, Victoria. Lawrence, J. F., Hastings, A. M., Dallwitz, Μ. J., Paine, T. A. & Zürcher, Ε. J. (1999 a): Beetle Larvae of the World: Descriptions, Illustrations, Identification, and Information Retrieval for Families and Subfamilies. CD-ROM, Version 1.1 for MS-Windows. CSIRO Publishing, Melbourne. - (1999 b): Beetles of the World: A Key and Information System for Families and Subfamilies. CD-ROM, Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne. Leschen, R. A. B., Lawrence, J. F. & Slipinski, S. A. (2005): Classification of basal Cucujoidea (Coleoptera: Polyphaga), with a cladistic analysis, descriptions of new genera and species, and keys to adults and larvae of cucujoid families. - Invertebrate Systematics 19:17-73.

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Matsuda, R. (1960): Morphology of the pleurosternal region of the pterothorax in insects. - Annals of the Entomological Society of America 53 (6): 712-731. Mikoleit, G. (1973): Über den Ovipositor der Neuropteroidea und Coleoptera und seine phylogenetische Bedeutung (Insecta, Holometabola). - Zeitschrift für Morphologie der Tiere 7 4 : 3 7 - 6 4 .

Scudder, G. G. E. (1961): The comparative morphology of the insect ovipositor. - Transactions of the Royal Entomological Society of London 113 (2): 2 5 - 4 0 . Stickney, F. S. (1923): The head-capsule of Coleoptera. -Illinois Biological Monographs 8 (1): 104 pp., 8 pis. Tanner, V. M. (1927): A preliminary study of the genitalia of female Coleoptera. - Transactions of the American Entomological Society 5 3 : 5 - 5 0 .

3. Adephaga (Addendum)

Introduction and Phylogeny Rolf G. Beutel, Ignacio Ribera and Michael Balke The recent discoveries of two new families - Aspidytidae and Meruidae - were doubtlessly rare highlights in adephagan studies. Aspidytidae is a monogeneric family with two species with a remarkable distribution in China and South Africa, respectively. It was described by Ribera etal. (2002). Specimens of Meru phyllisae Spangler & Steiner, the only species of Meruidae, were first collected in 1985, but only described and assigned to a new family 20 years later (Spangler & Steiner 2005). The adults of both groups and at least the larvae of Aspidytes niobe Ribera et al. inhabit different hygropetric habitats. Within a short time a considerable amount of morphological and molecular data was accumulated and the systematic position of both groups appears to be largely clarified. Both belong to the complex of dytiscoid families. Meruidae is likely the sistergroup of Noteridae, and Aspidytidae belongs to a clade which also includes Amphizoidae, Hygrobiidae and Dytiscidae (Beutel etal. 2006; Balke etal. 2008; see also ζ -3.1. and 3.2.).

Literature Balke, M., Ribera, I., Beutel, R., Viloria, Α., Garcia, M. & Vogler, A. P. (2008): Systematic placement of the recently discovered beetle family Meruidae (Coleoptera: Dytiscoidea) based on molecular data. - ZoologicaScripta 37:647-650. Beutel, R. G., Balke, M. & Steiner Jr., W. J. (2006): The systematic position of Meruidae (Coleoptera, Adephaga) and the phylogeny of the smaller aquatic adephagan beetle families. - Cladistics 22:102-131. Ribera, I., Beutel, R. G., Balke, M. & Vogler, A. P. (2002): Discovery of Aspidytidae, a new family of aquatic Coleoptera. - Proceedings of the Royal Society, Β 269: 2351-2157. Spangler, P. J. & Steiner, W. E., Jr. (2005): A new aquatic beetle family, Meruidae, from Venezuela. - Systematic Entomology 30:339-357.

3.1. Aspidytidae Ribera, Beutel, Balke and Vogler, 2002 Rolf G. Beutel, Michael Balke and Ignacio Ribera Distribution. Aspidytidae is a monogeneric family with only two known species with a widely disjunct distribution: Aspidytes niobe, discovered in 2001 in

the Western Cape Province of South Africa and described by Ribera etal. (2002); and Aspidytes wrasei, described one year later (Balke etal. 2003) although already discovered in 1995 by David Wrase and Michael Schtilke in the Chinese province Shaanxi (Qin Ling Shan, Hua Shan, 34.27N 110.06E). Biology and Ecology. Adults of both species and the larvae of Aspidytes niobe live in hygropetric habitats and are very likely predacious. A. niobe is known from two neighboring permanent water seepages flowing over exposed, nearly vertical rocks almost without vegetation, although areas around the seeps have a highly diverse plant cover. Adults were mostly resting on the rock surface, and walked rapidly when disturbed. Larvae were mostly found actively creeping over exposed rock surfaces in densely shaded areas (Alarie & Bilton 2005; I. Ribera & A. Cieslak pers. obs.). Aspidytes wrasei is known from a single locality, and was collected among stones and various plants at the base of a vertical hygropetric surface (Balke etal. 2003). Morphology, Adults (Ribera etal. 2002; Balke etal. 2003). Total length 4.8-7.0 mm. Body streamlined, without pronoto-elytral angle, convex dorsally. Dorsal side almost entirely black; parts of head and lateral margins of pronotum and elytra ferrugineous; ventral side rufous. Cuticle shiny. Head (Fig. 3.1.1 A) prognathous, distinctly shortened, wedge-shaped in lateral view. Compound eyes completely integrated in outline of laterally rounded head. Foramen occipitale wide. Posterior part of head distinctly retracted into prothorax, but without narrowed neck region. Tactile setae absent from head capsule. Clypeus distinctly narrowing towards labral articulation; frontoclypeal suture broadly interrupted medially. Clypeolabral suture almost straight. Labrum transverse, with wide median concavity. Anterior epipharynx with pair of large, sclerotized sensorial pegs. Antennae without pubescence or longer setae; inserted laterally, anterior to compound eyes, articulation not visible from above; scapus with globular basal piece and globular distal part separated by a distinct circular incision; distal part largely enclosing small pedicellus; length of 9segmented flagellum ca. 0.8 mm (A. wrasei); segments 3 and 11 longer than other antennomeres, 4 - 9 very slightly expanded, 6 - 1 0 with unusual fields of sensorial pores on ventral side; sensorial field present but vestigial on antennomere 5. Mandibles fairly short and compact, with semicircular outer margin; ventral condyle and dorsal socket strongly developed; dorsal side convex and ventral side flat; setae absent; dense row of hairs on ventral side extends from base to anterior edge close to mesal margin; apical part with pointed tooth and axe-like dorsal cutting edge;

22

Rolf G.Beutel, Michael Balkeand Ignacio Ribera

Fig. 3.1.1. SEM micrographs. Aspidytes niobe, adult; A, head, ventral view; Β, pro- and mesothorax, ventral view; C, metathorax and abdominal base, ventral view. semicircular emargination separates apical tooth from small retinaculum; mola and prostheca absent. Maxilla inserted in maxillary groove between submentum and compound eye; composed of semicircular cardo, basistipes, dististipes, lacinia, galea, palpifer and palp; most parts covered by mental lobes; galea palp-like, 2-segmented; palp 5-segmented, fairly short, scarcely overtopping labial palp. Submentum fused with gula posteriorly; border between both sclerites marked by small but distinct posterior tentorial pits; lateral margin rounded; posterior part largely smooth, with transverse meshes, distinctly convex like anterior gula. Anterior part flat, narrowing towards median line, laterally covered by stiff, anteriorly directed setae; mentum with well developed parabolic lateral lobes enclosing deep median emargination for reception of prementum; mental lobes with group of anteriorly directed setae; prementum with prominent median part and distinctly developed palpigers; palpiger with distinct

Fig. 3.1.2. SEM micrographs. Aspidytes niobe, male paratype; A, foreleg, ventral view, with attachment devices; B, modified adhesive hairs. lateral emargination; palpomere 1 short, curved, with deep lateral concavity for reception of base of palpomere 2; palpomere 2 longer, with curved outer margin, extended distally, with two short but distinct spines at anterior margin; palpomere 3 larger than 2, with strongly convex outer margin and apical field of sensilla. Anterior tentorial grooves long, fissure-shaped. Gular ridges high, with posterior apodeme for attachment of extrinsic head muscles; connected by thin tentorial bridge; medially fused laminatentorium and mid-gular apodeme absent.

Aspidytidae Ribera, Beutel, Balke and Vogler, 2002

Pronotum laterally equally rounded, with distinct lateral and posterior beads; disc black, lateral and posterior margin rufous; inflected pronotal margin distinctly widening posteriorly; posterior edge with dense row of hairs. Proventrite separated from propleura by distinct pleurosternal suture; anterior margin with fringe of thin hairs. Trochantin completely covered by propleural lamella. Prosternal process well developed, posteriorly rounded (A. wrasei) or truncate, with rounded posterolateral edges (A. niobe) (Fig. 3.1.1 B). Procoxal cavities open, withsclerotized, rufous internal bridge. Profurca short b u t extensive. Foreleg fairly short; procoxae globular, with short ventral condyle; femora with short row of stiff hairs at anterior margin (profemoral cleaning device); tibia with truncate apex and two curved terminal spurs; external spur slightly stronger; tarsomeres 1 - 4 short; tarsomere 5 ca. 2.5 times as long as basal tarsal segments; male tarsomeres 1 and 2 laterally slightly dilated and ventrally with numerous adhesive hairs with minute apical discs (Fig. 3.1.2); claws equal. Mesothorax slightly shorter than prothorax. Scutellum exposed. Elytra with lateral bead extending to apex; epipleura broad anteriorly, strongly narrowing towards abdominal apex; with polygonal meshes of different size and few punctures. Distinct pentagonal groove of mesoventrite articulates with prosternal process anteriorly (prothorax in retracted position) and with anteromedian process of metaventrite posteriorly (Fig. 3.1.1 C). Faintly impressed pleural suture separates anepisternum from moderately broad epimeron. Mesocoxal cavity laterally bordered by mesal edge of epimeron and narrow apical part of metathoracic anepisternum (= complex type cf. Bell 1967). Middle legs similar to front legs; femur without t u f t of spines; tibia and tarsus slightly

23

longer; tibia only very slightly extended distally; tarsomeres 1 and 2 of males laterally slightly dilated and ventrally with numerous adhesive hairs with minute apical discs (as in Fig. 3.1.2). Metathorax slightly longer than prothorax. Anepisternum triangular, with distinct bead along lateral and mesal margin. Epimeron covered by elytral epipleura. Ventrite, composed of praeepisternum and katepisternum, moderately shortened, not reaching lateral metacoxal margin laterally; metaventral process well developed, with rounded anterolateral edges. Transverse suture present as faintly impressed line and internal ridge or as internal ridge only (A. niobe); about as long as width of internal lamina of metacoxa, separating posterior katepisternum from anterior praeepisternal part of ventrite. Median longitudinal suture (= discriminal line or discrimen) very short, not surpassing posterior half of katepisternum. Metacoxa moderately extended, slightly longer than ventrite; lateral margin broadly contiguous with epipleural margin; mesal walls extensively fused, forming large intercoxal septum; anterior paramedian angle absent; inner lamina and coxal plates distinct, broadened posteriorly, distinctly curved outwards anteriorly (Fig. 3.1.1 C); metatrochanter larger than pro- and mesotrochanter; tibia and tarsus longer than those of middle leg, appearing thin and longish (length of femur 0.80 m m , tibia 1.05 m m , tarsus 0.95 m m ; ratio total body length/length of tibia + tarsus = 2.4 (ratio in Hygrotus impressopunctatus 1.7; Amphizoa lecontei 2.0; Carabus auratus 1.5). Wings well developed (examined in A. niobe), with distinct oblongum cell and moderately elongate medial setal binding patch (Beutel etal. 2006: fig. 11) (Fig. 3.1.3). Abdomen with six exposed ventrites equivalent to sternites II, III, IV, V, VI, VII (= last visible

Fig. 3.1.3. Aspidytes niobe, hind wing. From: Beutel etal. 2006 (modified, nomenclature of venation following Kukalovä-Peck & Lawrence 2004).

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Rolf G. Beutel, Michael Balke and Ignacio Ribera

Fig. 3.1.4. A-C, Aspidytes niobe, female paratype genitalia. Α, genital tract, lateral view: ra, ramen; gc, gonocoxa; lt, laterotergite; dp, dorsal pocket; va, vagina; vg, ventral gland; ovi, common oviduct; fd, fertilisation duct; sd, spermathecal duct; sp, spermatheca; B, gonocoxa, ramen and laterotergite (gc, ra, lt) in ventral view; C, gonocoxosternum, ventral view. D - F , Aspidytes wrasei female paratype genitalia. D, genital tract in lateral view, simplified (be, bursa copulatrix; dg, dorsal gland; dp, dorsal pocket; fd, fertilization duct; gc, gonocoxa; lt, laterotergite; ovi, common oviduct; sd, spermathecal duct; sp, spermatheca; va, vagina); E, gonocoxa, ventral view, some setae omitted; F, gonocoxosternum, ventral view. From: Balke, M., Ribera, I. & Beutel, R. G. (2003) in Water Beetles of China Vol. Ill, with kind permission.

Aspidytidae Ribera, Beutel, Balke and Vogler, 2002

sternite); length decreasing from III to VI. Sternite II only visible laterally of metacoxa (Fig. 3.1.1 C). Sternites III and IV partly fused, but with very distinct separating suture. Lateral bead present on all sternites, most prominent along lateral margin of sternite VII. Shape of sternite VII roughly semicircular, evenly rounded posteriorly, and devoid of modifications of its surface or hind margin. Male and female genitalia distinctly different between species. A. wrasei with median lobe of aedeagus simply curved. Symmetrical parameres of roughly triangular shape; apex stylus-like, with few short setae anteriorly; margin devoid of setae (see Balke et al. 2003: figs. 16, 17). A. niobe with median lobe of aedeagus composed of several lobes and membranous sacs (ultrastructural features could not be assessed due to lack of fully sclerotized specimens). Parameres broadly oval; apex rounded with few short setae. Gonocoxae (Fig. 3.1.4.) (nomenclature of Miller 2001) (= genital appendages IX) of females of A. wrasei roughly rectangular in ventral view, with very slightly curved lateral margins and blunt apex; cranially with short external apodeme; laterally with rather long setae; distally with shorter setae. Proximal portion rather broad in lateral view. Gonocoxa of A. niobe elongate in ventral view, with apex broadly rounded; cranially with short external apodeme; devoid of recognisable setation. Laterotergite (= tergite IX; "Tergumhälften IX" or "Tergum IX" of Burmeister 1976) of both species articulated with cranial tip of gonocoxa, extended caudally, shorter than gonocoxa. Gonocoxosternite (Coxosternum; = invaginated sternum VIII; "Gonocoxosternum" of Burmeister 1976) of A. wrasei somewhat rectangular, with short anterior lobe (= antero-external angle) and median anterior concavity; antero-internal angle also forming lobe equally in size to anterior lobe; posterior margin with fringe of conspicuous long setae, comparable in length to height of gonocoxosternite. Ramen of A. wrasei ("Vulvarsklerite": Burmeister 1976) short, triangular, rufous. Of similar shape in A. niobe, with only few short, scarcely visible setae along posterior margin. Vagina of A. wrasei rather short and saclike, with cranial expansion (probably representing bursa copulatrix) and conspicuous, approximately triangular dorsal pocket (Fig. 3.1.4 D: dp). Vagina and sac-like expansion not distinctly separated. Longish gland located in dorsomesal position ("dorsal gland", dg). Spermathecal duct (sd) comparatively short and broad, originating very close to opening of common oviduct (co) into bursa or vaginal expansion; spermatheca (sp) without any obvious modifications (50x) in both species; spermathecal gland always absent. Spermatheca and insertion of oviduct connected by narrow fertilisation duct (fd). Vagina of A. niobe with short, triangular caudal pocket on dorsal side (dp); caudal part of vagina covered with numerous spiny scales (dotted in Fig. 3.1.4 A); compact gland located in ventromesal position ("ventral gland", vg); spermathecal duct (sd) originating dorsally to opening of ommon oviduct (ovi) into vagina.

25

Morphology, Larvae (Alarie & Bilton 2005; Balke et al. 2005) (Figs. 3.1.5-3.1.7). Total length of mature larvae (excl. urogomphi) ca. 7 mm. Robust, almost onisciform anteriorly, but abdomen more subcylindrical towards apical segment IX. Head and body strongly pigmented on dorsal side. Cuticle smooth. Sclerites set with short setae and also setae of medium length (see Alarie & Bilton [2005] for a detailed account of the distribution of setae and pores). Head prognathous, very slightly inclined (Fig. 3.1.6 A, B). Posteriorly retracted into prothorax but without distinct neck region. Moderately flattened, slightly broader than long, distinctly narrower than prothorax. Greatest width (ca. 1 mm) at posterior 1/3 of head; head capsule abruptly narrowing towards foramen occipitale and gradually narrowing towards clypeal region. Foramen

Fig. 3.1.5. SEM micrographs. Aspidytes niobe, larvae; A, head and thorax, ventral view; B, thorax, dorsolateral view; C, abdomen, dorsal view. From: Balke etal. 2005 (modified).

26

Rolf G.Beutel, Michael Balkeand Ignacio Ribera

Fig. 3.1.6. SEM micrographs. Aspidytes niobe, larva; A, head, dorsal view; Β, head, ventral view; C, clypeolabral region, dorsal view; D, clypeolabrum and mouthparts, frontal view; E, abdominal tergite I, with spiracle; F, abdominal tergite IV, with spiracle. occipitale fairly large, broadly oval. Six moderately sized stemmata present posterad of antennal articulatory region, nearly arranged in a circle. Frontal suture (=frontal arms) present, lyriform, but with very indistinct indentation. Frons distinctly elongated posteriorly. Coronal suture accordingly shortened, ca. 1/3 as long as entire head capsule. Labrum fused with frontoclypeus; adnasalia (=epistomal lobes) distinct, slightly asymmetric; with several long setae and dense brush of microtrichia along mesal part of anterior margin and ventral surface; nasale without teeth, separated from adnasalia by very deep grooves; with several major sensilla (probably six) embedded in very dense field of smaller sensilla and microtrichia (Fig. 3.1.6 C, D). Antennae 4-segmented; antennomere 1 short; antennomere 2 ca. 4 times as long as wide, slightly

extended distally; antennomere 3 slightly shorter than 2, also extended distally, with distinct sensorial appendage on ventral side of apical part; antennomere 4 very small and spindle-shaped, inserted on apex of segment 3. Mandibles moderately long, with fairly broad basal part; sickle-shaped distal part distinctly marked-off, with deep mesal concavity enclosed by ventral and dorsal cutting edge; retinaculum small but distinct; mola and prostheca absent. Maxillae inserted in shallow groove at anterior margin of head capsule, between anterior labium and mandibular articulation; cardo slightly narrower than stipital base, almost vertically oriented; stipes fairly short, with three long setae close to lateral margin; lacinia absent; galea inserted on stipital apex, 2-segmented. Submentum completely fused with gula posteriorly and with ventrolateral

Aspidytidae Ribera, Beutel, Balke and Vogler, 2002

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Fig. 3.1.7. SEM micrographs. Aspidytes niobe, larva; Α, abdominal apex with urogomphi, ventral view; Β, segments VIII and IX, dorsal view. From: Balke etal. 2005 (modified). wall of head capsule laterally; mentum membranous, short; prementum short, without ligula; palpiger well developed; palp ventrally directed, 2-segmented; both segments about equally long, but basal segment distinctly wider. Posterior tentorial pits small but distinct, widely separated, distant from posterior margin of head capsule. Gula moderately long and broad. Tentorium with elongate caudal arms serving as attachment area of strongly developed M. tentoriopharyngalis. Prepharynx fairly short but not transverse; M. clypeopalatalis composed of several subcomponents; M. tentoriobuccalis anterior present and large; precerebral and postecerebral pharyngeal dilators strongly developed. Cerebrum located in posterior head region; suboesophageal ganglion large, located below well developed, nearly straight tentorial bridge and ventral wall of anterior pharynx. Prothorax largest segment of body (Fig. 3.1.5); cuticle smooth and shiny, with numerous short setae inserted in small pores and longer setae along lateral margin and anterior margin of posterior collar. Pronotum shield-like, rounded anteriorly and laterally, with a distinct bead along lateral margin; epipleura, i. e., inflected part of pronotum (paratergal lobe) distinctly developed; posterior collar fairly broad, with parallel longitudinal riffles. Presternum not separated from epipleura laterally. Basisternum small, with narrow transverse anterior part and narrow median part between procoxae. Procoxae large and prominent, cone-shaped; distinct lateral concavity for reception of profemora in repose delimited by sharp anterior edge and less conspicuous posterior edge; anterior edge with several long setae; trochanter well developed; femur rather short, ca. 2 times as long as wide, with 4 spines along ventral edge; tibia shorter and narrower, moderately extended distally, with several apical setae; tarsus shorter than tibia, cylindrical, with two equal, strongly developed claws; single seta present between claws. Mesothorax shorter than prothorax, otherwise similar. Paratergal lobes slightly broader. Anterior margin straight, overlapped by posterior pronotum. Trochanters and femora slightly larger than on prothorax.

Metathorax very similar to mesothorax. Coxae larger; femora and tibiae slightly longer. Abdominal (Figs. 3.1.5-3.1.7) segmentl approximately half as long as metathorax; with lateral bead, narrow paratergal lobes and posterior collar; spiracle small, located on lateral part of tergite. Sternal part with two distinct oblique ridges. Tergites of segments II-VIII laterally fused, forming ring-shaped structure; paratergal lobes absent; small spiracles positioned on tergites, close to posterior margin. Segments II-V distinctly longer than I, about equally broad. Segments VI and VII shorter; VI narrower than V, and VII narrower than VI; VII about as long as V and distinctly narrowing posteriorly. Strongly developed urogomphi inserted on segment VIII ventrolaterally, enclosing small but distinctly developed segment IX (visible in ventral and dorsal view); basal segment of urogomphi (Fig. 3.1.7) ca. 1.5 times as long as segment VIII; with broad bases almost contiguous ventromedially; evenly tapering towards apex; with several long setae; distal segment very slender, cylindrical, slightly more than half as long as basal segment; with long setae inserted on apex. Pupa. Unknown Eggs. Unknown. Phylogeny and Taxonomy (Ribera et al. 2002; Balke et al. 2005). Combined analyses of morphological and molecular data strongly support a position of Aspidytes within a clade which also comprises Hygrobiidae, Amphizoidae and Dytiscidae. Typical dytiscoid features are the extensive fusion of the mesal metacoxal walls, the loss of two of three metathoracic furcacoxal muscles (Mm 81, 83), and the loss of the abdominal segment X in larvae. A feature less derived than in other dytiscoid groups is the incomplete reduction of the larval abdominal segment IX (Alarie & Bilton 2005). The phylogenetic relationships among the families of this clade are much less understood (Balke etal. 2005; Beutel etal. 2006). The most complete analyses to date (Balke etal. 2005, with fifty-three morphological characters

28 of adults and larvae and ca. 6 Kb from six nuclear and mitochondrial genes) placed Aspidytes as sister to Amphizoa, although with relatively low support.

Acknowledgements We are grateful to Hans Fery for many valuable comments and corrections.

Literature Alarie, Y. & Bilton, D. T. (2005): Larval morphology of Aspidytidae (Coleoptera: Adephaga) and its phylogenetic implications. - Annals of the Entomological Society ofAmerica 98:417-430. Balke, M., Ribera, I. & Beutel, R. G. (2003): Aspidytidae: on the discovery of a new beetle family: detailed morphological analysis, description of a second species, and key to fossil and extant adephagan families. Pp. 53-66 in Jäch, Μ. Α. & Ji, L. (eds.) Water Beetles of China. Vol. III. -Zoologisch-Botanische Gesellschaft in Österreich und Wiener Coleopterologenverein, Wien. Balke, M., Ribera, I. & Beutel, R. G. (2005): The systematic position of Aspidytidae, the diversification of Dytiscoidea (Coleoptera, Adephaga) and the phylogenetic signal of third codon positions. -Journal ofZoological Systematics andEvolutionary Research 43 (3): 223-242. Bell, R. T. (1967): Coxal cavities and the classification of the Adephaga (Coleoptera). - Annals of the Entomological Society ofAmerica 60:101-107. Beutel, R. G., Balke, M. & Steiner Jr., W. J. (2006): The systematic position of Meruidae (Coleoptera, Adephaga) and the phylogeny of the smaller aquatic adephagan beetle families. - Cladistics 22:102-131. Burmeister, Ε. G. (1976): Der Ovipositor der Hydradephaga (Coleoptera) und seine phylogenetische Bedeutung unter besonderer Berücksichtigung der Dytiscidae.-Zoomorphologie 85:165-257. Miller, Κ. B. (2001): On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. -InsectSystematicsandEvolution 32:45-92. Kukalovä-Peck, J. & Lawrence, J. F. (2004): Use of hind wing characters in assessing relationships among coleopteran suborders and major endoneopteran lineages. -European Journal of Entomology 101 (1): 95-144. Ribera, I., Beutel, R. G., Balke, M. & Vogler, A. P. (2002): Discovery of Aspidytidae, a new family of aquatic Coleoptera. - Proceedings of the Royal Society, Β 269: 2351-2157.

3.2. Meruidae Spangler and Steiner 2005 Rolf G.Beutel, Michael Balke and Warren E. Steiner Distribution. The only known species Meruphyllisae Spangler & Steiner 2005 occurs in Venezuela, district of Amazonas, at El Tobogän de la Selva, 40 km south

Rolf G. Beutel, Michael Balke and Warren E. Steiner of Puerto Ayacucho, 5.23.1461 N, 67.36.5995 W, c. 140 m elevation. Biology and Ecology (Spangler & Steiner 2005). The type series of Meru phyllisae was collected on the broad, shallow surface and margins of a cascade flowing over open bedrock. The cascade is fed by a white-water stream, the Rio Coromoto. Leaves adhering to the wet rock are present in the slowly flowing sections. A water analysis provided the following data: pH 5; oxygen 12 ppm; hardness 0. The water temperature was 28°C when most of the specimens were collected. Some specimens of M. phyllisae were taken by collecting leaves from the seepage areas into a finemesh dip-net. Most specimens were found in partly submerged root mats, which held decaying leaves, sticks and other debris at the edges of the open or semi-shaded stream margins. Species of the following groups were collected in the same microhabitat: Dytiscidae (e.g., Copelatus, Laccodytes, Desmopachria, various Bidessini), Noteridae (Notomicrus), Elmidae (many genera, e. g., Cylloepus, Gyrelmis, Heterelmis, Austrelmis), Hydraenidae (Adelphydraena orchymonti Perkins, Hydraena spp.), Hydrophilidae (Anacaena, Berosus, Chaetarthria, Notionotus, Oocyclus, Phaenonotum), Hydroscaphidae (Scaphydra), Sphaeriusidae (Sphaerius), Torridincolidae (Hintonia). In captivity, beetles spent most of the time sitting still or crawling about on dead leaves. They were not observed feeding. Rarely beetles floated after becoming detached from the leaves and were observed walking along under the surface film as has been observed in Hydraenidae and some small hydrophilids. To submerge, the beetles turn over with the head directed downward and 'kick' (with an alternating leg motion as in walking) their way downward until finding a foothold on the substrate; if a beetle stopped 'kicking' on the way down, it would immediately float back to the surface, then start another descent. While the beetles were moving about, a silvery bubble-like area seemed to be present under each elytron. No mortality was noted until almost 4 months after capture, when two dead beetles were found. The last two specimens held in captivity lived in the glass finger bowl for 196 days. Morphology, Adults (Spangler & Steiner 2005; Beutel et al. 2005) (Figs. 3.2.2 A, 3.2.3, 3.2.4). Total length 0.85-0.9 mm, width c. 0.4 mm. Body form ovate, with indistinct pronoto-elytral angle; widest at basal l/4th of elytra; convex dorsally, slightly less so on ventral side. Colour dark brown to yellowish brown. Most surfaces bearing flat, wrinkled, unsclerotized setae arising from anterior side of large, shallow punctures. Head (Figs. 3.2.1, 3.2.2 A) prognathous, not shortened and broadened. Neck region moderately narrowed, round in cross section, covered by anterior collar of prothorax. Compound eyes broadly elliptic, moderately prominent; large ommatidia

Meruidae Spangler and Steiner 2005

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Fig. 3.2.1. Meruphyllisae, SEM micrograph, habitus. From: Spangler & Steiner 2005 (modified).

n u m b e r i n g 5 X 9 across axes of eye. Frons s m o o t h , slightly convex, w i t h reticulate microsculpture; laterally a n d basally w i t h scattered shallow p u n c tures a n d fine setae; frontoclypeal s u t u r e recognisable, nearly straight, n o t i n t e r r u p t e d medially. L a b r u m trapezoidal, w i t h scale-like microsculp t u r e ; connected w i t h nearly straight anterior clypeal m a r g i n by an internal m e m b r a n o u s fold; w i t h a shallow m e d i a n concavity covered by opposing, overlapping, s p a t u l a t e setae a n d w i t h t h r e e tapered setae o r i g i n a t i n g f r o m small pits lateral to base of spatulate setae; pair of small, flat, tooth-like setae i m b e d d e d in anterior m a r g i n of concavity. A n t e n n a (Fig. 3.2.3 A-C) longer t h a n h e a d , filif o r m , glabrous, 11-segmented; scapus very s h o r t a n d almost completely concealed; pedicellus almost globular, u r n - s h a p e d , b e a r i n g a few slender setae; a n t e n n o m e r e III as l o n g as pedicellus b u t m o r e narrow, nearly twice as l o n g as wide, subcylindrical, thickest t o w a r d apex; a n t e n n o m e r e s 4 - 1 0 of similar f o r m , b u t 5, 7 a n d 9 longer a n d thicker t h a n adjacent ones; a n t e n n o m e r e s 7 a n d 9 b e a r i n g a plate-like sensorial patch ventrally; a n t e n n o m e r e 11 longer t h a n preceding two comb i n e d , thickest at m i d d l e a n d obliquely t a p e r i n g to a narrow apex, w i t h two plate-like sensorial patches near apex on ventral side (Fig. 3.2.3 C). Mandibles asymmetrical (Fig. 3.2.4 A), nearly twice as long as wide, w i t h single, b l u n t , falcate, apical incisor a n d mesal terebral tooth; brushes of trichia, mola a n d retinaculum absent; left m a n d i b l e

slightly longer t h a n right, terebral tooth broadly angular; inferior terebral ridge w i t h one or two m i n u t e teeth; r i g h t mandible w i t h terebral tooth t r u n cate a n d closer to apex t h a n that of left mandible; basal face w i t h several small wrinkles. Maxilla (Figs. 3.2.3 E, F, 3.2.4 B) w i t h small and wedge-shaped cardo; mediostipes a n d lacinia fused; basistipes well developed, triangular, narrowing toward base of palp; lacinia stout, elongate, glabrous, w i t h a curved, robust apical spine and seven spines on inner side; galea palp-like, 2-segmented, shorter t h a n lacinia, w i t h apical s e g m e n t elongate, cylindrical, smoothly r o u n d e d at apex; palpifer n o t recognisable as separate element of maxilla; palpus 4-segmented; apical palpomere slightly longer t h a n combined length of segments 1 - 3 , w i t h sensory structures on distal part. S u b m e n t u m broad, w i t h distinct posterior tentorial pit at basolateral corner (Fig. 3.2.2 A); m e n t u m flat, s m o o t h , slightly m o r e t h a n half as wide as head, w i t h two long setae on each side at a b o u t middle; p r e m e n t u m recessed, inserted between anterolateral r o u n d e d lobes of m e n t u m ; base of palpiger w i t h single small seta; ligula moderately extended and r o u n d e d apicomedially, w i t h 1 2 - 1 4 apical setae (Fig. 3.2.3 D); labial palp 3 segmented; apical s e g m e n t swollen and slightly longer t h a n combined length of segments 1 a n d 2, w i t h apical sensory structures similar to those of maxillary palp (Fig. 3.2.3 E). Gular ridges high; t e n t o r i u m well developed except for reduced dorsal arms; l a m i n a t e n t o r i u m extensive, Y-shaped in cross section. Head musculature

Fig. 3.2.2. Meru phyllisae, SEM micrographs. A, head a n d prothorax, ventral view; B, pterothorax, ventral view; C, abdomen, ventral view. From: Beutel et al. 2006 (modified).

Meruidae Spangler and Steiner 2005

31

Fig. 3.2.3. Meruphyllisae, SEM micrographs. A, antenna, dorsal view; Β, antennomeres 6-8, with sensorial patch on segment 7; C, apex of antennomere 11, ventral view; D, labrum and mouthparts, frontal view; E, maxillary and labial palp; F, apex of maxillay palp. From: Spangler & Steiner 2005 (modified). well developed, not distinctly affected by miniaturisation. Prepharynx, pharynx and oesophagus extremely wide, round in cross section. Brain large in relation to head size; largest part located in posterior head region; circumoesophageal connectives short; suboesophageal ganglion enclosed by gular ridges. Cervical sclerites absent. Prothorax (Fig. 3.2.2 A) about 4/5 as wide as elytra. Pronotum widest at base, slightly wider than long; lateral margin sinuate, not beaded; anterior margin straight, anterior corners narrowly rounded; posterior corners angled; posterior margin nearly straight between corner and middle, tightly fitted against base of elytron and forming an angle of about 125° with other side at middle; disc evenly convex, with four shallow depressions on each side; coarsely, densely punctate; each puncture with a long, robust, flattened and wrinkled seta, pleated at base, arising from anterior edge of puncture and recumbent across it; length of seta about twice diameter of a puncture; scattered, hair-like setae also present, arising from smooth areas between large punctures. Thoracic ventrites with similar combination of setae and punctures, but larger and more sparsely placed laterally. Prosternum (Fig. 3.2.2 A) with notopleural and pleurosternal sutures visible; anterior prosternal margin evenly arcuate, with fine fringe of micro-setae along edge

and on each side with one large, anteriorly directed, club-shaped seta, with apex flattened and toothed, brush-like. Prosternal process very narrow between procoxae but wider and truncate behind them. Propleuron large, extending behind coxa. Procoxal cavity closed internally. Procoxa globular; trochanter smooth, unmodified. Musculature of prothorax normally developed. Mesoventrite fused with metaventrite and with lateral pleural parts; pentagonal groove present, partly covered by prosternal process. Mesocoxa globular. Musculature well developed. Scutellar shield not exposed. Elytra convex, twice as wide as long, dorsally with setae and punctures like those on pronotum (Spangler & Steiner 2005: fig. 5 A, B) in addition to rows of large, deep, strial punctures without setae; sutural stria absent and sutural edge not beaded; lateral margin arcuate in lateral view; epipleuron indistinct; inner surface smooth, laterally with large, locking pit at about mid length and round patch of granular surface (presumably for wing-folding) anterior to it (Spangler & Steiner 2005: fig. 5 C, D). Metaventrite fused with anepisternum and metacoxae, without discernable sutures; transverse ridge absent. Mesal wall of metacoxa extensively fused (intercoxal septum); metacoxal plates very reduced, not covering base of trochanter, with lateral margins indistinct anteriorly; median lamina of metacoxae

32

Rolf G. Beutel, Michael Balke and Warren E. Steiner

Fig. 3.2.5. Meru phyllisae, SEM micrographs. A, front and middle legs, posterior view; B, mesotibial spurs. From: Spangler & Steiner 2005 (modified).

Fig. 3.2.4. Meruphyllisae, mouthparts. A, mandibles; Β, maxilla. From: Spangler & Steiner 2005 (modified).

n o t distinctly separated f r o m lateral part; posterior m a r g i n s t r a i g h t , w i t h small m e d i a n e m a r g i n a t i o n a n d d e e p incisions for trochanteral articulations (Fig. 3.2.2 B); m e t a t r o c h a n t e r large, a b o u t twice size of pro- a n d m e s o t r o c h a n t e r , globular, swollen posteriorly; j u n c t i o n w i t h m e t a f e m u r oblique a n d sinuate. M e t a f u r c a simplified, Y-shaped, w i t h o u t lateral a r m s (Fig. 3.2.6 B); attached to extensive intercoxal s e p t u m a n d to m e t a v e n t r i t e for t w o - t h i r d s of its l e n g t h . M m . furcacoxalis anterior a n d posterior absent; large indirect flight muscles also reduced in b o t h specimens e x a m i n e d a n d large parts of m e t a t h o r a x filled w i t h f a t b o d y tissue. H i n d w i n g s d i m o r p h i c (3.2.6 A; Spangler & Steiner 2 0 0 5 : figs. 10-12), e i t h e r f u l l y developed w i t h v e n a t i o n moderately developed a n d distal half folded, or b r a c h y p t e r o u s , as l o n g as elytron, w i t h venation very reduced a n d distal half appeari n g s h r u n k e n ; m a r g i n s w i t h f r i n g e of l o n g hairs; m e d i a l b i n d i n g patch (katastigma) absent; oblong u m cell n o t discernable. Distal parts of legs (Fig. 3.2.5 A) w i t h f e m o r a s m o o t h , club-shaped,

widest at a b o u t apical 1/3, w i t h scattered sparse hair-like setae, some b r u s h - t i p p e d , a n d row of five to seven thick, branched setae on dorsal surface; m e t a f e m u r very slender basally, attached dorsolate r a l ^ to trochanter; distal two-thirds swollen; tibiae simple, straight, gradually t h i c k e n i n g f r o m base to near apex, w i t h sparse, t h i n setae, irregularly distributed, becoming m o r e s t o u t toward apex of tibia; each tibia bearing paired, toothed apical spurs; b o t h spurs t r i d e n t on pro- a n d mesotibia, one t r i d e n t and one laterally pectinate on metatibia (Fig. 3.2.5 B); tarsal f o r m u l a 5-5-5; last tarsomeres longer t h a n combined lengths of preceding two; tarsomeres 1 and 2 each w i t h f o u r stout, ventral setae, 3 a n d 4 w i t h two setae, and 5 w i t h one seta; h i n d tarsus w i t h basal tarsomere longest; l e n g t h equals combined l e n g t h of tarsomeres 2 - 4 ; tarsal claws large, pectinate, w i t h 4 - 5 teeth; e m p o d i u m small, s m o o t h , pad-like. Proventriculus present (Spangler & Steiner 2005: fig. 16), w i t h f o u r larger plates s m o o t h , each w i t h single m e d i a n a c u m i n a t e a n d serrulate flap seemingly capable of folding against base; smaller f o u r plates vase-shaped in lateral view, s m o o t h , w i t h fringe of fine hairs. A b d o m e n w i t h six visible ventrites (morphological sternites II-VII) (Fig. 3.2.2 B, C). Ventrites 1 , 2 a n d 3 completely f u s e d (=sternites II-IV),

Meruidae Spangler and Steiner 2005

33

A

0.4 mm

0.2 mm

D

0.1 mm 0.1 mm Fig. 3.2.6. Meru phyllisae, A, hind wing; B, metafurca; & Steiner 2005 (modified).

male genitalia; D, female genitalia. From: Spangler

w i t h sutures indistinct (positions m a r k e d by row of setae); first ventrite deeply a n d broadly divided medially for reception of metacoxal processes, internally w i t h heavily sclerotized transverse sept u m along posterior m a r g i n . Posterior m a r g i n s of ventrites 1 - 5 b e a r i n g a comb-like row of robust,

flattened a n d grooved setae a n d fewer hair-like setae (Fig. 3.2.2 B, C). Ventrites 1 a n d 2 w i t h p u n c tures b e a r i n g r o b u s t wrinkled setae scattered over surface; flat pleated setae on ventrites 3 - 5 w i t h o u t associated p u n c t u r e s (unlike pleated setae on other body regions). Apical ventrite (sternite VII) wider

34

Rolf G. Beutel, Michael Balke and Warren E. Steiner

than long, apical margin rounded; surface bearing irregular transverse band of flattened and grooved setae and fewer hair-like setae (Fig. 3.2.2 C). Male genitalia (Fig. 3.2.6 C) asymmetric; median lobe with curved, tubular base, becoming more straight, dorsoventrally flattened and tapering toward apex; parameres vestigial, scale-like; left paramere more rounded, laterally positioned at base of median lobe; right paramere smaller, more narrow, positioned on ventral side of base of median lobe; aedeagus with only slight torsion in repose; ring sclerite (sternum IX) complete. Female genitalia lacking sclerotized structures except for reduced short gonocoxostyli, each bearing a few small apical setae (Fig. 3.2.6 B).

The clade comprising Meruidae and Noteridae is suggested by the absence of the transverse ridge of the metaventrite, the fusion of abdominal sternites III and IV, the shape of the strongly asymmetric parameres, and the enlargement of antennomeres 5, 7, and 9. It is also supported by molecular data (Balke etal. 2008). The Meruidae + Noteridae clade is the sister group of the remaining Dytiscoidea.

Larvae, pupae and eggs. Larvae recently discovered (A. Short, pers. comm.), but not described yet. Other immature stages unknown.

Literature

Phylogeny and Taxonomy (Spangler & Steiner 2005; Beutel etal. 2006; Balke etal. 2008). The results of comprehensive cladistic analyses of morphological (Beutel etal 2006) and molecular data (Balke etal. 2008) clearly demonstrate that Meruidae belong to Dytiscoidea. Autapomophic conditions of this lineage including Meru are the extensively fused metacoxae (large intercoxal septum), the origin of the metafurca from the intercoxal septum, and the loss of Mm furcacoxales anterior and posterior.

Acknowledgements Hans Fery critically read and commented on the manuscript. This is gratefully acknowledged.

Balke, M., Ribera, I., Beutel, R., Viloria, Α., Garcia, M. & Vogler, A. P. (2008): Systematic placement of the recently discovered beetle family Meruidae (Coleoptera: Dytiscoidea) based on molecular data. - Zoologica Scripta 37:647-650. Beutel, R. G., Balke, M. & Steiner, W. E. (2006): On the systematic position of Meruidae (Coleoptera, Adephaga) and the phylogeny of the smaller hydradephagan families. - Cladistics 22: 102-131. Spangler, P. J. & Steiner, W. E., Jr. (2005): A new aquatic beetle family, Meruidae, from Venezuela. -Systematic Entomology 30:339-357.

4. Elateroidea

Introduction, Phylogeny John F. Lawrence, Ladislav Bocak, Milada Bocakova, Rolf G. Beutel and Jyrki Muona The constitution of Elateriformia has varied over time, as discussed in detail by Beutel & Leschen (2005) (see 1-14). The series was first proposed by Crowson (1960) for his Dascilliformia (Crowson 1955) minus the family Dascillidae (which was combined with Scarabaeoidea to form the series Scarabaeiformia) and the families Eucinetidae, Clambidae and Scirtidae, which were placed in a superfamily Eucinetoidea. This classification was also used in Crowson (1981) except that a series Eucinetiformia was recognized and Rhipiceridae was added to Scarabaeiformia-Dascilloidea, based on Crowson (1971). In all of Crowson's classifications, the superfamily Elateroidea included Perothopidae, Eucnemidae, Throscidae, Cebrionidae, Elateridae, and Cerophytidae, although the last was omitted in error f r o m the 1981 work. Lawrence & Newton (1982) followed Crowson's classification in most respects. They did not define series as such, b u t considered all Eucinetiformia, Scarabaeiformia and Elateriformia as belonging to an "Elateriform lineage". Although Scarabaeoidea was tentatively included in this "lineage", some d o u b t was expressed about the relationship of the group to Dascilloidea. The families normally included in Elateroidea and Cantharoidea, plus the Artematopodidae and Brachypsectridae were considered to form a monophyletic group. In the first cladistic analysis of Elateriformia, Lawrence (1988) excluded Scarabaeoidea altogether, while Eucinetoidea were included as an outgroup in some analyses. The monophyly of Elateroidea + Cantharoidea + Artematopodidae + Brachypsectridae was confirmed in analyses based on both adult and larval characters, with the family Rhinorhipidae (known f r o m adult characters only) at the base of this clade. Elateroidea was restricted by Crowson (1955) to those taxa the adults of which have more or less rounded procoxae with concealed trochantins, no transverse metakatepisternal suture, contiguous metacoxae, h i n d wing with an apically truncate wedge cell, acutely projecting h i n d pronotal angles, head without a distinct frontoclypeal suture, trilobate aedeagus with freely articulated parameres, and 4 free Malpighian tubules, while larvae lack a free l a b r u m or epicranial stem and have simple, non-channeled mandibles. Artematopodidae (then in Dryopoidea) were considered to be separable f r o m elateroids on little more than exposed trochantins in the adult and a free

labrum in the larva, and Brachypsectridae were considered to be even more difficult to separate on adult features. The superfamily Cantharoidea was considered to be the most likely group to be merged with Elateroidea. Lawrence & Newton (1982) followed Crowson in considering Artematopodidae, Brachypsectridae, Elateroidea and Cantharoidea to form a monophylum, and Lawrence (1988) formally recognized an expanded Elateroidea to include all of these groups. The position of Rhinorhipus Lawrence at the base of the elateroid clade was considered to be tentative because of lack of information on the larva, combined with the fact that there are six free Malpighian tubules (instead of four as in all other members of the group). Furthermore, in cladograms produced by Lawrence etal. (1995), Rhinorhipus usually formed a clade with Dascillus Latreille (Dascillidae), Sandalus Knoch (Rhipiceridae) and Dystaxia LeConte (Buprestidae or Schizopodidae) and was never placed within the elateroid-cantharoid group. In cladograms produced by Beutel (1995) and based on larval characters, Elateroidea (sensu lato) was always monophyletic, b u t this was true of neither Elateroidea (sensu stricto) nor Cantharoidea. M o s t c a n t h a r o i d f a m i l i e s p l u s Brachypsect r i d a e formed a clade sister to Cerophytidae + Throscidae + Eucnemidae, while Cantharidae formed a clade with Artematopodidae and Elateridae. The non-monophyly of the Cantharoidea was also supported by Bocakova et al. (2007) in cladograms based on nuclear and mitochondrial gene sequences. While Elateroidea (sensu lato) was strongly supported in all cladograms, the soft-bodied groups usually placed in Cantharoidea never formed a monophyletic group. The major clusters were formed by 1) Lampyridae (including Ototretinae) + Cantharidae, 2) Elateridae (including Drilidae and usually Omalisidae) + Phengodidae (including Rhagophthalmidae), 3) Lycidae and 4) Eucnemidae. The positions of the genera Drilonius Kiegenwetter, Telegeusis Horn, Trixagus Kugelann and sometimes Omalisus Geoffroy varied with type of alignment and analysis: 1) Drilonius, Telegeusis and Trixagus formed a clade with Chelonariidae and outside Elateroidea; 2) Drilonius and Telegeusis formed a clade sister to Elateroidea and Trixagus was sister to Elateroidea minus Drilonius and Telegeusis; 3) Drilonius was in Eucnemidae, Telegeusis sister to Elateroidea minus Eucnemidae, and Trixagus sister to Lycidae; or 4) Drilonius and Telegeusis formed a clade sister to remaining elateroids, and Trixagus and Omalisus formed a clade within Eucnemidae. Similar results were published by Sagegami-Oba etal. (2007) and Bocak etal. (2008). The Elateroidea, as here delimited, exhibit several major evolutionary trends which deserve

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John F. Lawrence, Ladislav Bocak, Milada Bocakova, Rolf G. Beutel and Jyrki Muona

f u r t h e r mention: 1) development of a type of defensive behavior k n o w n as "clicking" in adults of the families Cerophytidae, Eucnemidae, Throscidae and Elateridae, 2) reduction in sclerotization of the cuticle, often accompanied by chemical defense mechanisms and aposematic color patterns in adults of various families formerly included in Cantharoidea, and 3) retention of larval features (neoteny) in adults of at least some of these families; 4) the evolution of bioluminescence in both adults and larvae; 5) the occurrence of an elateroid type of ecdysis associated with biforous spiracles and the loss of the spiracular closing apparatus in larvae; and 6) consolidation of the larval maxillae and labium to form a maxillolabial complex. The cuticular and muscular modifications which make the clicking maneuver possible have been discussed by Evans (1972, 1973) for Elateridae, b u t precursors of these conditions are exhibited by members of various families of Dascilloidea, Buprestoidea and Byrrhoidea. The evolution of a pro-mesothoracic interlocking device involving projections and concavities or crenulate edges at the posterior end of the prothorax, anterior ends of the elytra, scutellum and/or mesanepisterna, combined with a mesoventral cavity for reception of the prosternal process, allow these beetles to combine mobility with structural integrity, by the unlocking or locking of this device. The transformation of this condition to form the clicking mechanism involves the enlargement of the prothorax, increase in the mass of the M4 muscle (Larsen 1966), reduction of the size of the exposed portion of the procoxa, enclosure of the trochantin and (except in Cerophytidae) its fusion to the n o t u m , enlargem e n t and deepening of the mesoventral cavity combined with the formation of a prosternal rest and an oblique slide at the anterior end of the cavity. Based on the topology given by Bocakova et al. (2007), this condition could have arisen independently f r o m three to five times in the Elateroidea. Vahtera et al. (2009), however, suggested that, given the complexity of the clicking mechanism, it could have evolved at the base of the elateroid clade and been subsequently lost on n u m e r o u s occasions, usually in association with the development of soft-bodiedness (see below). Cantharoidea were defined mainly on the shared morphological traits resulting f r o m soft-bodiedness (generally reduced body sclerotisation and a soft, flexible abdomen with extensive intersegmental membranes reminiscent of those in the larvae). The molecular phylogenies mentioned above (Bocakova et al. 2007; Sagegami-Oba et al. 2007; Bocak et al. 2008) rejected monophyly of Cantharoidea and proposed multiple origin of both softbodiedness and probably closely related female neoteny of some groups within broadly defined Elateroidea. The hypothesis of frequent shifts to neotenic development opens a possibility that the morphological disparity suggesting establishment of families like Drilidae or some subfamilies like

Leptolycinae (Lycidae) is not a result of the long evolutionary history, b u t a consequence of relatively recent modified function of the endocrine system. These events potentially led to homoplasious modifications of morphology. The resulting similarity of soft-bodied or neotenic lineages is therefore difficult to interpret in morphology based analyses. Crowson (1972) postulated that some neotenous groups, specifically the Southeast Asian lycid genera Duliticola Mjöberg and Lyropaeus Waterhouse, are members of primitively neotenous lineages and that fully metamorphosed winged forms re-developed f r o m neotenic ancestors. Similar scenarios of evolutionary 're-imaginalisation' were proposed for Lycidae by Kazantsev (2005), and equally for the closely related Lampyridae by Cicero (1988). Bocak et al. (2008) hypothesized that soft-bodiedness represents a first level of incomplete metamorphosis. Soft-bodied adults of both sexes are k n o w n in Telegeusidae, Omethidae, Cantharidae, Lycidae, Lampyridae, Phengodidae, Rhagophthalmidae, Drilidae, and Omalisidae. Some adult females within these families are neotenic, i. e., they maintain apparently juvenile features resulting in incomplete metamorphosis and, in extreme cases, the lack of adult stages. The neotenic development of females is obligatory in all Omalisidae, Drilidae, Phengodidae and Rhagophthalmidae, and in many lineages of Lampyridae and Lycidae. The modifications include females with vestigial wings, b u t adult-like thorax (Omalisidae, Lampyridae part), wingless females (Lampyridae part) or females with only mouthparts and head adult-like (Drilidae, Lampyridae part). Lineages affected by neoteny to the highest degree are found in Lycidae where females lack both pupal and adult stages and retain a larvae-like morphology after the last ecdysis (Wong 1996). Some neotenic lycids reach body sizes of five centimeters and more and are frequently referred to as 'trilobite larvae' due to their appearance (Gravely 1915; Mjöberg 1925). The corresponding males are regularly fully metamorphosed and only seldom brachelytrous (Alyculus Kazantsev in Lycidae and Phosphaenus Laporte in Lampyridae). Several elateroid groups (Lampyridae, Phengodidae, Rhagophthalmidae and two independent groups of Elateridae (Agrypninae: Pyrophorini and Thylacosterninae: Balgus Fleutiaux) are known for their bioluminescence. Previous morphological studies (Crowson 1972, Beutel 1995) often suggested close relationships of cantharoid luminescent lineages (Lampyridae, Phengodidae). Latest molecular analyses (Bocakova et al. 2007; Sagegami-Oba et al. 2007) showed that bioluminescent groups have arisen at least four times in Elateroidea. Likewise, recent morphological analysis (Branham and Wenzel 2001,2003) supported several independent originations of bioluminescence in Elateroidea. Although superficially similar, molecular conclusions differ substantially. While morphological study separates Rhagophthalmidae from Phengodidae, and Drilaster Kiesenwetter and Stenocladius Fairmaire from Lampyridae,

Elateroidea

neither of these conclusions were confirmed in the molecular studies, hence upholding the traditional view of the constitution of Phengodidae and Lampyridae (Crowson 1972; Lawrence et al. 1995). Conversely, latest molecular analyses f o u n d cantharoid luminescent groups Lampyridae and Phengodidae deeply separated which is also supported by the structural and biochemical differences of the luciferases in either group (Viviani 2002). Vahtera et al. (2009) presented an hypothesis connecting the clicking m e c h a n i s m w i t h the evolution of bioluminescence. If the clicking m e c h a n i s m evolved in ancestors of t h e entire elateroid complex, t h e b i f u n c t i o n a l role of t h e pre-luciferase e n z y m e in c o m b i n a t i o n w i t h t h e high-energy d e m a n d of t h e pronotal muscle were t h e preadaptive features for t h e luminescence to evolve in the p r o t h o r a x . Lineages evolving away f r o m the compact elateridtype body s t r u c t u r e retained t h e predisposition for luminescence, once a suitable luciferin was available. T h e sources of luciferin type c o m p o u n d s in beetles, w h e t h e r of symbiotic origin or not, facilitated t h e p r o n o t a l l i g h t spots at t h e m u s c u l a r a t t a c h m e n t points as well as t h e fat b o d y region. This scenario predicts t h a t t h e source for luciferin is m o s t likely external a n d after b e c o m i n g available for any elateroid clade could be picked u p repeatedly. It also explains why this f e a t u r e is restricted to this one g r o u p beetles - t h e clicking m e c h a n i s m b e i n g u n i q u e w i t h i n beetles. Larval h e a d structures of Elateroidea are q u i t e characteristic, even t h o u g h , as p o i n t e d o u t in Beutel (1995), several derived features are also f o u n d in larvae of all or m o s t g r o u p s presently assigned to Byrrhoidea (see 1 - 2 ) . A tendency to concentrate or reduce t h e s t e m m a t a is f o u n d in b o t h lineages. Well separated s t e m m a t a occur in so m e g r o u p s of Byrrhoidea (e. g., Byrrhidae, Dryopidae, Heteroceridae) b u t in others (e. g., Psephenidae, Ptilodactylidae) they f o r m t i g h t clusters a n d in Eulichadidae there is a single large lens b e n e a t h which are two to five p i g m e n t spots. In Elateroidea there is never m o r e t h a n a single s t e m m a on each side. As in all Byrrhoidea, elateroid larvae lack a basal m a n d i b u l a r mola, a n d as in Byrrhoidea (excluding Byrrhidae) the head is distinctly prognathous. Both conditions have evolved i n d e p e n d e n t l y in d i f f e r e n t lineages of Coleoptera, notably in g r o u p s w i t h predacious larvae (e. g., Adephaga, Hydrophiloidea, Cleroidea [see 1 - 7 , 1 - 1 0 , 2-9]). A characteristic f e a t u r e f o u n d in larvae of Elateroidea (and Byrrhoidea excluding Byrrhidae, some Ptilodactylidae a n d Eulichadidae) is a maxillolabial complex, w i t h closely connected l a b i u m a n d maxillae (Beutel 1995). T h e ventral m o u t h p a r t s are moved only as a structural u n i t vertically. T h e extrinsic tentoriomaxillary muscles are vertically arranged. Similar conditions have evolved i n d e p e n d e n t l y in Cleroidea a n d in s o m e supposedly related g r o u p s of Cucujoidea (Beutel & Slipinski 2001). As in m o s t byrrhoid g r o u p s (excluding Byrrhidae, Ptilodactylidae, Eulichadidae a n d Callirhipidae) t h e t e n t o r i u m of elateroid

37 larvae is strongly modified, w i t h posterior a r m s very strongly developed, cranially directed a n d completely detached f r o m t h e tentorial bridge. T h e dorsal a n d anterior parts of t h e t e n t o r i u m are reduced. Interestingly, again a similar condition is f o u n d in cleroid larvae a n d in some g r o u p s of Cucujoidea (Beutel & Slipinski 2001). Apparently this condition is linked w i t h t h e f o r m a t i o n of a maxillolabial complex. A set of features distinctly s e p a r a t i n g Elateroidea f r o m Byrrhoidea is the presence of a strongly developed lateral tentoriohypopharyngeal muscle, a dense, preoral filter f o r m e d by l o n g microtrichia, the immobilisation of t h e l a b r u m , a n d t h e loss of t h e labral muscles. A l a b r u m separated f r o m t h e clypeal region is preserved only in A r t e m a t o p o d i d a e a n d Brachypsectridae. T h e preoral filter is apparently an a d a p t a t i o n to liquid feeding. A similar condition has evolved in Carabidae a n d Histeroidea (Beutel 1993, 1999). U n u s u a l modifications of t h e m a n d i b u l a r apparatus are characteristic for larvae of m o s t E u c n e m i d a e a n d Throscidae, w h e r e mandibles may be fixed or e x o d o n t . A n o t h e r specific modification is t h e presence of m a n d i b u l a r sucking channels occurring in Brachypsectridae, Lampyridae, a n d a few other g r o u p s . An u n u s u a l f e a t u r e apparently linked w i t h highly specialized liquid feeding habits is t h e origin of very strongly developed extrinsic maxillary muscles of t h e sclerotized ventral wall of the h y p o p h a r y n x . A s o m e w h a t similar condition has evolved in Cleroidea a n d s o m e cucujoid groups, w h e r e an anterior b u n d l e of M. tentoriostipitalis originates f r o m the ventral prepharyngeal wall or f r o m the posteriormost h y p o p h a r y n x (Beutel & Slipinski 2001).

Literature Beutel, R. G. (1993): Phylogenetic analysis of Adephaga (Coleoptera) based on characters of the larval head. -Systematic Entomology 18:127-147. - (1995): Phylogenetic analysis of Elateriformia (Coleoptera: Polyphaga) based on larval characters. - journal of Zoological Systematics and Evolutionary Research 33:145-171. - (1999): Morphology and evolution of the larval head of Hydrophiloidea and Histeroidea (Coleoptera: Staphyliniformia). - Tijdschriftvoor Entomologie 142: 9 - 3 0 . Beutel, R. G. & Leschen, R. A. B. (2005): 14. Elateriformia Crowson, 1960. Introduction, Phylogeny. Pp. 427-429 in Handbuch der Zoologie/Handbook of Zoology. Band/Volume IV Arthropoda: Insecta Teilband/ Part 38. Coleoptera, Beetles. Volume 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). W. DeGruyter, Berlin. Beutel, R. G. & Slipinski, S. A. (2001): Comparative study of larval head structures of Sphindidae and Protocucujidae (Cucujoidea, Coleoptera). - European Journal ofEntomology 98: 219-232. Bocak, L., Bocakova, M., H u n t , T. & Vogler, A. P. (2008): Multiple ancient origins of neoteny in Lycidae

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38 (Coleoptera): consequences for ecology and macroevolution. - Proceedings of the Royal Society B, 275: 2015-2023. Bocakova, M., Bocak, L., H u n t , T., Teraväinen, Μ. & Vogler, A. P. (2007): Molecular phylogenetics of Elateriformia (Coleoptera): evolution of bioluminescence and neoteny. - Cladistics 23:477-496. Branham, M. A. & Wenzel, J. W. (2001): The evolution of bioluminescence in cantharoids (Coleoptera: Elateroidea). - Florida Entomologist 8 4 : 5 6 5 - 5 8 6 . - (2003): The origin of photic behavior and the evolution of sexual communication in fireflies (Coleoptera: Lampyridae). - Cladistics 19:1-22. Cicero, J. M. (1988): Ontophylogenetics of cantharoid larviforms (Coleoptera: Cantharoidea). Coleopterists Bulletin 4 2 : 1 0 5 - 1 5 1 . Crowson, R. A. (1955): The Natural Classification of the Families of Coleoptera. 187 pp. Nathaniel Lloyd, London. - (1960): The phylogeny of Coleoptera. - Annual Review ofEntomology 5 : 1 1 1 - 1 3 4 . - (1971): Observations on the superfamily Dascilloidea (Coleoptera: Polyphaga), with the inclusion of Karumiidae and Rhipiceridae. - Zoologicaljournal of the Linnean Society 5 0 : 1 1 - 1 9 . - (1972): A review of the classification of Cantharoidea (Coleoptera), with the definition of two new families, Cneoglossidae and Omethidae. - Revista de la Universidad de Madrid 2 1 : 3 5 - 7 7 . - (1981): The Biology of Coleoptera. 802 pp. John Murray, London. Evans, Μ. E. G. (1972): The j u m p of the click beetle (Coleoptera, Elateridae) - a preliminary study. -Journal ofZoology, London 167:319-336. - (1973): The j u m p of the click beetle (Coleoptera: Elateridae) - energetics and mechanics. -Journal of Zoology, London 1 6 9 : 1 8 1 - 1 9 4 . Gravely, F. H. (1915): The larvae and pupae of some beetles f r o m Cochin. Records of the Indian Museum 11:353-366. Kazantsev, S. V. (2005): Morphology of Lycidae with some considerations on evolution of the Coleoptera. Elytron 1 9 : 4 9 - 2 2 6 . Larsen, O. (1966): On the morphology and function of the locomotor organs of the Gyrinidae and other Coleoptera. - Opuscula Entomologica Supplementum 30:1-242. Lawrence, J. F. (1988): Rhinorhipidae, a new beetle family from Australia, with comments on the phylogeny of the Elateriformia. Invertebrate Taxonomy 2 : 1 - 5 3 . Lawrence, J. F. & Newton, A. F., Jr. (1982): Evolution and classification of beetles. - Annual Review ofEcology and Systematics 13: 261-290. Lawrence,J.F.,Nikitsky,N.B.,Kirejtshuk,A.G.(1995): Phylogenetic position of Decliniidae (Coleoptera: Scirtoidea) and comments on the classification of Elateriformia (sensu lato), pp. 375-410. In J. Pakaluk and S. A. Slipiriski (eds.), Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. M u z e u m i Instytut Zoologii Polska Akademia Nauk, Warsaw, Poland. Mjöberg, Ε. (1925): The mystery of the so called "trilobite larvae" or "Perty's larvae" definitely solved. Psyche 3 2 : 1 1 9 - 1 5 7 .

Sagegami-Oba, R., Takahashi, N., Oba, Y. (2007): The evolutionary process of bioluminescence and aposematism in cantharoid beetles (Coleoptera: Elateroidea) inferred by the analysis of 18S ribosomal DNA. Gene 400 (1-2): 104-113. Vahtera, V., Muona, J., Stahls, G. & Lawrence, J. F. (2009): The phylogeny of Thylacosterninae beetles (Coleoptera, Elateridae). - Cladistics 2 5 : 1 4 7 - 1 6 0 . Viviani, V. R. (2002): The origin, diversity, and structure function relationships of insect luciferases. Cellular and Molecular Life Sciences 5 9 : 1 8 3 3 - 1 8 5 0 .

4.1. Rhinorhipidae Lawrence, 1988 John F. Lawrence

Distribution. Rhinorhipus tamborinensis Lawrence h a s b e e n collected in a f e w localities in s o u t h e r n Q u e e n s l a n d , Australia, all at h i g h e r elevations in t h e vicinity of closed forest. It is likely t h a t t h e species also occurs in m o n t a n e regions in n o r t h e r n N e w S o u t h Wales.

Biology and Ecology. T h e largest series of a d u l t s were collected d u r i n g t h e day o n leaf surfaces of a n i n t r o d u c e d weed, Ageratina adenophora (Asteraceae), at t h e e d g e b e t w e e n r a i n f o r e s t a n d cleared areas. U n f o r t u n a t e l y , this site h a s n o w b e c o m e a s u b u r b a n d recent collecting e x p e d i t i o n s h a v e failed to

Fig. 4.1.1. Rhinorhipus tamborinensis Lawrence, adult male, dorsal (from Lawrence 1988; © CSIRO Australia).

Rhinorhipidae Lawrence, 1988

Fig. 4.1.2. Rhinorhipus taniborinensis Lawrence, adult female, dorsal (from Lawrence 1988; © CSIRO Australia).

produce more specimens. At another locality, a few beetles were found in an open area on low vegetation bordering a creek. When disturbed, the beetles exhibited a death-feigning reaction, dropping to the ground. Males greatly o u t n u m b e r e d females in this habitat. It is likely that these clearings were the sites of mating aggregations and that the beetles flew to t h e m from within the rainforest. Red m u d was present on a n u m b e r of the specimens, which suggests that they either emerged f r o m the soil after eclosion or sheltered there. The structure of the metacoxae and h i n d legs also suggests fossorial habits. The ovipositor is relatively unspecialized, so it is unlikely that the eggs are embedded in plant tissue or placed deep in soil. One female laid several eggs in the laboratory, b u t none of t h e m hatched. Morphology, Adults (Figs. 4.1.1-3). Length 5 - 8 . 5 m m . Body about 3 times as long as wide; slightly flattened above b u t moderately convex below. Heavily sclerotized and clothed with relatively stout and somewhat flattened, decumbent hairs. Head longer than wide, strongly declined, abruptly constricted immediately behind eyes, so that no temples are present. With very short, median occipital endocarina b u t no transverse line. Eyes moderately large, protuberant, more or less circular, finely faceted, without interfacetal setae; o m m a t i d i u m of exocone type with thick cornea and clear zone narrow or absent (Caveney 1986). Antennal insertions exposed, located in large, oblique, slightly raised circular fossae; subantennal grooves absent. Frontoclypeal region deflexed, strongly narrowed anteriorly, forming an elongate clypeal lobe which is apically rounded; frontoclypeal suture absent. Labrum elongate and slender, membranous, almost completely concealed by clypeus. Antennae 11-segmented, filiform to very slightly subserrate in both sexes, m u c h longer in male than in female; sensory elements consisting of 2 types of trichoid sensilla evenly distributed along anterior edges of segments 4 - 1 1 . Mandibles elongate, slightly and gradually curved mesally, unidentate, with simple, carinate incisor edge; deep, setose, dorsal cavity present at base (Fig. 4.1.3 B); mola absent; prostheca represented by short brush of hairs. Galea and lacinia reduced,

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membranous, densely setose; galea 1-segmented, truncate; laciniasubacute; apical palpomere slightly expanded near middle, apically truncate. Subment u m very short and broad; m e n t u m subtrapezoidal, biexcavate apically, with median tooth; ligula reduced, bilobed, setose; labial palps approximate; apical palpomere slightly expanded near middle, apically truncate. Hypopharyngeal and epipharyngeal surfaces clothed with distally inclined hairs. Subgenal ridges absent. Gular sutures moderately widely separated; pregular area subequal in length to gula, slightly concave. Corpotentorium and medially connected laminatentoria fused, f o r m i n g single, broad bridge. Cervical sclerites small. P r o n o t u m about as long as wide, distinctly narrower than basal width of combined elytra, abruptly constricted at anterior third, with strongly produced and rounded anterior edge partly concealing head; lateral carinae absent; trisinuate posterior edge lacking margin or crenulations; promesothoracic interlocking device highly reduced, without lateral tubercles; sides of p r o n o t u m obliquely vertical, each with narrowly elongate, deep, oblique cavity at anterior fourth; posterior angles blunt and rounded, not produced, each one with smaller, deep cavity in front of it. Prosternum in front of coxae shorter than mid length of procoxal cavity, obliquely elevated, with strongly concave anterior edge; prosternal process long, narrow and slightly curved, narrowly rounded at apex, which fits into cavity on mesoventrite. Notosternal suture complete. Procoxae conical and projecting well below prosternum, without plates or concealed lateral extensions. Trochantin exposed, closely adpressed to sternum and only slightly movable (Fig. 4.1.3 A). Endopleuron free, with narrow basal stalk. Procoxal cavities widely open externally and internally; notal projections barely indicated. Scutellar shield relatively small, subtriangular, with base abruptly elevated and simple. Elytra about twice as long as combined width and 3 times as long as pronotum; more or less parallel-sided, with narrowly rounded apices; sutural flange deflected beyond apical fourth; each elytron with twelve more or less regular rows of deep, rounded, window punctures, in between which are rows of setiferous punctures; scutellary striole and apical interlocking tongue absent; epipleura narrow, weakly developed, not well marked b u t more or less complete. Mesoventrite about half as long as wide, narrowed and truncate anteriorly; on same plane as metaventrite, separated by complete sutures from mesanepisterna, which are separated from one another; discrimen absent; mesoventral cavity shallow, formed between paired, oblique carinae, which form mesal edges of procoxal rests, formed primarily by mesanepisterna. Mesanepisterna subtriangular, partly divided by oblique ridge anterior to which is major portion of concave procoxal rest. Mesepimeron smaller, partly fused to mesanepisternum, the pleural suture partly effaced at mesal end. Mesocoxa conical and projecting, without plate; with broadly exposed trochantin

40

John F.Lawrence

Fig. 4.1.3. Rhinorhipus tamborinensis Lawrence, adult features: A, prothorax, ventrolateral, showing coxal articular region and protrochantin abutting sternum; B, frontoclypeal region and base of left mandible, showing setose cavity; C, male, ventrolateral, with legs removed; D, male abdomen, dorsolateral, with pregenital sclerites extruded; E, male hind wing; F, aedeagus, ventral view; G, mesal portions of metacoxae and attached metendosternite, posterodorsal view; H-I, female abdominal apex: H, dorsal, I, ventral (all from Lawrence 1988; © CSIRO Australia).

41

Rhinorhipidae Lawrence, 1988

and weakly developed secondary condyle (fitting into sides of metaventral process). Mesocoxal cavities narrowly separated, internally contiguous, about as long as wide, open laterally (partly closed by mesanepisternum and mesepimeron), with solid joint between meso- and metathoracic portions of coxal cavities. Mesometaventral junction with deeply notched mesoventral process overlapping narrowly rounded metaventral process. Metaventrite about 0.75 times as long as wide, strongly convex, with short discrimen, and no transverse (katepisternal) suture. Exposed portion of metanepisternum broad, about 2.5 times as long as wide, subquadrate with straight inner and outer edges and with outer carina and groove which fit against elytral epipleuron. Metepimeron fused to metanepisternum; pleural suture absent posteriorly. Metacoxae large and contiguous, extending laterally to meet elytral epipleura; coxal plates weakly developed. Metendosternite (Fig. 4.1.3 G) with long narrow stalk, most of which is attached to the mesal internal edges of the metacoxae, and moderately long, curved lateral arms; anterior process (which bears anterior tendons) with two halves separated basally so that a foramen is formed between them; ventrolateral processes expanded apically to form flat muscle attachments. Hind wing (Fig. 4.1.3 E) about 2.6 times as long as wide; apical field less than 0.2 times wing length; transverse folds meeting at point between anterior, oblique, linear sclerite and radial cell, which is well developed and more than 4 times as long as wide, with inner posterobasal angle right or slightly obtuse; cross-vein r3 short and slightly oblique, partly fused at base with r4; RP with basal section long, extending almost to mid wing, and apical section absent; R-M loop relatively narrow; medial spur more or less straight and extending to wing margin; medial field with five free veins, wedge cell well developed, apically acute; anal notch absent; AP undivided. Trochanterofemoral joints on fore and mid legs slightly oblique with base of femur well separated from coxa; that on hind leg strongly oblique, with trochanter produced along inner edge of tibia; femora and tibiae tuberculate; outer edges of tibiae denticulate; tibial apex slightly expanded, bearing two distinct spurs; metatibial apex more strongly expanded in female; tarsi 5-5-5; segments simple, without membranous lobes or setose pads beneath; claws pectinate; empodium well developed, bi- or trisetose. Abdomen (Figs. 4.1.3 C, D) with five strongly convex ventrites, the first three of which are connate b u t separated by distinct sutures. Ventrite 1 (sternite III) deeply excavate for reception of metacoxae, with short triangular intercoxal process and distinct carina. Sternite VII (last ventrite) with narrow deep groove bordering free edge. Laterosternites well developed, those on segment VII delimited by distinct ridge; parasternites narrow, absent on posterior half of segment VII. Tergites all heavily pigmented, with microsetae sparsely and generally distributed around edges. Sternites V and

VI each with paired sternomedian convergent and sternolateral divergent muscles; tergosternal muscles particularly well developed on sternites III-V, and represented externally by paired, glabrous elevations of the cuticle. Functional spiracles present on abdominal segments I-VIII, located in pleural membrane. Anterior edge of sternite VIII in male with short, triangular apodeme. Sternite IX broadly rounded anteriorly, withoutspiculum gastrale. Sternite VIII in female without median strut (spiculum ventrale) or anterolateral struts. Tergite IX in male emarginate. X well developed and free. Aedeagus (Fig. 4.1.3 F) trilobate, symmetrical, with short phallobase, long, narrow parameres, and longitudinally divided penis, with very short paired, basal struts and no ventral sclerotized lobe. Internal male reproductive system with large spermatophoral glands and closely associated seminal vesicles, large, thin-walled accessory glands, and testes consisting of ten pedicillate sperm tubes. Ovipositor (Fig. 4.1.3 H-I) short and broad, with divided coxites and spindle-shaped styli; paraprocts slightly longer than gonocoxites and styli combined, with ventral longitudinal baculi; coxites each with short basal section, transverse baculum, and elongate, narrow apex. Proctiger well developed; ventral sclerite present. Internal female reproductive system with vagina expanded to form weakly defined bursa copulatrix at point of entry of common oviduct bursa without spines or sclerotized plates; spermatheca consisting of long narrow lightly sclerotized tube continuous with anterior end of genital chamber; spermathecal gland attached by short duct to base of spermatheca. Alimentary canal consisting of long narrow oesophagus, large, thin-walled crop, simple proventricular area, without spines or plates, bilobed ventriculus without gastric caecae or regenerative crypts, short, bulbous pylorus bearing six free Malpighian tubules, moderately short ileum and somewhat enlarged rectum. Ventral nerve cord with metathoracic and first abdominal ganglia fused; ganglia VII and VIII completely fused, and ganglion VI partly fused to VII-VIII; connectives paired. [Caveney 1986; Lawrence 1988; Lawrence & Britton 1991, 1994; Lawrence & Newton 1995; Lawrence etal. 1995,1999.]

Morphology, Larvae. Larvae unknown. Phylogeny and Taxonomy. In cladograms produced by Lawrence (1988) (87 adult and 25 larval characters; all taxa of Elateriformia sensu Crowson 1973; Eucinetoidea outgroup), Rhinorhipidae was placed at the base of one of three clades: a) one consisting of all Elateroidea in the broad sense (including Artematopodidae, Brachypsectridae and the cantharoids), b) one consisting of Callirhipidae, Eulichadidae, Ptilodactylidae, Cneoglossidae, Psephenidae and Chelonariidae (Psephenoidea), or c) one consisting of both of the previous groups. Cladograms produced by Lawrence et al. (1995) (99 adult and 23 larval characters; all taxa of

42 Elateriformia sensu Crowson 1973 plus Scirtoidea; outgroups from Archostemata, Myxophaga and/or Staphylinoidea) produced different results. In all cladograms, Rhinorhipidae was included in Elateriformia and in most it formed a monophyletic group with Dascillidae, Rhipiceridae and Buprestidae. Based on these conflicting results, the family was considered to be Elateriformia incertae sedis by Lawrence & Newton (1995). Perhaps the discovery of the larva of Rhinorhipus and/or the collection of fresh specimens for DNA analysis will shed some light on the phylogenetic relationships of the group.

John F.Lawrence

Lawrence, J. F., Hastings, A. M., Dallwitz, Μ. J., Paine, T. A. & Zürcher, Ε. J. (1999): Beetles of the World: A Key and Information System for Families and Subfamilies. CD-ROM, Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne.

4.2. Artematopodidae Lacordaire, 1857 John F. Lawrence

Acknowledgements CSIRO Australia is gratefully acknowledged for research support and for allowing me to use copyrighted figures. The following artists are acknowledged: S. Monteith (Figs. 4.1.1, 2, 3 C), S. P. Kim (Figs. 4.1.3 D-I). Thanks are also due to S. A. Slipinski and A. Seago for assistance in obtaining and scanning published illustrations and A. Hastings for preparation of plates.

Literature Caveney, S. (1986): The phylogenetic significance of ommatidium structure in the compound eyes of polyphagan beetles. - Canadian Journal of Zoology 64:1787-1891. Crowson, R. A. (1973): On a new superfamily Artematopoidea of polyphagan beetles, with the definition of two new fossil genera from the Baltic amber. -Journal of Natural History 7 : 2 2 5 - 2 3 8 . Hlavac, T. F. (1975): The prothorax of Coleoptera (except Bostrichiformia - Cucujiformia). -Bulletin of the Museum of Comparative Zoology 147(4): 137-183. Lawrence, J. F. (1988): Rhinorhipidae, a new beetle family from Australia, with comments on the phylogeny of the Elateriformia. - Invertebrate Taxonomy 2(1987): 1 - 5 3 . Lawrence, J. F. & Britton, Ε. B. (1991): Coleoptera (Beetles). Pp. 543-683 in CSIRO Division of Entomology (ed.) Insects of Australia: a Textbook for Students and Research Workers, Second Edition. Vol. 2. Melbourne University Press, Carlton, Victoria. - (1994): Australian Beetles, χ + 192 pp. Melbourne University Press, Carlton, Victoria. Lawrence, J. F. & Newton, A. F., Jr. (1995): Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names). Pp. 779-1006 in Pakaluk, J. & Slipinski, S. A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: Tapers Celebratingthe80thBirthdayofRoyA. Crowson. Muzeum i Instytut Zoologii PAN, Warsaw. Lawrence, J. F., Nikitsky, Ν. B. & Kirejtshuk, A. G. (1995): Phylogenetic position of Decliniidae (Coleoptera: Scirtoidea) and comments on the classification of Elateriformia (sensu lato). Pp. 3 7 5 - 4 1 0 in Pakaluk, J. & Slipinski, S. A. (eds.) Biology, Thylogeny, and Classification of Coleoptera: Tapers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warsaw.

Distribution. The genus Artematopus Perty includes about 45 described species and many undescribed occurring from Nicaragua to Brazil. The closely related genus Carcinognathus Kirsch contains a single species from Peru, but probably occurs more widely in northern South America. Electribius Crowson contains two extant species from Mexico and El Salvador and four additional fossil species from Baltic amber. Ctesibius eumolpoides Champion is known from the Mexican states of Nuevo Leon and San Luis Potosi, while the related Brevipogon confusus (Fall) occurs throughout the southern half of California. Allopogonia villosus (Horn) is restricted to extreme southern California and northern Baja California. The genus Macropogon Motschulsky occurs throughout the northern part of North America and in eastern Asia, while species of Eurypogon Motschulsky are known from North and Central America, Italy, Japan and Taiwan. The two monotypic genera Proartematopus Crowson and Electrapate IablokovKhnzorian are known only from Baltic amber.

Fig. 4.2.1. Brevipogon confusus (Fall), adult, dorsal (from Lawrence 2005; © CSIRO Australia); length = 4 mm.

Artematopodidae Lacordaire, 1857

[Iablokoff-Khnzorian 1962; Crowson 1973, Lawrence 1995; Hörnschemeyer 1998; Lawrence 2005.] Biology and Ecology. There seems little doubt that species of Eurypogon and Macropogon are closely associated with and probably feed on mosses growing on boulders. Cooper (1991) collected larvae and pupae of Macropogon sequoiae Hopping in mixed mats of moss, dominated by a species of Grimmia (Grimmiaceae) in the Sierra Nevada of California, while Macropogon piceus LeConte and Eurypogon harrisi (Westwood) are found in beds of a Paraleucobryum (Dicranaceae) on granitic boulders in New

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Hampshire. The latter were reared from early instars to adult on that moss alone. Gut contents are composed entirely of fine, unidentifiable particles, which appears to be due to the fact that the mouth cavity in these larvae are entirely blocked by dense patches of hairs, which filter out larger particles and create capillary forces. It is also likely that extra-oral digestion is involved. Crowson (1973) reported finding a teneral adult and full grown larva of Eurypogon niger (Melsheimer) under growths of a lichen (Umbillicara sp.) on granitic rocks in the Great Smoky Mountains of Tennessee. According to Champion (1897), adult Artematopus

ÄSiigS!

Fig. 4.2.2. Α-B, E-F, Eurypogon harrisi Westwood, larva: A, head, dorsal; B, head, anterior; E, abdominal apex, ventral; F, left mandible, ventral (© J. F. Lawrence); C, Macropogon sequoiae Hopping, larva, dorsolateral (from Cooper 1991; © K. W. Cooper), length = 1 3 mm; D, Artematopus sp. (Ecuador), adult prothorax, ventral, with right coxa and trochantinopleuron removed (© CSIRO Australia); G, Electribius crowsoni Lawrence, adult, dorsal (from Lawrence 1995; © CSIRO Australia and Muzeum i Instytut Zoologii Polska Akademia Nauk, Warsaw), length = 2.82 mm; H, Macropogon testaceipennis Motschulsky, adult, dorsal (© CSIRO Australia), length = 8 mm; I, Artematopus sp. (Panama), adult, dorsal (© CSIRO Australia), length = 9 mm.

44 "are found upon bushes in dry places, chiefly on the savannas of the 'tierra caliente'". Larvae of Artematopus discoidalis Pic were collected in "the superficial layer of forest soil, just below litter" at Salesopolis, Säo Paulo, Brazil, and reared through to the adult stage (Costa etal. 1985). In the laboratory, live termites and tenebrionid larvae offered as food were not attacked, but cut up insect pieces were readily eaten. Although this indicates possible carnivorous or saprophagous habits, it does not eliminate the possibility that Artematopus larvae are also bryophagous. Morphology, Adults (Figs. 4.2.1, 2 G, Η, I). Length 2 . 5 - 1 0 mm. Body about 1.76-2.89 times as long as wide. Slightly flattened to moderately convex. Densely clothed with decumbent and erect or suberect hairs. Head strongly declined, not abruptly constricted posteriorly. Posterior edge above occipital foramen distinctly bi-emarginate, forming pair of vertical impressions; without median endocarina. Transverse occipital ridge weakly developed or absent. Frontal region usually not or only slightly, gradually declined (moderately strongly declined in Electribius). Eyes well developed, slightly to strongly protuberant, entire, finely facetted, without or with very short, fine interfacetal setae; ommatidium of exocone type with reduced clear zone (Caveney 1986). Antennal insertions widely separated and exposed, located slightly in front of eyes, not raised. Subantennal groove usually absent or very weakly developed (well developed with deep, setose cavity in Electribius). Frontoclypeal suture absent; anterior clypeal margin emarginate or multidentate in some Neotropical species. Labrum free, well sclerotized and completely exposed, or sometimes lightly sclerotized, especially mesally, and partly concealed by clypeus; slightly to strongly transverse; anterior margin usually subtruncate, but sometimes cleft or emarginate. Antennae 11segmented, filiform or serrate, moderately long, sometimes very long in male; antennomeres 2 and 3 or 2 - 4 sometimes much shorter than following ones; antennomeres 4 - 1 1 clothed with erect hairs in males of most species. Mandible unidentate and acute; incisor edge usually with one or two subapical teeth (absent in Allopogonia Cockerell); mola and prostheca absent. Maxilla with galea and lacinia short, broad, lightly sclerotized and pubescent; lacinia without uncus; apical maxillary palpomere fusiform to subtriangular or securiform. Labium with mentum subtrapezoidal; ligula short, broad, lightly sclerotized, undivided or cleft; apical labial palpomere fusiform or apically widened. Subgenal ridges absent. Gular sutures well separated; gula longer than wide. Corpotentorium absent. Cervical sclerites usually well-developed. Pronotum about 0 . 4 5 - 0 . 7 times as long as wide; usually widest posteriorly; sides moderately straight, slightly sinuate or moderately to strongly curved anteriorly; base usually not to somewhat

John F.Lawrence

narrower than elytral bases; lateral carinae usually complete, sometimes vaguely indicated, simple, with or without raised margin; anterior angles right or rounded, not produced; posterior angles obtuse or right to acute, but not strongly produced; posterior edge usually bi- or trisinuate, simple, not margined; disc usually simple, sometimes with uneven glabrous elevations, with pair of basal cavities joined by transverse impression in Electribius. Promesothoracic interlocking mechanism weakly developed. Prosternum in front of coxae longer than mid length of procoxal cavity; slightly to strongly convex, with paired carinae (Fig. 4.2.2. D) (except in Allopogonia), with paired anterior cavities in Artematopus and Carcinognathus; anterior edge not produced, without mesal excavation. Prosternal process complete, parallel-sided, flat, overlapping mesoventrite; apex narrowly to broadly rounded; notosternal suture complete. Procoxae not or only slightly projecting, without concealed lateral extensions. Trochantins exposed, not closely adpressed to notum. Procoxal cavities strongly transverse, moderately narrowly separated, externally broadly open, without narrow lateral extensions, internally open. Scutellar shield well developed, anteriorly simple, abruptly elevated or not, posteriorly acute to broadly rounded. Elytra 1.36-2.47 times as long as combined width and 2 . 8 9 - 5 . 8 8 times as long as pronotum; punctation usually distinctly seriate (less so in Ctesibius), with eleven or twelve distinct puncture rows and no scutellary striole; sutural stria deeply impressed apically in Electribius·, apices conjointly rounded, each with ventrally interlocking tongue; epipleura very narrow, sometimes incomplete. Mesoventrite separated by complete sutures from mesanepisterna, which are well separated from one another; anterior edge on same plane as metaventrite, with paired, slightly to strongly declined procoxal rests; discrimen present but incomplete; mesoventral cavity small and shallow to large and moderately deep. Mesocoxae not projecting. Trochantins exposed. Mesocoxal cavities moderately to widely separated, slightly transverse, open laterally (partly closed by mesepimeron, occasionally with mesanepisternum); with distinct joint between meso- and metathoracic portions of coxal cavities. Mesometaventral junction usually complex, monocondylic or dicondylic. Metaventrite moderately to strongly convex; discrimen moderately to very long; transverse (katepisternal) suture almost complete to short or absent, but usually located close to posterior edge of ventrite; postcoxal lines sometimes forming distinct axillary spaces; exposed portion of metanepisternum moderately elongate, three to four times as long as wide, broader anteriorly. Metacoxae contiguous, extending laterally to meet elytra, plates more or less complete but often weakly developed. Metendosternite with short lateral arms, no laminae, short to very long anterior process and approximate to well separated anterior tendons. Hind wing usually 2 . 2 - 2 . 5 times as long as wide, with apical field 0 . 1 8 - 0 . 2 4

Artematopodidae Lacordaire, 1857

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times total wing length (0.37 times as long in Electribius)·, apical folds meeting apical wing margin separately; radial cell slightly to moderately elongate, with inner posterobasal angle right to acute; cross-vein r3 usually long, oblique, somewhat sinuate and meeting RP (although often incomplete in middle), sometimes shorter and more or less horizontal; basal portion of RP long to very long; apical portion absent; medial field with four or five free veins and no medial fleck; wedge cell usually well developed and apically truncate; anal lobe well developed, with AP undivided; anal embayment absent. Legs moderately long and slender, without tibial modifications; tibial spurs small, simple, equal; tarsi 5-5-5, tarsomeres 2 - 4 or 3 - 4 each with ventral, deeply bifid or occasionally emarginate lamella; basal pro- and mesotarsomeres with ctenidium in males of most Macropogon; pretarsal claws usually slightly expanded at base (with distinct basal tooth in Carcinognathus); empodium not prominent, usually concealed. Abdomen with five connate ventrites, the sutures between them mesally obscured in Electribius. Ventrite 1 distinctly shorter than 2, without postcoxal lines; intercoxal process acute to broadly rounded or angulate. Patches of glandular hairs on ventrites 2 - 4 in male Allopogonia. Ventrite 5 longer than preceding ones, often bearing glandular hairs near the apex, much longer in males of Brevipogon and Ctesibius, where it is completely, densely clothed with glandular hairs. Abdominal tergites moderately to heavily sclerotized. Spiracles located in pleural membrane, those on segment VIII functional. Anterior edge of sternite VIII in male without median strut (except in Electribius). Tergite IX in male usually slightly to moderately emarginate and X well developed and free or partly fused to IX (IX and X completely fused in Electribius); sternite IX usually broadly rounded at base, without spiculum gastrale. Aedeagus of trilobate type, symmetrical; parameres usually individually articulated, basally fused in Allopogonia, fused to form complete tube in Electribius. Penis undivided with short anterior struts. Sternite VIII in female with long spiculum ventrale. Ovipositor long and moderately slender, lightly sclerotized; paraprocts much longer than gonocoxites, which are subdivided, with apical styli. Internal female tract usually greatly enlarged in vicinity of gonopore to form uterine bursa bearing patches of asperities or window-like sclerites; anteriorlyattached spermatheca usually complex and multichambered, with gland attached near its apex; Electribius with large anterior bursa, with spermatheca and spermathecal gland attached by a long duct near its base. [Forbes 1926; Crowson 1955, 1973, 1975; Lawrence 1982, 1988, 1995, 2005; Caveney 1986; Kukalovä-Peck & Lawrence 1993, 2004; Lawrence etal. 1 9 9 5 , 1 9 9 9 b; Young 2002.] Morphology, Larvae (Figs. 4.2.2 A-C, E, F, 3 A, B). Body relatively straight, elongate, more or less

Fig. 4.2.3. Artematopus discoidalis Pic, larva: A, dorsal; B, ventral (from Costa etal. 1985; © Museu de Zoologia, Universidade de Säo Paulo); length = 1 5 mm. parallel-sided, subcylindrical to somewhat flattened. Dorsal surfaces usually moderately heavily pigmented, ventral surfaces lightly so. Vestiture consisting of fine hairs or setae only. Head protracted and prognathous, somewhat flattened, with rounded sides. Posterior edge distinctly emarginate. Epicranial stem absent or very short; frontal arms V-shaped or U-shaped, with bases usually well separated (apparently contiguous in Artematopus); endocarina absent. One stemma present on each side, with well developed lens. Frontoclypeal suture absent or only vaguely indicated; clypeus broadly emarginate anteriorly. Labrum separated from clypeus by suture but more or less immobile, strongly transverse, broadly, deeply emarginate anteriorly (in Artematopus slightly concave with median tooth and triangular epipharyngeal sclerome). Antennae short, 3-segmented, with sensorium on antennomere 2 usually half as long as 3 (as long as 3 in Artematopus), conical or palpiform. Mandibles symmetrical, moderately stout; apex obliquely bidentate with two dorsal retinacula, or unidentate with a single large retinaculum (Artematopus); accessory ventral process absent; mola and prostheca absent; mandibular base with brush of hairs in Artematopus. Ventral mouthparts

46 strongly retracted, forming maxillolabial complex; maxillary articulating area highly reduced or absent. Cardines undivided, transverse, well separated; stipes elongate; galea and lacinia short and subequal, the former 2-segmented and palpiform, the latter densely setose; palp 4-segmented. Labium consisting of prementum, mentum and submentum, the last sometimes unpigmented and not clearly separated from mentum, but distinctly separated from gula; ligula absent or shorter than labial palps, which are 2-segmented and separated by more than width of first palpomere. Hypopharyngeal sclerome present. Hypostomal rods absent; ventral epicranial ridges weakly developed. Gular sutures separate; gula longer than wide. Short and stout posterior tentorial arms arise immediately posterad of maxillolabial complex; other parts of tentorium reduced (Macropogon). Extrinsic and intrinsic labral muscles absent. Tentoriocardinal and tentoriostipital muscles strongly developed, almost vertical. Median premental retractor absent; M. tentoriopraementalis inferior bipartite, longer part originating from hind margin of head capsule likeM. tentoriopraementalis superior. Closed prepharyngeal tube present, strongly flattened; posterior precerebral pharyngeal dilator strongly developed; ventral dilators and dorsal postcerebral dilators absent. Brain and suboesophageal ganglion shifted to prothorax (Beutel 1995). Prothorax longer than meso- or metathorax but not as long as the two combined. Thoracic terga without special armature. Prothoracic presternum divided into anterior and posterior sclerites, the latter diamond-shaped with a longitudinal endocarina. Biforous mesothoracic spiracle located on spiracular sclerite (anterior laterotergite); atrophied metathoracic spiracle present;. Legs short, stout, 5-segmented, armed with short spines and setae; pretarsus claw-like, with two setae lying side by side; coxae large and narrowly separated. Abdominal terga I-VIII simple and subequal. Tergum IX distinctly longer than VIII, extending onto ventral surface; posterior three-fourths of dorsal surface slightly concave and lined laterally and posteriorly by a continuous ridge, which is often crenate or lined with slightly upturned teeth; sternum IX strongly transverse and apically emarginate. Segment X forming a short, cylindrical, ventrally projecting pygopod. Spiracles biforous, located on tergal plates above laterotergites. [Boving & Craighead 1931; Costa etal. 1985,1988; Cooper 1991; Lawrence etal. 1999 a.] Phylogeny and Taxonomy. Artematopodidae were included in Dascillidae in older works (Champion 1897; Pic 1914; Arnett 1963), moved (as Eurypogonidae) to the superfamily Dryopoidea by Crowson (1955), and later combined with Callirhipidae and Brachypsectridae in a separate elateriform superfamily Artematopoidea (Crowson 1973). Lawrence & Newton (1982) suggested that both Artematopodidae and Brachypsectridae (but

John F.Lawrence

not Callirhipidae) were part of a monophyletic group containing those families usually placed in Elateroidea and Cantharoidea. In cladograms produced by Lawrence (1988) and Lawrence et al. (1995), based on adult and larval characters, and Beutel (1995), based on larval features only, artematopodids were usually placed within Elateroidea in this broad sense. There are currently three subfamilies recognized: Electribiinae, Allopogoninae and Artematopodinae. Lawrence (1995) redescribed the fossil subfamily Electribiinae Crowson based on recentspecies,andHörnschemeyer(1998)described additional fossil forms. Ctesibiinae, proposed by Crowson (1973), was combined with Macropogoninae and Artematopodinae into a single subfamily by Lawrence (1995) and in a later study (Lawrence 2005) he elevated the tribe Allopogoniini Crowson (1973) to subfamily rank.

Acknowledgements CSIRO Australia is acknowledged for support of research and CSIRO Australia, Muzeum I Instytut Zoologici, PAN, Warsaw, Museu de Zoologia, Universidade de Säo Paulo and Prof. K. W. Cooper are gratefully acknowledged for allowing me to use copyrighted figures. The following artists are acknowledged: S. P. Kim (Figs. 4.2.1,2G) and S. Ide (Figs. 4.2.3 Α, Β). I am grateful to S. A. Slipiriski and A. Seago for assistance in obtaining and scanning published illustrations, Rolf Beutel for calling my attention to internal structures of the larval head, F. Dolambi for assistance with digital photography, and Anne Hastings for preparation of the plates. Portions of this Chapter benefited from research carried out with the support of a National Science Foundation grant "Assembling the Beetle Tree of Life" (DEB-0531768).

Literature Arnett, R. H. (1963): The Beetles of the United States (a Manual for Identification), xi + 1112 pp. Catholic University of America Press, Washington, D. C. Beutel, R. G. (1995): Phylogenetic analysis of Elateriformia (Coleoptera: Polyphaga) based on larval characters. - Journal ofZoological Systematics and Evolutionary Research 33:145-171. Boving, A. G. & Craighead, F. C. (1931): An illustrated synopsis of the principal larval forms of the order Coleoptera.-EntomologicaAmericanaiN.S.) 11 (1930): 1-351. Caveney, S. (1986): The phylogenetic significance of ommatidium structure in the compound eyes of polyphagan beetles. - Canadian Journal of Zoology 64:1787-1891. Champion, G. C. (1897): Fam. Dascillidae. Pp. 586-662, pis. 26-27 in F. D. Godman and O. Salvin (eds.) Biologia Centrali-Americana. Insecta, Coleoptera. Vol. III. Part 1. Porter, London. Cooper, K. W. (1991): Artematopidae (Elateroidea) (= Eurypogonidae). Pp. 407-409 in F. W. Stehr

47

Brachypsectridae Horn, 1881 (ed.) Immature Insects. Volume 2. Kendall/Hunt Publishing Co., Dubuque, Iowa. Costa, C., Casari-Chen, S. A. & Vanin, S. A. (1985): Larvae of Neotropical Coleoptera. XII. Artematopoidea, Artematopidae. - Revista Brasileira de Entomologia 29:309-314. Costa, C., Vanin, S. A. & Casari-Chen, S. A. (1988): Larvas de Coleoptera do Brasil. 282 pp. + 266 pis. Museu de Zoologia da Universidade de Säo Paulo, Säo Paulo. Crowson, R. A. (1955): The Natural Classification of the Families of Coleoptera. 187 pp. Nathaniel Lloyd, London. - (1973): On a new superfamily Artematopoidea of polyphagan beetles, with the definition of two new fossil genera from the Baltic Amber. - Journal of Natural History 7:225-238. - (1975): The evolutionary history of Coleoptera, as documented by fossil and comparative evidence. -Atti delXCongresso Nazionaleltaliano di Entomologia. Sassari, 20-25 Maggio 1974:47-90. Forbes, W. Τ. M. (1926): The wing folding patterns of the Coleoptera. -JournaloftheNew YorkEntomological Society 34:42-115, pis. 7-18. Hörnschemeyer, Τ. (1998): New species of Electribius Crowson 1973 (Coleoptera: Artematopodidae) from Baltic amber. - Paläontologische Zeitschrift 72 (3/4): 299-306. Iablokoff-Khnzorian, S. M. (1962): Representatives of Sternoxia (Coleoptera) in Baltic amber. - PaleontologicheskiiZhurnal 1962 (3): 81-89 (in Russian). Kukalovä-Peck, J. & Lawrence, J. F. (1993): Evolution of the hind wing in Coleoptera. - Canadian Entomologist 125:181-258. - (2004): Use of hind wing characters in assessing relationships among coleopteran suborders and endoneopteran lineages. - European Journal of Entomology 101: 95-144. Lacordaire, J. T. (1857): Histoire Naturelle des Insectes. Genera des Coleopteres... Vol. 4.5 79 pp. Libraire Encyclopedique de Roret, Paris. Lawrence, J. F. (1982): Coleoptera. Pp. 482-553 in Parker, S. P. (ed.) Synopsis and Classification of Living Organisms. Vol. 2. McGraw-Hill, New York. - (1995): Electribius Crowson: alive and well in Mesoamerica, with notes on Ctesibius Champion and the classification of Artematopodidae. Pp. 411-431 in: Pakaluk, J. & Slipiriski, S. A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warsaw. - (2005): Brevipogon, a new genus of North American Artematopodidae (Coleoptera). - The Coleopterists Bulletin 59 (2): 223-236. Lawrence, J. F. & Newton, A. F., Jr. (1982): Evolution and classification of beetles. - Annual Review of Ecology and Systematics 13:261-290. - (1995): Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names). Pp. 779-1006 in Pakaluk, J. & Slipiriski, S. A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warsaw. Lawrence, J. F., Nikitsky, N. B.&Kirejtshuk, A. G. (1995): Phylogenetic position of Decliniidae (Coleoptera:

Scirtoidea) and comments on the classification of Elateriformia (sensu lato). Pp. 375-410 in Pakaluk, J. & Slipiriski, S. A. (eds.) Biology, Phylogeny, and Classification ofColeoptera: Papers Celebrating the 80th Birthday ofRoy A. Crowson. Muzeum i Instytut Zoologii PAN, Warsaw. Lawrence, J. F., Hastings, A. M., Dallwitz, Μ. J., Paine, T. A. & Zürcher, Ε. J. (1999 a): Beetle Larvae of the World: Descriptions, Illustrations, Identification, and Information Retrieval for Families and Subfamilies. CD-ROM, Version 1.1 for MS-Windows. CSIRO Publishing, Melbourne. - (1999 b): Beetles of the World: A Key and Information SystemforFamiliesandSubfamilies. CD-ROM, Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne. Pic, M. (1914): Coleopterorum Catalogus. Pars 58. Dascillidae, Helodidae, Eucinetidae. 65 pp. W. Junk, Berlin, Young, D. K. (2002): 53. Artematopodidae Lacordaire, 1857. Pp. 146-147 in Arnett, R. H., Jr., Thomas, M. C., Skelley, P. E. & Frank, J. H. (eds.) American Beetles. Volume 2. polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Boca Raton, Florida.

4.3. Brachypsectridae Horn, 1881 Cleide Costa, Sergio A. Vanin, John F. Lawrence, Sergio Ide and Marc A. Branham Distribution. The family contains the single genus Brachypsectra LeConte with four described extant species: B. fulva LeConte occurring in the more arid parts of the southwestern United States and northern Mexico, B. vivafosile Woodruff from the Dominican Republic, B. lampyroides Blair from southern India and B.fuscula Blair from Singapore. In addition, an undescribed species from northern Australia is known from larvae only, and an adult and larvae of B. moronei Branham were described from Miocene amber in the Dominican Republic. [Lawrence & Newton 1995; Wu 1996; Woodruff 2004; Costa etal. 2006.] Biology and Ecology. Adult males of Brachypsectra are usually attracted to lights at night, but are probably short-lived. Most known females have been reared from larvae in the laboratory. Brachypsectra larvae are usually collected beneath loose bark, in cracks in rock, in leaf bases of monocolyledonous plants or under leaves or other debris on the ground. The larvae appear to be non-specific ambush predators, remaining inactive until approached by prey. Some of the prey items recorded for B. fulva are small spiders, an ant, an immature solpugid (Solifugae) and a larva of a tenebrionid beetle. When approached by a spider or other invertebrate of appropriate size, the larva was observed to arch its back, so that the prey is wedged between the articulated, spine-like tergum IX and the perforated sucking mandibles. It has been

Fig. 4.3.1. A-F, Adult habitus. A, Brachypsectrafulva (male); B, B.fulva (female) C, B. vivafosik (male); D, B. lampyroides (male); Ε, B. lampyroides (female); F, B.fuscula (male) (after Costa etal. 2006).

Fig. 4.3.2. A-H, Adult structures. Α-D, antennae: A, Brachypsectrafulva (male); B, B.fulva (female); C, B. lampyroides (male); D, B. vivafosik (male). E-H, mouthparts of B.fulva (male): E, mandible, dorsal; F, right maxilla (ventral); G, labium; H, labrum (after Costa etal. 2006).

Brachypsectridae Horn, 1881

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Fig. 4.3.3. A-C, Adult structures, Brachypsectrafulva (male). A, prothorax (ventral); B, hind wing; C, metendosternite (dorsal) (after Costa etal. 2006).

hypothesized that the small papillae on the dorsum of the larva may produce an allomone for attracting prey. Pupation occurs in a very wide-laced silken cocoon. [Ferris 1927; Neck 1993; Fleenor & Taber 1999; Costa etal. 2006.] Morphology, Adults (Fig. 4.3.1 A-F). Length about 4 to 8 mm. Body oblong to somewhat elongate, about 1.9-2.7 times as long as wide, more or less flattened and finely pubescent. Head slightly shorter than wide, slightly declined, deeply inserted into prothorax. Strongly constricted behind eyes forming very short neck consisting of little more than an expanded rim on either side of occipital foramen; dorsal rim of foramen slightly biemarginate. Occipital region without carina. Eyes large, more or less globular, strongly protruding and finely facetted, with exocone ommatidia (Caveney 1986) and no interfacetal hairs. Frontoclypeal region strongly declined anteriorly, so that mouthparts directed ventrally; transverse or longitudinal carinae absent. Antennal insertions slightly visible from above, placed within saucer-like fossae which are separated by flat area about two-thirds the width of one fossa; each continuing ventrally and laterally to form a short, oblique subantennal groove. Frontoclypeal suture absent; anterior edge of frontoclypeus broadly emarginate. Labrum (Fig. 4.3.2 H) relatively small; free, well sclerotized, slightly transverse and strongly rounded apically, with tormae straight, simple, and apically acute. Antennae (Figs. 4.3.2 A-D) 11-segmented, with antennomeres 4, 5 or 6 to 10 expanded apically on one side to form a pectinate

club with the terminal antennomere; scape slightly longer than wide and slightly longer than pedicel, which is subquadrate; antennomere 3 distinctly elongate, the following one or two slightly so. Mandible (Fig. 4.3.2. E) small, subtriangular, unidentate, without mola or prostheca. Maxilla (Fig. 4.3.2 F) with galea and lacinia subequal in length; galea more or less hyaline and lined with setae; lacinia lightly pigmented and densely setose, without hook or spine; apical palpomere fusiform. Labium (Fig. 4.3.2. G) with mentum transverse and trapezoidal; ligula short, slightly emarginate apically and setose; apical palpomere fusiform. Subgenal ridges absent. Gular sutures well separated posteriorly and strongly converging anteriorly. Corpotentorium absent. Cervical sclerites moderately well developed; anterior ones straight, slender and lightly sclerotized; posterior ones slightly longer, thicker and slightly curved. Pronotum about 0.5-0.6 times as long as wide; widest posteriorly, with distinct lateral carinae the anterior third of which are concealed from above; anterior angles more or less oblique, not produced forward; posterior angles acute, produced laterally and posteriorly, embracing elytral bases, bearing carinae which extend anteriorly and end at posterior third of disc, which is slightly convex in male but somewhat inflated anteriorly in female; posterior edge trisinuate with moderately developed interlocking device. Prosternum in front of coxae (Fig. 4.3.3 A) at least twice as long as shortest diameter of procoxal cavity, slightly convex, with short chin piece, slightly curved ventrally and anteriorly truncate; head rest strongly transverse and

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Cleide Costa, Sergio A.Vanin,John F. Lawrence, Sergio Ide and Marc A.Branham

slightly declined. Prosternal process moderately narrow, parallel-sided, slightly curved dorsally; apex broadly rounded, extending posteriorly to fit into mesoventral cavity. Notosternal sutures complete and more or less open (with exposed membrane). Procoxae not projecting below prosternum, with well-developed articulating area and slender trochantin more or less concealed by expansion of prosternal cowling; endopleuron free, not fused to notum. Procoxal cavities strongly transverse, externally broadly open, with narrow lateral extensions; internally open. Scutellar shield abruptly elevated, with straight, carinate basal edge, slightly rounded lateral edges and rounded to subtruncate or slightly emarginate apex. Elytra about 1.5-2 times as long as greatest combined width and 3.75-5 times as long as pronotum; sides subparallel or slightly wider at posterior third; apices conjointly, broadly rounded; disc flattened, with nine very weakly impressed punctate striae; sutural stria slightly more deeply impressed than others and scutellary striole absent; humeri moderately well developed; carinate anteriorly, the carinae continuing mesally and continuous with the scutellar carina; epipleura narrowed from base to about level of anterior third of metanepisternum, more or less uniform in width beyond that point and complete to apex. Mesoventrite separated by complete sutures from mesanepisterna, which are well separated from one another; anterior edge on same plane as metaventrite, with pair of large, shallow procoxal rests, which flank a raised median boss and are contiguous laterally with similar rests on mesanepisterna; posteriorly with a relatively large, deep mesoventral cavity, which extends well beyond anterior edges of coxal cavities; discrimen absent. Mesocoxae transverse, notprojecting, with exposed trochantins. Mesocoxal cavities separated by a little more than one-third shortest diameter of one of them; laterally open (partly closed by mesanepisternum and mesepimeron); without distinct joint between meso- and metathoracic portions of coxal cavities. Mesometaventral junction complex with metaventral knob fitting into notch on mesoventral process. Metaventrite relatively long and flat; discrimen well developed, vaguely indicated anteriorly but extending beyond base of metaventral process; transverse (katepisternal) suture absent; visible portion of metanepisternum moderately narrow, more or less parallel-sided and distant from mesocoxal cavity; metepimeron concealed beneath elytra. Metacoxae strongly transverse, extending laterally to meet epipleura; coxal plates narrow but more or less complete. Metendosternite (Fig. 4.3.3 C) with long stalk, short, broad arms, weakly developed ventrolateral processes and well developed, bilobed anterior process, with tendon at apex of each lobe and deep excavation between lobes. Hind wing (Fig. 4.3.3 B) about 1.85-1.9 times as long as wide, with very short apical field with a pair of strongly oblique, apically diverging sclerotizations; radial cell moderately large, elongate, its base

complete, forming right inner posterobasal angle; cross-vein r3 short and slightly oblique; RP extending to basal third of wing, without apical extension; medial field with five free veins; MP 3 + 4 with basal cross-vein and spur, apically forked; CuAj meeting MP 4 just beyond fork; wedge cell absent; anal lobe well developed, with undivided AP; anal embayment absent. Legs slender and simple; trochanters moderately elongate; trochanterofemoral joints on fore and mid legs slightly oblique, those on hind legs strongly so; tibial spurs absent; tarsi 5-5-5; tarsomeres 1 - 4 combined more than twice as long as 5; tarsomere 4 slightly reduced with weak ventral lobe; pretarsal claws simple; empodium visible, bisetose. Abdomen with five free ventrites. Ventrite 1 not much longer than 2, without postcoxal lines; intercoxal process acute. Ventrites 1 - 4 subequal in length, 5 broadly rounded at apex. Abdominal spiracles on segments I to VIII, located in pleural membrane. Sternite VIII in male anteriorly bisinuate, forming pair of short lateral struts and short, broad, rounded median plate. Sternite IX broadly rounded anteriorly, subtruncate posteriorly. Tergite IX in male distinctly emarginate and separated from tergite X, which is subtruncate. Sternite VIII in female trapezoidal, posterior edge sinuous and setose; spiculum ventrale well developed. Aedeagus (Figs. 4.3.4 Α-B) of trilobate type, symmetrical; phallobase about as long as wide, slightly emarginate at base; parameres with apical, laterally curved hooks; penis undivided, with short basal struts. Ovipositor moderately elongate, lightly sclerotized; paraprocts slightly longer than coxites, which are transversely divided into two lobes; proximal lobe with obliquely transverse baculum; distal lobe narrowed and palp-like; stylus well developed, terminal. Internal tract with long anterior bursa and spermathecal duct entering between gonopore and base of bursa; spermatheca lightly pigmented. Malpighian tubules four, free. Midgut with regenerative crypts. [Hlavac 1975; Kasap & Crowson 1975; Lawrence & Britton 1991, 1994; Lawrence etal. 1999 b; Young 2002; Woodruff 2004; Costa etal. 2006.] Morphology, Larvae (Figs. 4.3.5 A-B; 5 A-B; 7 Α-B). Length up to 15 mm. Body broadly ovate, strongly flattened and disc-like, with moderately long, branched projections on all thoracic segments and abdominal segments I—Villi; head and tergite IX darkly pigmented, upper surfaces unevenly pigmented, varying from white to yellow or brown, covered with small setiferous tubercles, flattened pigmented areas of varying size and shape, and a median row of subconical projections lined with setiferous tubercles; lower surfaces very lightly pigmented, except for presternal area, thoracic spiracular sclerites, and apical portions of legs. Vestiture consisting mainly of expanded, modified setae, which on most of the dorsal surface form an irregularly multisided, flattened scales with a median stalk and spiculate upper surface.

Brachypsectridae Horn, 1881

51

Fig. 4.3.4. Α-B, Aedeagus, Brachypsectrafulva, A, ventral; B, dorsal (after Costa etal. 2006).

Fig. 4.3.5. Α-B, Larval habitus. A, Brachypsectra lampyroides; B, Brachypsectra sp. (after Costa etal. 2006).

Head (Figs. 4.3.6 A-B, 7 Α-B) p r o g n a t h o u s a n d protracted, elongate a n d more or less parallel-sided, m u c h narrower t h a n prothorax; usually slightly elevated a n d capable of being strongly elevated. Posterior edge as seen f r o m above broadly emarginate. Epicranial stem very short; frontal arms slightly lyriform, sometimes vaguely indicated. Endocarinae absent. One large s t e m m a on each side, w i t h welldeveloped lens. Frontoclypeal s u t u r e present or absent. Labrum free, slightly transverse, e x p a n d e d apically w i t h biemarginate apex bearing f o u r s t o u t

setae. Antennae well developed, a b o u t two-thirds as l o n g as greatest head w i d t h , 3-segmented; antenn o m e r e 1 a b o u t as long as wide, and 2 a b o u t 8 times as l o n g and irregularly club-shaped, heavily pigm e n t e d and clothed w i t h modified setae; antennomere 3 a n d palpiform sensorium highly reduced a n d lying side by side at outer edge of oblique apex of a n t e n n o m e r e 2. Mandibles symmetrical, narrow a n d falcate, strongly curved mesally, perforate w i t h completely enclosed internal channel; widely separated at base, w i t h o u t mola or prostheca. Ventral

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CleideCosta,Sergio A.Vanin,John F.Lawrence,Sergio Ide and Marc A.Branham

Β Fig. 4.3.6. Α-B, Larval head, Brachypsectrafulva. A, dorsal; ventral (after Costa etal. 2006).

mouthparts moderately retracted, forming maxillolabial complex without articulatory area; maxillae with cardines fused together forming single median, trapezoidal plate; stipes elongate; galea small, palpiform and articulated, with single seta at apex; lacinia fixed, slender, curved and falcate; palp 4-segmented, with palpomere 2 as long as 1, 3 and 4 combined. Labium with m e n t u m and subment u m fused; postmentum completely fused to stipes on each side, separated by slight impression only; palps 2-segmented, approximate, with palpomere 1 much longer than 2; ligula absent. Hypopharynx without sclerome. Hypostomal region (ventral head closure) longer than maxillolabial complex; gular sutures fused. Hypostomal rods and ventral epicranial ridges absent. Thoracic terga and abdominal terga I-VIII moderately to strongly transverse; the former each with a pair of narrow subacute, slightly posteriorly curved, branched processes on each side; the latter each with a single similar process in f r o n t of which is a reduced process without branches. Protergum slightly longer t h a n either meso- or metathoracic terga; preceded by membranous cervical region bearing transverse row of setiferous tubercles; body of protergum with single median oval projection and pair of moderately large, irregular p i g m e n t patches, which may be broken u p or accompanied by smaller patches laterally; centers of patches or entire small patches with smooth surfaces, in contrast to other areas, which are more or less uniformly covered with setiferous tubercles. Meso- and metaterga similar, b u t with main p i g m e n t patches smaller and more broken up, and with median projections larger t h a n that on protergum. Prothoracic s t e r n u m unpigmented, except for pair of patches on cervicosternum; clothed mainly with short,

simple stout setae, except in f r o n t of procoxae and on cervicosternum, where most setae arise f r o m tubercles; procoxae separated by slightly more than greatest coxal diameter; meso- and metacoxae separated by about two coxal diameters. Legs well developed, with tibiotarsus slightly longer than f e m u r and pretarsus forming distinct claw bearing a pair of setae. Abdominal segments I-VIII about 1.5 times as long as thorax; terga I-VII all with median projection, smaller paramedian patches, and one or two small, rounded p i g m e n t patches just behind spiracular opening; paramedian patches often small and transverse on anterior segments b u t larger on posterior ones. Tergum VIII w i t h o u t median projection and with paramedian patches occupying most of dorsum. Tergum IX f o r m i n g long, narrow, heavily sclerotized, articulated plate, which has several lateral branches at base, narrows towards subacute apex, and is covered with setiferous tubercles. Anterior abdominal sterna without or with one pair of small lateral pigment spots, these increasing in size and n u m b e r on sterna V to VII. Sternum VIII with single pair of larger pigment patches. Sternum IX m e m b r a n o u s and segment X not distinguishable. Spiracles biforous b u t reduced in size, borne on short spiracular tubes; mesothoracic spiracles located ventrally on anterolateral transverse spiracular sclerites; abdominal spiracles located dorsally on lateral portion of tergite; all spiracles with closing apparatus. [Barber 1905; Blair 1930; Boving & Craighead 1931; Lawrence 1991; Lawrence & Britton 1991, 1994; Lawrence etal. 1999 a; Young 2002; Costa etal. 2006.] Phylogeny a n d T a x o n o m y . LeConte (1874) placed his genus Brachypsectra in Rhipiceridae, based on its

Brachypsectridae Horn, 1881

53

Fig. 4.3.7. Α-B, Larval head, Brachypsectra sp. (Australia). A, dorsal; B, ventral (after Costa etal. 2006).

resemblance to Zenoa Say (now placed in Callirhipidae), and H o r n (1881) proposed the tribe Brachypsectrini w i t h i n Dascillidae. Forbes (1926) f o u n d t h a t t h e type of w i n g folding in Brachypsectra was similar to t h a t f o u n d in Lampyridae a n d Elateridae, b u t n o t in Dascillidae, a n d Blair (1930) concluded t h a t the tribe s h o u l d be raised to family rank, allied m o r e closely to Elateridae t h a n Dascillidae. Crowson vacillated between placing the family in Cantharoidea (1955) or in a g r o u p also c o n t a i n i n g the families Artematopodidae a n d Callirhipidae (1973). Kasap & Crowson (1975) r e t u r n e d the family to Cantharoidea based mainly on the free a b d o m i n a l ventrites (with m o r e or less complete longitudinal musculature), a condition considered to be derived t h r o u g h neoteny in Elateriformia and unlikely to have evolved i n d e p e n d e n t l y in the two groups. In cladograms produced by Lawrence (1988) a n d Lawrence et al. (1995) based on b o t h a d u l t a n d larval characters, Brachypsectridae was placed at or near the base of a clade containing Cerophytidae, Eucnemidae, Throscidae a n d Elateridae (Elateroidea sensu stricto) or a clade including these families plus the Cantharoidea (Elateroidea sensu lato). In Beutel's (1995) analysis of elateriform larvae, the family always clustered w i t h Cantharoidea based mainly on the channeled mandibles. [Lawrence & N e w t o n 1982,1995.]

Acknowledgements T h a n k s are d u e to Simone P. Rosa for h e l p i n g in t h e electronic t r e a t m e n t of t h e figures. T h e E n t o m o l o g ical Society of America is gratefully acknowledged for allowing us to use copyrighted illustrations. T h a n k s are also d u e to CNPq for the Research G r a n t 302721/2007-0 to C. Costsa.

References Barber, H. S. (1905): Illustrations of an undetermined coleopterous larva. - Proceedings of the Entomological Society of Washington 7:117-121. Beutel, R. G. (1995): Phylogenetic analysis of Elateriformia (Coleoptera: Polyphaga) based on larval characters. -Journal of Zoological Systematics and Evolutionary Research 33:145-171. Blair, K. G. (1930): Brachypsectra, Lec. - the solution of an entomological enigma. - Transactions of the Royal Entomological Society ofLondon 7 8 : 4 5 - 5 0 , 1 pi. Boving, A. G. & Craighead, F. C. (1931): An illustrated synopsis of the principal larval forms of the Coleoptera. -Entomologica Americana (N.S.) 11:1-351. Caveney, S. (1986): The phylogenetic significance of ommatidium structure in the compound eyes of polyphagan beetles. - Canadian Journal of Zoology 64:1787-1891. Costa, C., Vanin, S.A., Lawrence, J. F., Ide, S. & Branham, M. A. (2006): Review of the family Brachypsectridae (Coleoptera: Elateroidea).-Annals ofthe Entomological Society of America 99 (3): 409-432. Crowson, R. A. (1955): The Natural Classification of the Families of Coleoptera. 187 pp. Nathaniel Lloyd, London. - (1973): On a new superfamily Artematopoidea of polyphagan beetles, with the definition of two new fossil genera from the Baltic Amber. - Journal of Natural History 7:225-238. Ferris, G. F. (1927): Notes on an entomological enigma. - The Canadian Entomologist 59:279-281. Fleenor, S. B. & Taber, S. W. (1999): Review of Brachypsectra with a new record of the Texas Beetle (B. fulva LeConte; Coleoptera: Brachypsectridae). - The ColeopteristsBulletin 53:359-364. Forbes, W. Τ. M. (1926): The wing folding patterns of Coleoptera. - Journal of the New York Entomological Society 34 (2): 4 2 - 6 8 , 9 1 - 1 3 6 .

54 Hlavac, T. F. (1975): The p r o t h o r a x of Coleoptera: (except Bostrichiformia - Cucujiformia). - Bulletin of the Museum of Comparative Zoology 147: 137-183. Horn, G. W. (1881): Notes on Elateridae, Cebrionidae, Rhipiceridae, and Dascyllidae. - Transactions of the American Entomological Society 9 : 7 6 - 9 0 , 2 pis. Kasap, H. & Crowson, R. A. (1975): A comparative anatomical study of Elateriformia and Dascilloidea (Coleoptera). - Transactions of the Royal Entomological Society ofLondon 126:441-495. Lawrence, J. F. (1988): Rhinorhipidae, a new beetle family from Australia, with comments on the phylogeny of the Elateriformia. - Invertebrate Taxonomy 2:1-53. - (1991): Brachypsectridae(Cantharoidea).Pp.421-422 in Stehr, F. W. (ed.) Immature Insects, Vol. 2, Kendall/ Hunt Publishing Co., Dubuque, Iowa. Lawrence, J. F. & Britton, Ε. B. (1991): Coleoptera (Beetles). Pp. 543-683 in CSIRO Division of Entomology (ed.) Insects of Australia: a Textbookfor Students and Research Workers, Second Edition. Vol. 2. Melbourne University Press, Carlton, Victoria. - (1994): Australian Beetles. Melbourne University Press, Carlton, Victoria, χ + 192 pp. Lawrence, J. F. & Newton, A. F., Jr. (1982): Evolution and classification of beetles. - Annual Review ofEcology andSystematics 13:261-290. - (1995): Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names). Pp. 779-1092 in Pakaluk, J. & Slipinski, S. A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warszawa. Lawrence, J. F., Hastings, A. M., Dallwitz, Μ. J., Paine, T. A. & Zürcher, Ε. J. (1999 a): Beetle Larvae of the World: Descriptions, Illustrations, Identification, and Information Retrieval for Families and Subfamilies. CD-ROM, Version 1.1 for MS-Windows. CSIRO Publishing, Melbourne. - (1999 b): Beetles of the World: A Key and Information System for Families and Subfamilies. CD-ROM, Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne. Lawrence, J. F., Nikitsky, Ν. B. & Kirejtshuk, A. G. (1995): Phylogenetic position of Decliniidae (Coleoptera: Scirtoidea) and comments on the classification of Elateriformia (sensu lato). Pp. 3 7 5 - 4 1 0 in Pakaluk, J. & Slipinski, S. A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. M u z e u m i Instytut Zoologii PAN, Warszawa. LeConte, J. L. (1874): Description of new Coleoptera chiefly from Pacific slope. - Transactions ofthe American Entomological Society 5:43-72. Neck, R. W. (1993): Notes on habitats and rearing of Brachypsectra fulva LeConte (Coleoptera: Brachypsectridae). - The Coleopterists Bulletin 47: 291-292. Woodruff, R. E. (2004): A new species of the beetle genus Brachypsectra from the Dominican Republic, with fossil connections (Coleoptera: Brachypsectridae). -InsectaMundi 16:161-170.

Cleide Costa,Sergio A.Vanin,John F.Lawrence and Sergio Ide Wu, R. J. C. (1996): Secrets of a Lost World: Dominican Amber and its Inclusion. Ill pp. R. J. C. Wu, Santo Domingo, Dominican Republic (Amber West, Colorado Springs, Colorado). Young, D. K. (2002): 54. Brachypsectridae Boving and Craighead 1931. Pp. 148-149 in Arnett, R. H., Jr., Thomas, M. C., Skelley, P. E. & Frank, J. H. (eds.) American Beetles. Volume 2. polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Boca Raton, Florida.

4.4. Cerophytidae Latreille, 1834 Cleide Costa, Sergio A.Vanin, John F. Lawrence and Sergio Ide

Distribution. T h e family includes three genera: the Holarctic Cerophytum Latreille and the Neo-tropical Brachycerophytum Costa et al. a n d Phytocerum Costa et al. T h e distributions of the 21 k n o w n species are as follows: Nearctic Region: C. convexicolle LeConte (western U. S. A.) a n d C. pulsator( Haldeman) (eastern U. S. A.); Palaearctic Region: C. elateroides (Latreille) (southern and central Europe) a n d C. japonicum Sasaji (Japan); Neotropical Region: B. fuscicorne (de Bonvouloir) (Mexico to Colombia), B. sinchona Costa etal. (Peru a n d Bolivia), P. alleni Costa etal. (Costa Rica and Nicaragua), P. belloi Costa et al. (Brazil), P. birai Costa et al. (Brazil), P. boliviense (Golbach) (Bolivia), P. burakowskii Costa et al. (Trinidad), P. cayennense (de Bonvouloir) (Mexico, French Guiana, and Brazil),

Fig. 4.4.1. Phytocerum minutum, male (from Costa etal. 2003), Length = 5.1 m m .

Cerophytidae Latreille, 1834

P. distinguendum (Soares & Peracchi) (Brazil, Paraguay, and Argentina), P. golbachi Costa et al. (Argentina), P. ingens Costa et al. (Brazil), P. inpa Costa et al. (Brazil), P. minutum (Golbach) (Argentina), P. simonkai Costa et al. (Trinidad), P. serraticorne Costa et al. (Guatemala), P. trinidadense (Golbach) (Trinidad), and P. zikani (Soares & Peracchi) (Brazil). [Costa etal. 2003]. Biology and Ecology. Adult cerophytids have been collected at light traps or Malaise traps, by beating vegetation, or in association with rotten wood or bark. Cerophytum elateroides has been reported from a n u m b e r of deciduous tree genera, including Ulmus (Ulmaceae), Fagus and Quercus (Fagaceae), Salix and Populus (Salicaceae), Acer (Aceraceae), Tilia (Tiliaceae), Betula (Betulaceae), and Juglans (Juglandaceae). Adults are capable of "clicking" in the same manner as Elateridae (by the sudden release of the prothorax, which is held under tension by a small projection at the anterior edge of the mesoventral cavity), b u t it is also possible that the enlarged and modified profemora and the lack of metacoxal plates in Cerophytidae are both connected in some way with

55

escape behavior. The larva of C. elateroides was first described by Rey (1887) based on specimens collected with an adult in the rotten t r u n k of Sambucus (Caprifoliaceae). Mamaev (1978) redescribed and illustrated the Cerophytum larva based on a series of specimens found in the dark fungus-infested wood (brown rot) of a dead standing Ulmus in association with larvae of Oedemeridae. It is likely that these larvae feed in old b r o w n - r o t t e n wood belonging to a variety of hardwoods. [Buysson 1910; Horion 1953; Lawrence 1991; Steiner 2000; Johnson 2002; Costa etal 2003.] Morphology, Adults (Fig. 4.4.1). Length 4.3-9.2 m m . Body about 2.2-2.6 times as long as wide, slightly flattened above, strongly convex below, usually black or yellowish-brown in color and clothed with recumbent and/or suberect hairs. Head about as long as wide, deeply inserted into prothorax and only slightly declined, not abruptly constricted posteriorly. Occipital region with weak transverse carina continuing behind and below eyes to form short subgenal ridges. Median endocarina

Fig. 4.4.2. Cerophytum elateroides, A, fore leg; B, hind leg. Brachycerophytumfuscicorne; C, fore leg; D, hind leg; G, basolateral portion of left elytron. Phytocerum distinguendum; E, fore leg; F, hind leg. Phytocerum boliviense; H, basolateral portion of left elytron. Scale lines = 1.0 mm (A-F, G, H). (from Costa etal. 2003).

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Cleide Costa,Sergio A.VaninJohn F.Lawrence and Sergio Ide

Fig. 4.4.3. Aedeagus, ventral, dorsal, lateral, and lateral with parts separated. Α-D, Cerophytum pulsator, E-H, Brachycerophytum sinchona; I-L, Phytocerum cayennense; pa = parameres; pe = penis; ph = phallobase. Scale lines = 1 . 0 mm (A-C, E-G, I-K, D, H, L). (from Costa etal. 2003).

absent. Frons produced in front of eyes forming median prominence bearing antennal insertions, with or without median carina, strongly declined anteriorly, so that frontoclypeal region is more or less vertical and mouthparts are ventrally oriented; frontoclypeal suture absent. Eyes large, more or less globular, strongly protruding and finely facetted, with exocone ommatidia and no interfacetal setae. Antennal insertions moderately close together,

concealed by frontal ridges or partly exposed from above; subantennal grooves absent. Labrum free, well sclerotized, strongly transverse and truncate. Antennae 11-segmented, serrate to pectinate from antennomere 3 to 10, with rami arising from base of each; 11 simple and narrowly rounded or notched; scape more than 3 times as long as pedicel, which is attached subapically, so that antennae are more or less geniculate. Mandibles narrow, strongly

Cerophytidae Latreille, 1834

curved, unidentate and acute; without mola b u t with membranous prostheca. Maxillae with b o t h galea and lacinia densely setose at apex; apical palpomere more or less expanded and truncate, usually securiform or elongate oval. Ligula membranous, rounded apically; palps approximate; apical palpomere subtriangular. Gular sutures widely separated. Corpotentorium absent. Cervical sclerites well developed. Prothorax about 0.55-0.7 times as long as wide. P r o n o t u m usually widest anteriorly, without lateral carinae; anterior angles not produced; posterior angles acute and slightly produced laterally or obtuse; posterior edge more or less straight or slightly sinuate; disc moderately convex; interlocking device weakly developed. Notal hypomera extensive, obliquely vertical; notosternal sutures complete. Prosternum in f r o n t of coxae at least twice as long as shortest diameter of procoxal cavity, strongly convex; anterior edge truncate or with ventrally directed chin piece; head rest (upper surface of anterior edge of sternum) strongly oblique (from lateral perspective) and hemispherical to strongly transverse; prosternal process moderately broad and curved dorsally, laterally expanded at middle to form secondary condyle on each side, then narrowed to form acute apex which fits into mesoventral cavity. Procoxae not projecting ventrally, each with long, narrow internal extension, articulating with long, concealed trochantin and its attached endopleuron, the latter not fused to hypomeron. Procoxal cavities circular to slightly transverse, open internally and externally. Scutellar shield abruptly elevated, with straight basal edge; lateral edges slightly rounded and apex subacute to rounded. Elytra about 1.7-2 times as long as combined width and 3.4-4.4 times as long as p r o n o t u m ; with nine distinct rows of deep window punctures, sometimes with an additional subhumeral row (Fig. 4.4.2 G-H); scutellary striole absent; epipleuron wide anteriorly, abruptly narrowed just behind h u m e r i and extending almost to apex, narrow posterior portion weakly delimited and almost vertical (completely visible in lateral view). Mesoventrite on same plane as metaventrite, with large, deep mesal cavity, preceded by sclerotized lip, flanked anteriorly by a pair of concave areas and posteriorly by mesocoxal cavities, and extending posteriorly almost to edge of ventrite. Mesoventrite separated by complete sutures f r o m mesanepisterna, which are widely separated at midline, each posteriorly fused to mesepimera (a vague internal ridge probably indicating position of reduced pleural apophysis) and with a narrow, oblique housing into which the pronotal hypomeron fits; anterior edge of mesoventrite on same plane as metaventrite, with large, deep mesal cavity, preceded by a convex lip (sometimes with pair of small, slightly declined procoxal rests), flanked anteriorly by pair of concave areas and posteriorly by mesocoxal cavities, and extending posteriorly almost to edge of ventrite. Mesocoxal cavities oblique, separated by

57

more than shortest diameter of one of them; open laterally (partly closed by mesepimeron), with solid joint between meso- and metathoracic portions of cavity; meso-metaventral junction a straight line. Metaventrite strongly convex, w i t h o u t discrimen or transverse (katepisternal) suture; visible portion of metanepisternum very narrow and distant f r o m mesocoxal cavity; metacoxae large, somewhat oblique, strongly developed internally, and extending laterally to meet elytral epipleura; coxal plates absent. Metendosternite with long stalk, short, broad arms, and well developed anterior process bearing approximate tendons. H i n d wing about 2.25 times as long as wide, with very short apical field bearing two parallel, oblique, anterior sclerotizations and sometimes an additional oblique posterior one; radial cell elongate, its base complete, f o r m i n g a right or very slightly acute angle; cross-vein r3 moderately long and slightly oblique, extending almost to RP, which extends to basal third of wing; medial field with four free veins (MP 3 , MP 4 +CUAJ, CUA 2 a n d AA 3 ); w e d g e cell a n d

anal notch absent, AP 3+4 undivided. Legs (Fig. 4.4.2 A-F) moderately long and slender. Protrochanter less than one-fifth length of f e m u r with trochanterofemoral joint truncate or slightly oblique; prof e m u r subequal in length to protibia, with ventral longitudinal carina and slight concavity into which the protibia fits, sometimes with additional longitudinal carina on posterior surface. Meso- and metatrochanters more than half as long as corresponding femora with trochantofemoral joints strongly oblique; meso- and metafemora without longitudinal carina and only 0.75 to 0.85 times as long as corresponding tibiae. Tibial spurs paired and subequal on all tibiae; tarsi 5-5-5; tarsomeres 1 and 5 about equal in length, 2 shorter than 1 , 3 and 4 subequal, the latter with a ventral lobe; pretarsal claws pectinate; e m p o d i u m not visible. Abdomen strongly convex. Basal four ventrites connate, 1 slightly shorter than 2, with narrow, acute intercoxal process. Ventrites 2, 3 and 4 subequal; 5 broadly rounded at apex. Abdominal spiracles present on segments I to VIII, located in pleural membrane. Sternite VIII in male broadly rounded anteriorly, without median strut. Sternite IX similarly rounded at base, produced apically to form elongate, narrow, apically rounded, lightly sclerotized process. Tergite IX in male narrowly emarginate almost to base, distinctly separated f r o m tergite X, which is long and narrow with rounded apex. Aedeagus (Fig. 4.4.3 A-L) of trilobate type, symmetrical; phallobase, parameres and penis with extensive weakly sclerotized areas; phallobase ventrally less sclerotized in the center and more sclerotized at the proximal edge, with shallow to deep anterior emargination and long, narrow posterior process which is closely associated with ventral surface of the penis; parameres ventrally, loosely articulated to phallobase fused dorsally at base f o r m i n g a more or less developed convex anterior projection which is divided into two parts, the proximal one

58

Cleide Costa,Sergio A . V a n i n J o h n F.Lawrence and Sergio Ide

Fig. 4.4.4. Cerophytum elateroides, larva, dorsal. Scale line =1.0 m m . (from Costa etal. 2003).

Fig. 4.4.5. Cerophytum elateroides, larva, ventral. Scale line = 1.0 m m . (from Costa etal. 2003).

sclerotized a n d w i t h variable hook-like structures a n d t h e distal one m e m b r a n o u s , unilobate, bilobate or trilobate; penis m o r e or less flattened; base w i t h m e d i a n , dorsally curved s t r u t w h i c h joins parameres dorsally at their p o i n t of f u s i o n . Sternite VIII in female f o r m i n g l o n g s p i c u l u m ventrale. Ovipositor l o n g a n d narrow; coxites n o t divided, styli s h o r t a n d terminal, baculi elongate. Internal female tract consisting of elongate vagina, large t e r m i n a l b u r s a

w i t h one or two pairs of basal e m b e d d e d sclerites; spermatheca present or absent. [Caveney 1986; Costa etal. 2003.] M o r p h o l o g y , Larvae. L e n g t h 3 . 0 - 1 5 . 0 m m . Body (Figs. 4.4.4-5) elongate, r o b u s t b u t m o r e or less parallel-sided, slightly curved ventrally. W h i t e in color, except for legs, anterior p a r t of p r o t e r g u m a n d h e a d capsule, w h i c h are yellow, a n d pretarsus,

Fig. 4.4.6. Cerophytum elateroides, larval head, Α, dorsal; Β, ventral. Scale line = 1.0 mm. (from Costa et dl. 2003).

internal prothoracic skeleton and labial plate, which are dark brown; vestiture consisting of fine setae only. Head (Fig. 4.4.6 Α-B) prognathous and protracted or slightly retracted, m u c h narrower t h a n thorax, strongly transverse and flattened above. Posterior edge not emarginate. Frontal arms and dorsal endocarinae absent. One stemma with well developed lens present on each side of head. Frontoclypeal suture vaguely indicated. Labrum fused to head capsule, strongly transverse, deeply emarginate and lightly pigmented. Antennae short, 3-segmented; subconical sensorium on preapical antennomere longer t h a n apical antennomere, which bears a second conical sensorium. Mandibles flattened, symmetrical and non-opposable, more or less parallel to one another and fitting into lateral channels on the labial plate; each mandible with basolateral, rounded expansion, narrow, blade-like, sharply acute apex, and internal channel extending f r o m near base to apex; mola, prostheca and accessory ventral process absent. Ventral m o u t h p a r t s strongly retracted forming maxillolabial complex; maxillary articulating area absent. Maxillae well separated f r o m one another by labium; cardines not clearly delimited; stipes elongate; galea and lacinia f o r m i n g slender, blade-like, channelled mala, which is narrowly acute at apex; maxillary palpomeres three, b u t palpiger enlarged and partly articulated at base, so that palps appear 4-segmented. P r e m e n t u m and hypopharynx forming 5-dentate sclerome; p o s t m e n t u m subquadrate and strongly convex; labial palps 2-segmented with subacute apical palpomere, closely adpressed to side of sclerome and scarcely visible except in side view. Ventral longitudinal endocarinae present along lateral edges of maxillae and between maxillae and labium. Hypostomal region, hypostomal rods, ventral epicranial ridges and gula absent. Prothorax almost as long as meso- and metathorax combined. Thoracic terga simple, without sclerotization, asperities, rugosities or carinae. Venter with two pairs of anterior sclerotizations: lateral, subtrapezoidal and slightly curved pleurites, which

extend from the neck region to the anterior articulations of procoxae, and oblique, slender, sternal rods, which converge posteriorly and define a subtriangular area, which is lightly sclerotized and has a short endocarina near the posterior end. Legs relatively short. Procoxae large, oval, strongly oblique and moderately heavily pigmented, separated by about 0.25 times the shortest diameter of one of them; protrochanters short and lightly pigmented; femora stout, only slightly longer than wide, subcylindrical; tibiotarsus short and broad, about one-third as long as femur; pretarsus slightly shorter than tibiotarsus, heavily pigmented, deeply emarginate forming a pair of claw-like processes, one slightly shorter than the other; pretarsal setae two, lying side by side. Mesocoxae and metacoxae smaller than procoxae and separated by about 3 times the shortest diameter of one of them; trochanter similar to that on fore leg; femur more slender, about twice as long as wide; tibiotarsus about two-thirds as long as femur and slender; pretarsus slender, claw-like, with two setae. Abdomen about 3 times as long as thorax. Segments I-VIII each with three clearly defined lateral lobes on each side (laterotergites, pleurites and laterosternites). Terga densely clothed with long hairs b u t without sclerotizations, carinae or asperities. Paired gland openings present on all thoracic terga and abdominal terga I-VIII. Segment IX about half as long as VIII, tergum dorsal, simple, without urogomphi; X about one third as long as IX, cylindrical and terminal, w i t h o u t pygopods. Thoracic and abdominal spiracles biforous; closing apparatus present. [Mamaev 1978; Lawrence 1991; Lawrence etal. 1999 a; Costa etal. 2003.]

Phytogeny and Taxonomy. Latreille(1825) placed his genus Cerophytum in the group Sternoxi of the Familie Serricornia, along with Buprestidae and most elateroid genera k n o w n at the time. In 1834, h e proposed a family group based on Cerophytum, and this was recognized by Lacordaire (1857) who suggested a relationship with Eucnemidae. Crowson (1955) considered the family to be part of the

60 Elateroidea (sensu stricto), a n d this has been followed by m o s t workers. T h e u n u s u a l , apparently plesiom o r p h i c n a t u r e of t h e cerophytid propleurocoxal m e c h a n i s m p r o m p t e d Hlavac (1975) to remove t h e g r o u p f r o m Elateroidea a n d place it together w i t h two o t h e r elateriform families of d o u b t f u l affinities, A r t e m a t o p o d i d a e a n d Brachypsectridae. In cladog r a m s p r o d u c e d by Lawrence (1988) Cerophytidae f o r m e d a clade w i t h E u c n e m i d a e a n d Throscidae {sensu stricto) or was basal to a clade c o n t a i n i n g these two families plus Elateridae. A similar association was f o u n d in analyses conducted by Calder et al. (1993), Beutel (1995), a n d Lawrence etal. (1995). In M u o n a ' s (1993) revision of Eucnemidae, Cerophytidae were considered basal to t h e elateroid c o m p l e x (Elateroidea sensu Crowson 1955) a n d in a later cladistic analysis (Muona 1995) Cerophytidae a n d E u c n e m i d a e f o r m e d one clade whereas Throscidae (sensu Crowson 1955) represented a derived g r o u p w i t h i n Elateridae. T h e larva of Cerophytum is highly u n u s u a l a n d differs f r o m those of all other Elateroidea in h a v i n g u n i q u e m o u t h p a r t s involving paired s u c k i n g grooves, styliform m a n d i b l e s a n d m a x illae, paired thoracic a n d a b d o m i n a l glands a n d bifurcate pretarsi. The g e n u s Anischia F l e u t i a u x (1896) was considered by its a u t h o r to be related to Cerophytum, mainly because of t h e lack of metacoxal plates, a n d is o f t e n placed in Cerophytidae (Schenkling 1928; Crowson 1955). T h e g e n u s has also been placed in Elateridae (Fleutiaux 1936; Lawrence & N e w t o n 1995) or considered as a separate family (Lawrence etal. 1999 b), b u t a recent cladistic analysis of larval a n d a d u l t characters c o m b i n e d w i t h DNA sequence data strongly s u p p o r t s its inclusion in E u c n e m i d a e (Lawrence et al. 2007). T h e extinct g e n u s Aphytocerus Zherichin (1977) has also been included in this family based on two species in fossil resin (retinite) f r o m t h e U p p e r Cretaceous of Siberia. Its inclusion in the family m u s t r e m a i n tentative based on t h e limited i n f o r m a t i o n available on the s t r u c t u r e of t h e fossil specimens. Costa et al. (2003) carried o u t a phylogenetic analysis of t h e 21 k n o w n species in t h e family, w i t h o u t g r o u p s f r o m Elateridae, Throscidae a n d E u c n e m i d a e . Three m a i n clades were recognized: (Cerophytum (Brachycerophytum, Phytocerum)). Syna p o m o r p h i e s for the family include: 1) chin piece ventrally directed; 2) posterior p r o n o t a l angles p r o d u c e d laterally; 3) m e t a t r o c h a n t e r s m o r e t h a n half as l o n g as m e t a f e m o r a ; 4) base of tergite IX a n d sternite X fused; 5) parameres divided i n t o proxim a l sclerotized a n d distal m e m b r a n o u s regions; 6) phallobase Y - s h a p e d ; 7) parameres projecting anteriorly, b e n e a t h phallobase. Possible larval s y n a p o m o r p h i e s are: s u c k i n g m o u t h p a r t s f o r m e d by styliform m a n d i b l e s a n d maxillae enclosed in separate lateral channels; anterior pretarsi b i f u r cate; a n d l a b i u m f o r m i n g a five-toothed plate. T h e Holarctic g e n u s Cerophytum is characterized by h a v i n g the p r o f e m u r w i t h a l o n g i t u d i n a l carina a n d t h e dorsal region of t h e parameres f r i n g e d . Synapomorphies for t h e Neotropical clade include:

Cleide Costa,Sergio A.Vanin,John F.Lawrence and Sergio Ide posterior angles of p r o n o t u m reduced; u p p e r distal angle of p r o f e m u r acute a n d produced; a n d lateroposterior m a r g i n of phallobase p r o t r u d i n g over the bases of parameres. Brachycerophytum is d i s t i n g u i s h e d by t h e n i n t h elytral stria strongly convex, basal p o r t i o n of penis strongly constricted a n d b u r s a copulatrix w i t h o u t s m o o t h a n d elongate sclerites. A u t a p o m o r p h i e s of Phytocerum include the lack of a chin piece, presence of an additional row of p u n c t u r e s between stria 8 a n d 9, a n d t h e absence of a sclerotized spermatheca.

Acknowledgements TheRoyal Entomological Society of L o n d o n a n d Systematic E n t o m o l o g y are gratefully acknowledged for allowing us to reproduce copyrighted figures f r o m Costa etal. (2003). T h a n k s are d u e to Carlos E. Simonka for t h e final art a n d electronic t r e a t m e n t of Fig. 4.4.3. a n d to CNPq for t h e Research G r a n t 3 0 2 7 2 1 / 2 0 0 7 to C. Costa.

Literature Beutel, R. G. (1995): Phylogenetic analysis of Elateriformia (Coleoptera: Polyphaga) based on larval characters. -Journal ofZoological Systematics and Evolutionary Research 33:145-171. Buysson, H. du (1910): Materiaux pour servir ä l'histoire des insectes de l'aulne. - Annales de la Societe Entomologique de France 79:105-128. Calder, Α. Α., Lawrence, J. F. & Trueman, J. W. (1993): Austrelater, gen. nov. (Coleoptera: Elateridae) with a description of the larva and comments on elaterid relationships. - Invertebrate Taxonomy 7:1349-1394. Caveney, S. (1986): The phylogenetic significance of ommatidium structure in the compound eyes of polyphagan beetles. - Canadian Journal of Zoology 64:1787-1891. Costa, C., Vanin, S. Α., Lawrence, J. F. & Ide, S. (2003): Systematics and cladistic analysis of Cerophytidae (Elateroidea: Coleoptera). - Systematic Entomology 28: 375-407. Crowson, R. A. (1955): The Natural Classification of the Families of Coleoptera. 187 pp. Nathaniel Lloyd, London. Fleutiaux, E. (1896): Eucnemides nouveaux. -Memoires de la Societe Entomologique de France 9:300-315. - (1936): Les Elateridae de l'lndochine frangaise (6 lme partie). - Annales de la Societe Entomologique de France 105:279-300. Hlavac, T. F. (1975): The prothorax of Coleoptera (except Bostrichiformia - Cucujiformia). - Bulletin of the Museum of Comparative Zoology 147:137-183. Horion, A. (1953): Faunistik der mitteleuropäischen Käfer. Band III: Malacodermata, Sternoxia (Elateridae bis Throscidae). 280 pp. Museum G. Frey, Tutzing bei München. Johnson, P. J. (2002): 55. Cerophytidae Latreille 1834. Pp. 150-151 in Arnett, R. H., Jr., Thomas, M. C., Skelley, P. E. & Frank, J. H. (eds.) American Beetles. Volume2. polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Boca Raton, Florida.

Eucnemidae Eschscholtz, 1829 Lacordaire, J. T. (1857): Histoire naturelle des insectes. Genera des coleopteres ou expose methodique et critique de tous le genres proposes jusqu'ici dans cet ordre d'insectes, Vol. 4. 579 p p . Librairie Encyclopedique de Roret, Paris. Latreille, P. A. (1825): Families Naturelles du Regne Animal, Exposees Succinctement et dans un Ordre Analytique, avec Vindication de leurs Genres. 570 p p . J.-B. Bailliere, Paris. - (1834): D i s t r i b u t i o n m e t h o d i q u e de la famille des Serricornes. - Annales de la Societe Entomologique de France 3 : 1 1 3 - 1 7 0 . Lawrence, J. F. (1988): R h i n o r h i p i d a e , a n e w beetle family f r o m Australia, w i t h c o m m e n t s o n the p h y l o g e n y o f E l a t e r i f o r m i a .-Invertebrate Taxonomy 2(1987): 1 - 5 3 . - (1991): Cerophytidae (Elateroidea). Pp. 4 0 9 - 4 1 0 in Stehr, F. W. (ed.), Immature Insects, Vol. 2. Kendall/ H u n t P u b l i s h i n g Co., D u b u q u e , Iowa. Lawrence, J. F. & N e w t o n , A. F., Jr. (1995): Families a n d subfamilies of Coleoptera (with selected genera, notes a n d references, a n d data on family-group names). Pp. 7 7 9 - 1 0 0 6 in Pakaluk, J. & Slipiriski, S. A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: papers Celebrating the 80th Birthday of Roy A. Crowson. M u z e u m i Instytut Zoologii PAN, Warsaw. Lawrence, J. F., Hastings, A. M., Dallwitz, Μ. J., Paine, T. A. & Zürcher, Ε. J. (1999 a): "Beetle Larvae of the World: Descriptions, Illustrations, Identification, and Information Retrieval for Families and Subfamilies." CD-ROM, Version 1.1 for MS-Windows. CSIRO Publishing, Melbourne. - (1999 b): "Beetles of the World: A Key and Information System for Families and Subfamilies." CD-ROM, Version 1.0 for MS-Windows. CSIRO P u b l i s h i n g , Melbourne. Lawrence, J. F., M u o n a , J., Teräväinen, Μ., Stahls, G. & Vahtera, V. (2007): Anischia, perothops a n d the phylogeny of Elateroidea (Coleoptera: Elateriformia). - InsectSystematics and Evolution 3 8 : 2 0 5 - 2 3 9 . Lawrence, J. F., Nikitsky, Ν. B. & Kirejtshuk, A. G. (1995): Phylogenetic position of Decliniidae (Coleoptera: Scirtoidea) a n d comments on the classification of Elateriformia (sensu lato). Pp. 3 7 5 - 4 1 0 in Pakaluk, J. & Slipiriski, S. A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: papers Celebrating the 80th Birthday of Roy A. Crowson. M u z e u m i Instytut Zoologii PAN, Warsaw. Mamaev, Β. M. (1978): M o r p h o l o g y of the larvae of Cerophytum elateroides Latr. a n d t h e phylogenetic ties of the Cerophytidae (Coleoptera) family. - Doklady Akademii Νauk SSSR Zoologii 238: 1 0 0 7 1008 (in Russian). M u o n a , J. (1993): Review of t h e phylogeny, classification a n d biology of t h e family E u c n e m i d a e (Coleoptera). - Entomologica Scandinavica, S u p p l e m e n t 44:1-133. - (1995): The phylogeny of Elateroidea (Coleoptera), or which tree is best today? - Cladistics 1 1 : 3 1 7 - 3 4 1 . Rey, C. (1887): Essai d ' E t u d e s sur certaines larves des Coleopteres. A d d e n d u m . Larve s u p p o s e d u Cerophytum elateroides, Latreille. - Annales de la Societe Linneenne de Lyon (Ser. 2) 3 3 : 2 5 3 - 254. Schenkling, S. (1928): Coleopterorum Catalogus. Pars 101. Fam. Cerophytidae. 3 p p . W. J u n k , Berlin.

61

Steiner, W. E., Jr. (2000): Records a n d habitats of the 'rare click beetle', Cerophytumpulsator (HMeman), in Virginia a n d Maryland (Coleoptera: Cerophytidae). -Banisteria 1 5 : 4 3 - 4 5 . Zherichin, V. V. (1977): Mesozoic Coleoptera. Infraorder Elateriformia. Superfamily Cebrionoidea. Family Cerophytidae Latreille 1 8 3 4 . - T r u d y Paleontologicheski Instituta 1 6 1 : 1 3 0 - 1 3 4 .

4.5. Eucnemidae Eschscholtz, 1829 Jyrki M u o n a

Distribution.

E u c n e m i d a e are c o s m o p o l i t a n i n d i s t r i b u t i o n b u t m o s t a b u n d a n t a n d diverse in t h e t r o p i c s . S l i g h t l y less t h a n 2 0 0 g e n e r a a n d a b o u t 1 5 0 0 species are i n c l u d e d i n t h e f a m i l y . M a n y m o n o t y p i c g e n e r a , especially i n t h e s u b f a m i l y M a c r a u l a c i n a e , are b a s e d o n a u t a p o m o r p h i c f o r m s a n d m a y t u r n o u t t o b e s u p e r f l u o u s . T h e speciesr i c h w o r l d - w i d e g e n e r a Microrhagus D e j e a n , Fornax L a p o r t e , a n d Dromaeolus K i e s e n w e t t e r a p p e a r t o b e b a s e d o n p l e s i o m o r p h i c c h a r a c t e r s . M u c h of t h e diversity in tropical regions r e m a i n s undescribed a n d t h e t r u e n u m b e r of species is likely t o e x c e e d 3 0 0 0 . Anelastes Kirby, Melasis Olivier, Isoriphis B o i s d u v a l & L a c o r d a i r e , Hylochares L a p o r t e a n d

Fig. 4.5.1. Eucnemis americana (Horn), a d u l t , dorsal (from M u o n a 2000; © F i n n i s h M u s e u m of N a t u r a l History), l e n g t h = 6 m m .

Jyrki Muona

Fig. 4.5.2. Eucnemidae adults, dorsal: A, Anischia stupenda Fleutiaux (© CSIRO Australia), length = 3 mm; Β, Epiphanis cornutus Eschscholtz, length, = 5 mm; C, Schizophilus subrufus (Randall), length = 8 mm; D, Nematodes Humphreyi Muona, length = 10 mm; (B-D from Muona 2000; © Finnish Museum of Natural History).

Hylis de Gozis are typical Holarctic genera. Perothops Laporte, Schizophilus Bonvouloir, Onichodon N e w m a n and Palaeoxenus H o r n are Nearctic and Anelastidius DuVal is restricted to the Palaearctic. Anischia Fleutiaux, Dendrocharis Guerin-Meneville and Vitellius Bonvouloir represent Pantropical elements. Many clades suggest Gondwanan distributions and Arrhipis Bonvouloir, Entomosatopus Bonvouloir, Dyscharachthis Blackburn and Phaenocerus Bonvouloir have species on all Southern continents, whereas other putative Gondwanan clades have been split to two or three genera, and do not occur in south temperate regions (e. g., Poecilochrus Bonvouloir (Asia)+Idiotarsus Bonvouloir (Americas)). The Ethiopian fauna is characterized by many endemic species of Phyllocerus Lepeletier & Audinet-Serville. The Indomalayan region is very rich in species. The tribe Galbitini comprises 4 genera and about 50 species. The Australian fauna is phylogenetically diverse and rich with about 90 described species and about 90 undescribed species. Typical predominantly Australian genera include Hemiopsida MacLeay, Dyscharachthis Blackburn, and Euryptychus LeConte. The New Zealand fauna is phylogenetically poor with few species and include the d o m i n a n t endemic genera Talerax Sharp and Neocharis Sharp. [Schenkling 1928 a, b; Muona 1991 b, 1993 a, b; Alaruikka & Muona 2007; Muona & Briistle 2008.] Biology a n d Ecology. Adult Eucnemidae are rarely encountered although they can be regionally abundant, especially in the tropics. Most species develop in rotten wood and adults are usually encountered r u n n i n g on trunks and stumps, especially on broken surfaces. Many species are nocturnal and are attracted to light-traps and most are excellent fliers and often caught in Malaise traps. Most species of Eucnemidae click well although not as readily as most Elateridae. However,

Anischia species and some Melasini do not seem to click at all. Most species have sexually dimorphic antennae with several different types of receptors on the antennomeres and conspicuous excretory pits on the head, prosternum and hypomera (Fig. 4.5.3A). Chemical communication certainly plays an important role in their behavior. The well described swarming of Melasis buprestoides (Linnaeus) (Palmqvist 1952) is probably not typical of the family as a whole as observations of the mating in other species differ (Hylochares, Hylis, Dromaeoloides Fleutiaux, Microrhagus; J. Muona pers. obs.). Adult Eucnemidae are short-lived and it is unclear whether they feed. The few studied forms of Hylis and Xylophilus Mannerheim may lack a functional g u t altogether (Dodelin etal. 2005). Most Eucnemidae require forests with a good supply of dead wood and many are listed as endangered in heavily populated regions. Most eucnemid larvae are legless and lignicolous and the great majority do not construct galleries. Instead they move slowly with the help of their wedge-shaped head, internal hydrostatic pressure, and small densely spinose plates on their body. The spinose dorsal and ventral plates of the abdomen are pressed against the wood and at the same time the head is forced forward t h r o u g h the soft wood. Solid particles have not been f o u n d in the g u t and the larvae appear to obtain all their nutrition by sucking liquid f r o m the moist wood. Presumably this is a form of extra-oral digestion - the larvae vomit digestive juices in the wood in order to break down the fungal hyphae or possibly myxomycete Plasmodium. Eucnemidae larvae have roundish smooth plates b o t h dorsally and ventrally on most abdominal segments in addition to the spinose ones. These plates (usually referred to as areoles) function as organs for removing extra liquid f r o m the body. A few basal groups of Eucnemidae (e. g., Phyllocerus, Perothops) develop in soil. [Lucht 1981; H a m m o n d 1990;

Eucnemidae Eschscholtz, 1829

Penny & Arias 1992; Muona 1993 b, 2000; Muona & Brüstle 2008; Muona & Teräväinen 2008.] Morphology adults (Figs. 4 . 5 . 1 - 3 ) . Length 1.5 m m (Anischia, Microrhagus) to 4 0 m m (Phlegon). Body moderately elongate to very elongate, 2 - 6 times as long as wide, narrowing caudally or parallelsided; elytra slightly expanded basally and tapered apically; prothorax often curved laterally so that narrowest point is at pronoto-elytral junction (as in most Elateridae); moderately to strongly convex dorsally; ventrally either moderately convex or flat. Color frequently yellowish-brown to black and uniform (Fornax, Dromaeolus, Microrhagus), sometimes brightly bicoloured (Scython Laporte, Palaeoxenus, Spinifornax Fleutiaux), or even metallic (Nodema Fleutiaux). Vestiture rarely absent (Gastraulacus Guerin-Meneville), often consisting of fine decumbent, posteriorly oriented hairs (e. g., Fornax, Microrhagus); setae sometimes bicoloured (.Dromaeolus) or flat and scale-like forming striking dorsal patterns (Galbitini, Poecilochrus). Head strongly transverse, slightly to strongly declined, deeply inserted into prothorax, not abruptly narrowed posteriorly to form neck. Posterior edge bi-emarginate, forming median tooth, with or without median carina. Fine transverse occipital ridge usually present, often continuing below eyes as weak subgenal ridges. Median endocarina absent. Eyes highly variable, small to large, entire or almost so, flat to protruding, finely facetted. Antennal insertions exposed and usually separated by less than length of antennomere 1 but sometimes much more so (Dendrocharini, Gastraulacini); subantennal grooves variable, from absent to short and shallow to wide and deep with either group of pores or deep pits. Frontoclypeal area gradually declined, with mouthparts anteroventrally or ventrally oriented; frontoclypeal suture absent; anterior edge of clypeus mostly broadly rounded concave or bisinuate, rarely truncate. Labrum attached beneath edge of clypeus, membranous (many species) or partly visible, slightly sclerotized and densely covered with hairs (Melasini), or partly visible, strongly transverse, rounded anteriorly and with distinct median emargination (Anischia). Antennae usually 11-segmented (12-segmented in Phyllocerus males); variable in length, not extending to base of prothorax (many Melasinae) to longer than body (Plesiofornax Coquerel); antennomeres moniliform (many Macraulacini) to strongly flabellate (e. g., Galbites Fleutiaux), sometimes gradually expanded apically, incrassate or clavate, or more abruptly expanded to form weak 2- to 5segmented club (Anischia, Anelastes, and many Melasinae); scape always inflated; pedicel often tiny, attached subapically with sharp tooth just laterad of attachment; antennomere 3 usually distinctly elongate; sensory elements either evenly distributed (Anelastes) or present only on apical antennomere and at or around apex on others. Mandibles variable, either evenly curved and elongate or short

63

and broad, only slightly longer than wide at base, unidentate or bidentate; mola absent or reduced; if present sub-basal, consisting of group of asperities several of which form a transverse, comb-like structure; distinct hyaline area present at base of mesal edge; prostheca absent. Maxilla usually with unequal lobes; galea articulated; lacinia narrowed apically; terminal maxillary palpomere variable, often slightly widened at middle, narrowed apically and obliquely truncate at apex; frequently more strongly developed in males than in females. Labium with truncate ligula; terminal palpomere similar to that of maxillary palp but not sexually dimorphic. Rarely ventral mouthparts grotesquely developed, divided into dozens of branch-like structures (Cladus Bonvouloir, Hyperpalpus Lucht). Gular sutures variable, often widely separated. Corpotentorium very narrow, slightly arched; anterior tentorial arms expanded mesally and sometimes fused at midline to form broad anterior bridge (laminatentorium). Cervical sclerites well developed, each divided into two or three parts. Proventriculus variable from weakly developed (Perothops) to consisting of setose pads alternating with elongate hyaline processes lined with saw-like teeth. Pronotum variable, wider than long to more than twice as long as wide, usually widest at about middle; sides parallel to strongly rounded. Lateral carinae absent (Langurioscthton Heller), very fine (Anelastes), divided (most Dirhagini), or strongly developed and entire (most species), sometimes not visible for their entire lengths from above; anterior angles absent to right. Anterior edge simple or minutely serrate (Dirhagini). Posterior angles acute and mostly produced. Hind edge with well-developed interlocking device, sometimes including pair of deep sublateral cavities for receiving paired processes on anterior edges of elytra (Anischia). Disc usually convex or flattened and slightly grooved in front of scutellum; sometimes with large gibbosities (e. g., Galbites), rarely with two pairs of longitudinal carinae extending anteriorly from hind edge (Anischia). Hypomeron without antennal grooves or cavities in Phyllocerus, Perothops, Anelastes, Pseudomeninae and many Melasinae, with deep antennal grooves running along lateral edge in Eucneminae and Macraulacini (rarely wider than rest of hypomeron in some Macraulacini), with variablyshaped antennal grooves running across notosternal suture in Dirhagini (Fig. 4.5.3 A). Posterior surface of hypomeron usually with crural impression (often separated by carina from rest of hypomeron) for reception of profemora. Notosternal suture complete or partly to almost entirely obliterated (some Dirhagini). Prosternum well developed in front of coxae, moderately convex and produced anteriorly to form broad chin-piece; prosternal process variable, short and poorly developed (e. g., Hemiopsida) to moderately long and straight or parallel-sided except posteriorly, where sides taper to form subacute apex; about as wide as coxal cavity (many species) to long and knife-like; surface either

64 smooth or with median and/or lateral carinae. Procoxae globular, with very short internal extension. Trochantinopleuron reduced, concealed and fused to wall of hypomeron. Procoxal cavities moderately broadly open; notal projections short and subacute. Well developed scutellar shield either flat or abruptly elevated basally, with straight basal edge and parallel sides or with rounded lateral edges and subacute apex. Elytra 1.2 to 4 times as long as wide, subparallel to tapering posteriorly; anterior edge of each more or less carinate and produced at middle to form carinate lobe fitting into cavity at base of pronotum; fine sutural stria usually present and extending almost to apex; scutellary striole absent; other striae absent to well-developed and deep ('Vitellius and other taxa), with interstices flat to convex and odd ones sometimes keeled and/or wider than even ones (Galbites and other taxa); punctation absent to present and doubled, confused or seriate; epipleura moderately to strongly developed, anteriorly sometimes grooved (some Macraulacini and Eucneminae); usually tapering posteriorly and reaching elytral apex, sometimes with apical portion vertical. Mesoventrite longer than wide to distinctly wider than long; anterior edge at middle with deep notch bordered by a raised lip that is sometimes flanked by a pair of large, shallow, horizontal procoxal rests. Anterior lip contiguous posteriorly with diagonal slide leading into a deep mesoventral cavity extending well beyond the anterior edges of the mesocoxae. Mesocoxae globular, separated by a distance from about half the longest diameter of one (Perothops) to slightly more than one diameter. Mesocoxal cavity laterally open (partly closed by mesopleuron and not by meeting of meso- and metaventrites). Mesanepisternum separated from mesepimeron by complete pleural suture (Perothops) or solidly joined to mesanepimeron with no trace of pleural suture or pleural ridge. Meso-metaventral junction distinctly sinuate to straight, with metaventral knob fitting into cavity on mesoventrite. Metaventrite higly variable, from strongly transverse to more than twice as long as wide; without discrimen. Postcoxal lines usually absent (Anischia with two postcoxal lines arising from posterior edge of each mesocoxal cavity: one beginning at posteromesal edge of cavity and extending posterolaterally and the other beginning j ust behind mesepimero-metanepisternal j unction, extending mesally and then abruptly posteriorly near lateral edge of cavity); sometimes with distinct grooves for reception of tibiae (many Eucneminae) and tarsi (Gastraulacus etc.). Visible portion of metanepisternum variable; usually very narrow and more or less parallel-sided, sometimes wide and short or strongly widening caudad; anterior edge distant from mesocoxal cavity. Metacoxae slightly oblique, usually contiguous and extending laterally to meet epipleura (in Anischia well separated, extending laterally almost to epipleura but separated from them by posterior portions of metanepisternum and metepimeron; coxal plates

Jyrki Muona

usually well developed and complete but narrowed laterally, sometimes parallel-sided (e. g., Rhagomicrus Fleutiaux, Dendrocharis), rarely completely absent (Anischia). Metendosternite usually with moderately to very long, slender stalk and short lateral arms, each bearing a long tendon; anterior process usually long and narrow, with more or less approximate anterior tendons; sometimes short, broad and bilobed, with moderately widely separated t e n d o n s (e. g., Perothops a n d Phyllocerus); in

Anischia, stalk short and broad; anterior process absent and arms long and anteriorly oblique with tendons located near apices. Hind-wing short, about 2 times as long as wide (Farsus Jacquelin DuVal, Temnus Fleutiaux) or as much as 3 times as long as wide (Hylotastes Bonvouloir); apical field well developed, about 0.4 times total wing length (Anischia, Porraulacus F l e u t i a u x , Proxylobius

Fleu-

tiaux, etc.) or reduced, about 0.15 times total wing length; with one to five oblique sclerites, (usually with an anterior, posterior, and median one); radial cell 1.5-5 times as long as wide with posterobasal angle more or less right or rounded; cross-vein r3 usually very slightly oblique and almost longitudinal, sometimes absent; cross-vein r4 arising towards apex of cell, long and slightly sinuate; basal portion of RP variable. Medial field usually with five free veins all reaching wing margin, sometimes with three or four veins not reaching wing margin, without veins in Porraulacus; MP3+4 with well developed basal cross-vein and usually with CuAj joining it before MP3-MP4 fork; base of MP3 incomplete; wedge cell either absent or present and often longer than medial spur, 1.5 to 4 times as long as wide, with apex oblique; CuA1+2 arising near apical third of cell and often slightly longer than CuAj; AA3 meeting CuP near base of wedge cell or absent; anal notch present or absent. Legs stout to long and slender; hind legs somewhat longer than anterior pairs; trochanter rarely very long, almost half as long as femur in Anischia, usually much shorter (especially protrochanter); trochanterofemoral joint oblique; tibiae variable, often relatively slender, only slightly enlarged apically, rarely strongly flattened (Melasis) or flattened and with tarsal grooves (Dendrocharini); protibial apex usually with one spur, occasionally with two spurs of equal l e n g t h (e. g., Perothops, Phyllocerus, Anischia); tarso-

meres variable: either simple with 1-4 decreasing in length and 5 usually as long as previous two or three combined (many Dirhagini), or tarsomeres 1 4 densely pubescent and 5 as long as 1 - 4 (many Eucnemini), or tarsomeres 1-4 with distinct lobes (Galbitini, Dendrocharini); pretarsal claws often simple or basally dentate (many Macraulacini) or serrate (Perothops), rarely with basal setae (Pseudomenes Fleutiaux); empodium weakly developed but usually not visible beyond apex of tarsomere 5; protarsus often sexually dimorphic: male protarsomere 1 with complete or basal sex-comb (Macraulacini; Fig. 4.5.3 B) or with apical sex-comb (Dirhagini).

Eucnemidae Eschscholtz, 1829

Abdomen slightly to very elongate, with five ventrites. Ventrites 1 - 4 often subequal in length; 5 distinctly longer; all ventrites usually connate; in Anischia, ventrites 1 - 3 connate, 4 and 5 movable, and ventrite 1 with two postcoxal lines on each side of intercoxal process; in Perothops, ventrite 1 somewhat longer than 2 with acute intercoxal process, 2 - 4 slightly decreasing in length, 5 somewhat longer t h a n 4, ventrites 1 and 2 distinctly connate, 3 - 5 at least slightly movable and posterolateral corners of 2 - 4 produced and acute. Abdomen rarely with grooves for reception of tarsi (Gastraulacus, Temnus, Temnillus Bonvouloir). Spiracles on segments I to VIII located in pleural membrane. Tergites often membranous b u t sometimes all tergites well sclerotized (Dromaeolus, Asiocnemis Mamaev, etc.) or VIII-X lightly sclerotized (Anischia). Sternite VIII in male with paired lateral struts. Sternite IX in male with paired lateral struts only (Anischia, Temnillus, etc.) or with broadly rounded anterior lobe and usually with sclerotized border; tergites IX and X in male clearly separate (Perothops, Phyllocerus, etc.) or more or less fused together. Sternite and tergite VIII in female variable, often lightly sclerotized; spiculum ventrale absent (Perothops, Phyllocerus, etc.) or well developed and long, either basally articulated (Pseudomenes, Schizophilus, Anischia) or solidly fused (most taxa). Aedeagus of the trilobate type b u t quite variable. Several different types may be distinguished: 1) phallobase subquadrate, slightly flattened and dorsally open (sclerotized ventrally), symmetrical, between one-third and two-thirds as long as parameres which are broadly fused dorsally and narrowly so ventrally, ventrally each paramere tapering anteriorly, to form acute, dorsally or laterally curved strut (many Melasinae and Macraulacinae), divided penis with short or long basal struts (many Melasinae, all Macraulacinae); 2) phallobase slightly flattened anteriorly, with paired ventral struts and a dorsal process which is fused to parameres (Perothopinae and Phyllocerini); 3) phallobase divided in two lateral sclerites; parameres and penis strongly dorsoventrally flattened (Anelastini); 4) phallobase reduced, elongated, rod-shaped, resembling letter Y, parameres and penis slender, parameres soft, densely hairy mediad (Pseudomenes Bonvouloir); 5) phallobase short, asymmetrical, laterally compressed, about a third the length of parameres which are fused together at basal half to form a tube; apex of parameres narrowly rounded truncate or slightly expanded to form lateral tooth; penis relatively short, with body about half as long as parameres b u t with paired anterior struts which may extend almost to base of parameral tube; penis attached to parameres at point where parameral tube ends and free parameres begin (Anischia); 6) parameres and penis fused together forming a tube, phallobase well developed (Dirhagini), or strongly reduced (Dendrocharini); 7) penis shield-like and parameres apparently very small (most Eucneminae), rarely also phallobase reduced and membranous (Temnillus etc.) or penis divided in dorsal and

65

ventral lobes (Entomosatopus Bonvouloir). Ovipositor of three major types: 1) short and sclerotized and lacking distinct segments (e.g., Sarpedon Bonvouloir); 2) short and sclerotized and divided in two parts with styli absent (Hylochares, etc.) or present, 3) ovipositor moderately long and slender (most

Fig. 4.5.3. Eucnemidae adults: A, Microrhaguspygmaeus (Fabricius), ventral view of head and prothorax showing notosternal antennal grooves and excretory pits close to eye and on hypomere; B, Dicaptothorax koebeli (Blackburn), male protarsus showing complex sexcomb on base of tarsomere 1.

66

Jyrki Muona

Qu ts

Μ

Wm A

m

1

m:;·

I

a;

• i

μ • β

Fig. 4.5.4. Eucnemidae larvae: A, Asiocnemis morawitzi Semenov, ventral; B, Euryptychus lewisi Fleutiaux, dorsal; C, Thambus friwaldskyi Bonvouloir, ventral; D, Eucnemis zaitzevi Mamaev, dorsal; E, Isorhipis melasoides (Laporte), dorsal. (A-E from Mamaev 1976).

taxa) to longer than body and very slender (e. g., Nematodini, Hylis); paraprocts with longitudinal bacula 1.5 to 8 times as long as coxites which are membranous to sclerotized, narrowed apically and either undivided or divided into two parts; styli well developed apically (Perothopinae and Phyllocerinae) or laterally attached and often minute. Female genital tract variable, with three major types: 1) with large bilobed bursa with weakly sclerotized and elongate spermatheca attached by short duct between bases of two bursal lobes, and spermathecal gland attached by long narrow duct to base of spermatheca (Perothopinae, Phyllocerinae, Pseudomeninae, Phelgoninae); 2) bursa undivided with spermatheca well sclerotized and ribbed, undivided or divided into 2 - 3 parts (Melasinae, Eucneminae, Macraulacinae); 3) genital tract enlarged anteriorly to form an elongate to almost spherical uterus to which the common oviduct and spermathecal duct are separately attached; bursa copulatrix absent; spermatheca elongate and

cylindrical, finely transversely ribbed with a basal collar and apical invaginated pocket; spermathecal gland attached basally just beyond the collar. [Burakowski 1991; Lawrence et al. 2007; Muona 1991 b, 1993 b.] Description Larvae (Figs. 4.5.4-6). Body elongate, usually parallel-sided, sometimes widest at metathorax (Perothops, Melasini, Hylochares); moderately to strongly flattened (most) or subcylindrical (Perothops, Melasini, some Dirrhagini), rarely hypermetamorphic, swollen and maggot-like (some Eucnemini). Evenly soft or well sclerotized, rarely with dorsal surfaces more heavily sclerotized than ventral ones (Anischia, Phyllocerus). Color of head and apex of last tergum often darkened, rest of body white to yellowish, rarely very dark brown. All segments nearly equal in size or abdominal segments transversely subdivided forming pseudosegments (Melasini), or thoracic segments and abdominal segments I-VI more or less inflated at middle and strongly narrowed at either

67

Eucnemidae Eschscholtz, 1829

end (Perothops); surfaces of terga either smooth or with fine reticulation or rugosity. Vestiture consisting of a few macrosetae and numerous microsetae (some forming transverse rows) and dorsal and/or ventral patches of dense spicules and microtrichia (Fig. 4.5.5). Head usually, b u t not always, forming sclerotized, serrate, wedge-like plate that is horizontal and protracted; much wider than long or longer than wide and widest at base, narrowing anteriorly with broadly truncate anterior edge; tapering from base to apex in lateral view; head rarely with deep longitudinal grooves (Pseudomenes, Schizophilus Fig. 4.5.6 A). Stemmata almost always absent (one small stemma on each side in Anischia). Posterior edge of head with moderately broad mediodorsal emargination flanked by two narrower emarginations or not emarginate. Epicranial stem very short or absent; frontal arms absent. Paired endocarinae present or absent. Labrum entirely fused to head capsule with no indication of a clypeo-labral suture, or separated from head capsule by complete suture (Phyllocerus, Perothops, and Pseudomenes); clypeolabrum usually not projecting, rarely forming a nasale (Perothops). Antennae short and 2-segmented with large sensorium; often located in cavities with their insertions concealed from above. Mandibles usually entirely fused to head capsule (Fig. 4.5.6 B-C) or nearly so; occasionally movable and opposable (Fig. 4.5.6 A); mandibles in Anischia broad at base, abruptly narrowed and abruptly curved at basal third, straight apically with a subacute apex and grooved at mesal edge; those in Perothops about as long as head capsule, symmetrical, stout, about twice as long as wide at base, very slightly curved mesally b u t not or barely meeting at midline with mandibular apex forming single narrowly rounded lobe, incisor edge with short apicomesally projecting retinaculum and mesal surface at base with brush of hairs but no mola; and those in Phyllocerus similar b u t wider at base, narrow, falcate and grooved. Maxilla without apical lobes (most taxa) or with mala (Perothops, Phyllocerus); palps 2 - 4 segmented or absent. Labium with short ligula (Phyllocerus) or ligula absent; palps 1 - 2 segmented, widely (Anischia, Phyllocerus) or narrowly separated. Postmentum and maxillae elongate, usually partly connate, or entirely so and extending almost to base of head. Prothorax usually slightly shorter and narrower than meso- or metathorax (clearly so in Melasini and Hylochares); pro-, meso-, and metaterga usually with variably-sized sclerotized plates or t-shaped rods. Legs always highly reduced, almost always reduced to small dark spots with or without seta (Fig. 4.5.6 B). In Perothops legs articulated and 5-segmented, with short coxa, broad dome-like trochanter, and short, broad, disk-like femur less than half diameter of coxa, tibiotarsus about as long as wide, pretarsus longer than remaining segments combined, slender, without apparent setae. In Phyllocerus legs similar to those in Perothops b u t with pretarsus bearing well-developed setae. Legs in Anischia, Schizophilus and Psedudomenes reduced to single short and broad

segment bearing three or four long stout setae and several shorter apical setae. Abdominal segments I VIII subequal in length; apparently divided in two in Melasini. Segment IX shorter or longer than previous ones, often rugose and dark in color, caudally with asperities or rounded or with tiny urogomphi (Hylis, Fornax etc) or simple or bifid large urogomphi (Euryptychus, Pseudomenes, Schizophilus). Abdominal segments I-VI (and meso- and metathorax) of all derived forms with variably placed single or divided plates of two types: those with densely set small spicules and those which are entirely smooth and hymen-like (areoles, Fig. 4.5.5); Perothops with paired and complexly rugose areas on dorsum separated by longitudinal rugulae and bordered anteriorly and posteriorly by transverse rugulae; Anischia with longitudinal rugulae on lateral regions, venter with rugulose areas more or less similar to those on dorsum, surfaces of segments VII and VIII with simpler finely transversely rugulose, and segments with large spicule patches but no smooth plates. Segment IX ofPeroiftopswithshortstrongly upturned, approximate urogomphi about 0.1 times as long as basal width of segment. Segment X elongate oval, without teeth or asperities. Spiracles small or mediumsized, biforous with closing apparatus surrounded by sclerotized ring that is either rounded or with a pointed caudal edge; thoracic spiracle either equal in size to others or larger than those on abdomen; its sclerite either rounded or vertically oriented and slightly obliquely oval; abdominal spiracles with circular or oval sclerite located laterally in a membrane. [Boving & Craighead 1936; Burakowski 1991; Gardner 1935,1936; Ghilarov 1979; Lawrence etal. 2007; Leiler 1976; Otto & Young 1998; Muona & Teräväinen 2008.] Phylogeny and Classification. Eucnemidae includes two major groups, one with soil-dwelling larvae (Perothopinae and Phyllocerinae) and another more extensive radiation with lignicolous larvae (Pseudomeninae, Palaeoxeninae, Phlegoninae, Anischiinae, Melasinae, Eucneminae, and

ΜΗκ Fig. 4.5.5. Onichodon canadensis (Brown), larva, abdominal segment V with triangular spinose patch and small wide smooth plate (areole) (from Muona & Teräväinen 2008; © Μ. Teräväinen).

68

Jyrki Muona

Fig. 4.5.6. Eucnemidae larvae: A, Pseudomenes bakewelli (Blackburn), head, dorsal, showing inward biting welldeveloped mandibles and articulating labrum; B, ?Jenibuntor sp., head and portion of mesothorax, ventral, showing nasale-like projection (A), fused postmentum and maxillae (B), and tiny spot-like legs (C); C. Prodirrhagus sp., head, dorsal, showing tiny immobile mandibles and no trace of labrum or nasale;

Macraulacinae). This is essentially t h e general taxon o m i c view held by Lameere (1900), Crowson(1955, 1960), Cobos (1964), Lawrence (1987) a n d M u o n a (1993). T h e placement of Anischia in E u c n e m i d a e is based on a recent analysis i n c l u d i n g t h e newly discovered larva a n d molecular d a t a (Lawrence etal. 2007). A l t h o u g h there is little d i s a g r e e m e n t a b o u t t h e composition of this clade, its position w i t h other elateroids is still o p e n a n d even t h e m o n o p h y l y of t h e clicking elateroids requires f u r t h e r s t u d y (e. g., Lawrence 1988; Beutel 1992; Lawrence et al. 1995; M u o n a 1995; Calder et al. 1996; Bocakova et al. 2007). O n g o i n g analyses a n d m o r e molecular data will h o p e f u l l y clarify t h e situation. T h e E u c n e m i d a e classification is based o n recent phylogenetic analyses (Muona 1991,1993; Lawrence etal. 2007). Several of t h e traditional e u c n e m i d features have been s h o w n to be incorrectly described in literature. Most species click well a l t h o u g h t h e ability has b e e n lost i n d e p e n d e n t l y in at least two lineages (Anischia, Echtrogasterini). U n r e l a t e d genera s h o w a partly visible l a b r u m in adults (Anischia, Melasini) a n d t h e larval l a b r u m has b e e n f u s e d i n d e p e n d e n t l y at least two times (Schizophilus a n d all derived forms). Fossil material shows t h a t m o s t of t h e e x t a n t h i g h e r clades were present d u r i n g t h e Baltic Amber period a n d large shifts in d i s t r i b u t i o n have t a k e n place later o n a l t h o u g h t h e relative diversity of t h e m a i n subfamilies has r e m a i n e d the same (Muona 1993 a).

Acknowledgements Grants f r o m t h e F i n n i s h Academy, t h e E u r o p e a n U n i o n ("Hotspots"- consortium), t h e Helsinki Entomological Society a n d t h e F i n n i s h E n t o m o logical Society are gratefully acknowledged. T h e following artists are acknowledged: A. Smatana (Figs. 4 . 5 . 1 , 2 B-D) a n d S. P. Kim (Fig. 4.5.2 A).

Literature Alaruikka, D. & Muona, J. (2007): World Check-List of Eucnemidae. Database Finnish Museum of Natural History Helsinki. Beutel, R. G. (1995): Phylogenetic analysis of Elateriformia (Coleoptera: Polyphaga) based on larval characters. - Journal of Zoological Systematics and Evolutionary Research 33:145-171. Bocakova, M., Bocak, L., Hunt, T., Teraväinen, Μ. & Vogler, A. P. (2007): Molecular phylogenetics of Elate-riformia (Coleoptera): evolution of bioluminescence and neoteny. - Cladistics 23:477-496. Boving, A. G. & Craighead, F. C. (1931): An illustrated synopsis of the principal larval forms of the order Coleoptera. - Εntomologica Americana (N.S.) 11(1930): 1 - 3 5 1 . Burakowski, B. (1991) Klucze do Oznaczania Owadow Polski. CzescXIX. Chrzaszcze - Coleoptera. Zeszyt 35-37. CerophytidaeEucnemidaeThroscidaeLissomidae. 91 pp. Polskie Towarzystwo Entomologiczne Warsaw. Calder, Α. Α., Lawrence, J. F. & Trueman, J. W. H. (1993): Austrelater gen. nov. Coleoptera: Elateridae) with a description of the larva and comments on elaterid relationships. - Invertebrate Taxonomy 7:1349-1394. Cobos, A. (1964): Materiales para el estudio de la familia Eucnemidae. Primera parte. Eos 40:289-435. Crowson, R. A. (1955): The Natural Classification of the Families of Coleoptera. 187 pp. Nathaniel Lloyd London. - (1960): The Phylogeny of Coleoptera. - Annual Review ofEntomology 5:111-134. Dodelin, B. Pene, B. & Andre, J. (2005): L'alimentation des coleopteres saproxyliques et notes sur les contenus stomacaux de cinq especes. - Bulletin Mensuel de la Societe Linneenne de Lyon 74 (10): 335-345. Eschscholtz, F. (1829): Zoologischer Atlas enthaltend Abbildungen und Beschreibungen neuer Thierarten während des Flottcapitains von Kotzebue zweiter Reise um die Welt auf der Russisch-Kaiserlichen Kriegsschlupp Predpriaetiein den Jahren 1823-1826. Erstes Heft. 17 pp. G. Reimer, Berlin.

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Throscidae Laporte, 1840 Gardner, J. C. M. (1935): Immature stages of Indian Coleoptera(l 7) (Eucnemidae). - Indian Forest Records (New Series) Entomology 1 (4): 79-93. - (1936): A larva of the subfamily Balginae (Col. Elateridae). - Proceedings of the Royal Entomological Society ofLondon (B) 5:3-5. Ghilarov, M. S. (1979): A larva of Phyllocerus and the position of this genus in the system of Elateroidea (Coleoptera). - Zoologicheskii Zhurnal 58: 655-663 (in Russian). Hammond, P. M. (1990): Insect abundance and diversity in the Dumoga-Bone National Park N. Sulawesi with special reference to the beetle fauna of lowland rain forest in the Toraut region. Pp. 197-254 in Knight W. J. & Holloway J. D. (eds.) Insects and the Rain Forests ofSouth East Asia (Wallacea). The Royal Entomological Society of London, London. Lawrence, J. F. (1988): Rhinorhipidae a new beetle family from Australia with comments on the phylogenyofElateriformia .-Invertebrate Taxonomy 2(1987): 1-53. Lawrence, J. F. & Newton, A. F. Jr. (1995): Families and subfamilies of Coleoptera (with selected genera notes references and data on family-group names). Pp. 779-1006 in Pakaluk, J. & Slipiriski, S. A. (eds.) Biology Phylogeny and Classification of Coleoptera: papers Celebrating the 80th Birthday of Roy A. Crowson. XII + 1092 pp. Muzeum i Instytut Zoologii Polska Akademia Nauk, Warsaw. Lawrence, J. F.Nikitsky,N.B.&Kirejtshuk, A. G. (1995): Phylogenetic position of Decliniidae (Coleoptera: Scirtoidea) and comments on the classification of Elateriformia (sensu lato). Pp. 375-410 in Pakaluk J. & Slipiriski, S. A. (eds.) Biology Phylogeny and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii Polska Akademia Nauk Warsaw. Lawrence, J. F., Hastings, A. M., Dallwitz, Μ. J., Paine, T. A. & Zürcher, Ε. J. (1999): Beetles of the World: A Key and Information System for Families and Subfamilies. CD-ROM Version 1.0 for MS-Windows. CSIRO Publishing Melbourne. Lawrence, J. F., Muona, J., Teräväinen, Μ., Stahls, G., Vahtera, V. (2007): Anischia, Perothops and the phylogeny of Elateroidea. - Insect Systematics & Evolution 38: 205-239. Leiler, T.-E. (1976): Zur Kenntnis der Entwicklungsstadien und der Lebensweise nord- und mitteleuropäischer Eucnemiden (Col.). - Entomologische Blätter 72(1): 10-50. Lucht, W. (1981): Die Präimaginalstadien von Hypocaelus olexai Palm (Col. Eucnemidae) nebst Bestimmungstabelle der Larven Nord- und Mitteleuropäischer Hypocaelus-Arten. - Entomologische Blätter für Biologie und Systematik der Käfer 77 (1-2): 61-74. Mamaev, Β. M. (1976): Morphological types of xylophagous beetle larvae (Coleoptera, Eucnemidae) and their evolutionary importance. Pp. 136-155 in Mamaev, Β. M. (ed.) Evolutionary Morphology ofWoodBoring Larvae. Nauka, Moscow. Muona, J. (1991 a): The Eucnemidae of South-east Asia and the western Pacific - a biogeographical study. - Australian Systematic Botany 4:165-182.

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(1991 b): A revision of the Indomalesian tribe Galbitini new tribe (Coleoptera Eucnemidae). - Entomologica Scandinavica, Supplement 39: 1-67, 2 pis. - (1993 a): Eucnemidae and Throscidae in Baltic Amber. - Entomologische Blätterfür Biologie und Systematik der Käfer 89:15-45. - (1993 b): Review of the phylogeny, classification and biology of the family Eucnemidae (Coleoptera). - Entomologica Scandinavica, Suppplement 44,133 pp. - (1995): The phylogeny of Elateroidea (Coleoptera) or which tree is best today? - Cladistics 11:317-341. - (2000): A revision of the Nearctic Eucnemidae. - ActaZoologicaFennica 212,106 pp. Muona, J. & Brüstle, L. (2008): Observations on the biology of Hylochares cruentatus (Gyllenhal) (Coleoptera: Eucnemidae). - Entomologica Fennica 19:151-158. Muona, J. & Teräväinen, Μ. (2008): Notes on the biology and morphology of false click-beetle larvae (Coleoptera, Eucnemidae). - The Coleopterists Bulletin 62 (4): 475-479. Otto, R. L. & Young, D. K. (1998): Description of the larva of Schizophilus subrufus (Randall) (Coleoptera: Eucnemidae: Pseudomeninae) with notes on its natural history. - The Coleopterists Bulletin 52 (4): 306-312. Palmqvist, S. (1952): Nagra iakttagelser over förplantingsbiologin hos Melasis buprestoides L. (Col. Eucnemidae). - Opuscula Entomologicae 17 (1-2): 70-75. Penny, N. D. & Arias, J. R. (1982): Insects of an Amazon forest. 320 pp. Columbia University Press, New York. Schenkling, S. (1928 a): Coleopterorum Catalogus. Pars 96. Melasidae. 110 pp. W. Junk Berlin. - (1928 b): Coleopterorum Catalogus. Pars 101. Throscidae CerophytidaePerothopidae. 30 pp. W. Junk Berlin.

4.6. Throscidae Laporte, 1840 Jyrki Muona, John F. Lawrence and Adam Slipiriski Distribution. Throscidae are cosmopolitan in distribution with the greatest diversity in warm temperate and subtropical regions. They do not occur in New Zealand. Five extant genera and about 150 species are included in the family. Several new species have been described from well-known European regions recently and much of the diversity in less well-known areas may remain undescribed. Potergus Bonvouloir with a few described and undescribed species is widespread in the Asian, Australian and Pacific tropics and the monotypic Pactopus LeConte is restricted to Western North America. Trixagus Kugelann and Aulonothroscus Horn are species-rich world-wide genera, the latter being more diverse in subtropical and tropical regions. Fossil evidence suggests that the structure of the diversity

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Jyrki Muona, John F.Lawrence and Adam Slipinski

of the family has changed since the formation of Baltic Amber. Instead of two dominant genera, several about equal-sized ones seem to have existed. One of them is Pactopus, which had several now extinct species in the Old World, coexisting with Trixagus and Aulonothroscus. In addition to these extant genera a further, now extinct genus, Jaira Muona, is known from this fossil material. [Schenkling 1928; Yensen 1975 a, b; Muona 1993; Leseigneur 1998.] Biology and Ecology. Adult Trixagus and Aulonothroscus are frequently found in numbers in open, vegetated habitats. Trixagus dernestoides larvae develop in soil, sucking liquid from ecotrophic mycorrhizae attached to trees (e. g., Betula, Alnus and Ulmus). Adult beetles emerge early in spring and mate at dusk. Males have a complex mating behaviour. They court females by dancing around them, tapping with antennae and vibrating both the elytra and the alae. Copulation seems to take place on twigs or branches, even though the larvae develop in soil. Aulonothroscus species have been collected in hollow trunks and flight-intercept traps placed in them in order to catch lignicolous beetles. Their larvae have never been clearly associated with adults, which are less frequently collected than those of Trixagus species and probably live in soft, rotten wood. Larvae illustrated by Becker (1991) from decaying pine boards in Virginia and very similar larvae from red-rotten conifer logs in Vermont may well belong to Aulonothroscus. Pactopus pupae have been found under a log in a burnt forest area; little is known of Potergus except that they are repeatedly collected in rainforest fogging studies and have been seen running on logs. Most throscid species are nocturnal and are attracted strongly to light-traps. They are excellent fliers and are often encountered in Malaise trap catches as well. Trixagus spp. can be netted from low vegetation in numbers from late afternoon to early evening. When caught, they feign death for a considerable time, giving the impression of being dry seeds. Typically they become active all at the same time in the net, release antennae and legs from the protective furrows and start running with considerable speed. Members of all genera have been observed to click and they do it well, although not as readily as the Elateridae. Adult throscids are short-lived and their food remains unknown, but it has been shown that several different micro-organisms live in mycotomes in their fat bodies. [Burakowski 1975.] Morphology, Adults. (Figs. 4.6.1, 2A, B). Length 1.2-6.0 mm. Body usually oblong to slightly elongate, about 2.1-2.6 times as long as wide (narrowly elongate, up to 3.5 times as long as wide in Potergus), usually widest at junction of prothorax and elytra and tapering anteriorly and posteriorly (sides subparallel in Potergus); slightly flattened to moderately convex dorsally, distinctly convex ventrally. Color uniformly reddish-brown to black. Densely clothed with decumbent hairs, usually

Fig. 4.6.1. Aulonothroscus sp., adult, dorsal (from Lawrence & Britton 1991; © CSIRO Australia), length = 2.8 mm.

interspersed with suberect hairs, sometimes with silvery sheen. Head transverse, moderately to strongly declined, not abruptly constricted posteriorly and deeply inserted into prothorax. Posterior edge above occipital foramen not to slightly biemarginate, without median endocarina. Transverse occipital carina usually present. Frontal region convex and more or less declined, with long median carina in Potergus, sometimes with paired supraocular carinae. Eyes well developed, not protuberant, usually entire (moderately to deeply emarginate in many Trixagus); finely facetted, without interfacetal setae; ommatidium of exocone type with reduced clear zone (Caveney 1986). Sometimes with one to three deep pits in vicinity of eye. Antennal insertions moderately widely separated and exposed, more narrowly separated and located within circular fossae in Potergus. Subantennal grooves well developed. Frontoclypeal suture absent; clypeal apex more or less truncate. Labrum small, strongly transverse, well sclerotized, free and exposed, its anterior edge usually slightly to strongly convex. Antennae 11-segmented, usually with symmetrical to strongly asymmetrical, 2- to 5-segmented club (filiform to slightly incrassate in Pactopus and most Potergus); pedicel relatively wide, usually as wide as apex of scape. Mandible short and broad, somewhat flattened, unidentate; mola

Throscidae Laporte, 1840

71

Fig. 4.6.2. A, Aulonothroscus laticollis (Rybinski), adult, dorsal (from Burakowski 1991; © Polskie Towarzystwo Entomologiczne, Wroclaw), length = 3.5 mm; B, Trixagus carinifrons (Bonvouloir), adult, dorsal, length = 3.2 mm; C-D, Trixagusdermestoides (Linnaeaus); C, pupa, dorsal, length = 4.5 mm; D, larva, dorsal (from Burakowski 1975; © Polska Akademia Nauk, Warsaw), length = 6 mm.

and prostheca usually absent (reduced in Potergus). Maxilla with galea and lacinia short, broad, lightly sclerotized and pubescent; lacinia without uncus; apical palpomere usually expanded and truncate (fusiform in Potergus). Labium with m e n t u m subtrapezoidal; ligula short, broad, lightly sclerotized, cleft; apical labial palpomere fusiform or apically widened. Subgenal ridges present. Gular sutures widely separated; gula strongly transverse. Corpotentorium narrow. Cervical sclerites well-developed. P r o n o t u m about 0.5-1.0 times as long as wide, not narrower than elytral bases; usually widest posteriorly; with sides slightly to strongly curved and converging anteriorly, sometimes sinuate (subparallel in Pactopus); lateral carinae usually incomplete, not extending to anterior edge (complete b u t not visible f r o m above in Potergus), simple, with or without raised margin; anterior angles not produced forward; posterior angles moderately to strongly acute and produced posteriorly, fitting into impressions beneath elytral humeri; posterior edge biemarginate, fitting tightly against elytra; disc simple. Promesothoracic interlocking mechanism well developed. Prosternum in f r o n t of coxae longer t h a n mid length of procoxal cavity; slightly to strongly convex, with paired longitudinal carinae (except in Potergus); anterior edge more or less produced to form truncate or rounded chin piece. Prosternal process complete, parallel-sided or gradually expanded and then narrowed, flat or slightly elevated, b u t not curved behind coxae, extending well beyond coxae and almost to metaventrite; apex narrowly to broadly rounded. Notosternal suture complete, widely open anteriorly forming antennal grooves, which curve laterally onto hypomeron and extend almost to lateral edges.

Hypomera behind antennal grooves excavated to receive profemora. Procoxae not projecting, w i t h o u t concealed lateral extensions; trochantinopleuron reduced, concealed and fused to wall of hypomeron. Procoxal cavities more or less circular, moderately widely separated, externally broadly open, w i t h o u t narrow lateral extensions; internally open. Pro-mesothoracic clicking mechanism present. Scutellar shield well developed, abruptly elevated, anteriorly simple, laterally rounded, posteriorly subacute to broadly rounded. Elytra usually about 1.5-2.2 times as long as combined width and 2 - 4 times as long as p r o n o t u m (2.4-2.5 times as long as wide and 2 . 5 - 3 times as long as pronot u m in Potergus); anteriorly produced and carinate, fitting into transverse groove at base of p r o n o t u m ; punctation distinctly seriate with nine distinct puncture rows and no scutellary striole; interstices usually much wider than striae, flat, with microsculpture and fine punctation, sometimes doubled; apices conjointly rounded or angulate; epipleura narrow and incomplete; elytral vestiture sometimes sexually dimorphic; males with dense fringe of hairs (Trixagus exul Bonvouloir) or sparse group of long hairs (T. carinifrons Bonvouloir) along outer margin of each elytron. Mesoventrite separated by complete sutures f r o m mesanepisterna, which are well separated f r o m one another; anterior edge on different plane than metaventrite, at middle with raised area, which may be notched, flanked by large paired procoxal rests, and continued posteriorly as a diagonal slide leading to large, deep mesoventral cavity; discrimen absent. Mesocoxae not projecting, with concealed trochantins. Mesocoxal cavities widely separated, open laterally (partly closed by mesepimeron), with solid j o i n t between

72 meso- and metathoracic portions of coxal cavities. Mesometaventral junction a straight line or absent due to fusion. Metaventrite moderately to strongly convex; discrimen usually absent (moderately long b u t weakly indicated in Pactopus and Potergus); transverse (katepisternal) suture absent; deep, curved mesotarsal grooves extending f r o m mesocoxal cavities to posterolateral edges of metaventrite present in most genera (absent in Trixagus, which has pair of curved postcoxal lines defining leg impressions); exposed portion of metanepis t e r n u m narrowly elongate and parallel-sided; anterior part of metaventrite and metanepisterna slightly impressed, forming with mesepimeron a rest for mesofemur and mesotibia. Metacoxae contiguous, extending laterally to meet elytra, plates well developed and more or less complete to lateral edge. Metendosternite with moderately to very long lateral arms, no laminae, moderately long anterior process and approximate anterior tendons. H i n d wing relatively short and broad, with well developed apical field; radial bar heavily pigmented; radial cell vaguely indicated, w i t h o u t basal edge; cross-vein r3 absent, r4 incomplete or absent; basal portion of RP usually absent (long in Pactopus and Potergus); R-M loop either narrow and acute or absent; medial spur slightly to strongly curved, ending well before margin, where there is usually a distinct medial embayment; medial field usually with three free veins (five in pactopus); wedge cell absent; anal lobe well developed, anal embayment deep and notch-like. Legs adapted for h i g h degree of compaction, with f e m u r and tibia flattened, ventrally convex with normal punctation and vestiture and dorsally excavate, impunctate and shiny; fore legs somewhat shorter t h a n other pairs; trochanterofemoral joint on fore legs almost transverse, those on mid and h i n d legs more strongly oblique; metatrochanters larger, shorter and broader than meso- or metatrochanters; tibiae relatively slender, flattened, usually widest at middle, sinuate before apex, outer apical angle of protibia toothed in Potergus; tibial spurs absent; tarsi 5-5-5, tarsomeres relatively slender, 3 and 4 with slight ventral lobes, except in Potergus; pretarsal claws simple; empod i u m absent. Abdomen with five connate ventrites. Ventrite 1 not longer than 2, much shorter at midline; usually with paired transverse impressions for housing legs; with short, oblique, postcoxal lines in Pactopus and Potergus; intercoxal process narrow and acute. Ventrites 2 and 3 on each side with either vague, lateral impressions (Aulonothroscus, Trixagus) or paired, oblique, deep grooves (Potergus and Pactopus) for reception of metatarsi. Ventrite 5 at least slightly longer than preceding ones (much longer in Pactopus and Potergus), its apical edge sometimes finely crenulate. Abdominal tergites usually membranous (lightly sclerotized in Potergus). Spiracles located in pleural membrane, those on segment VIII non-functional or absent in Aulonothroscus and Trixagus. Anterior edge of

Jyrki Muona,John F.Lawrence and A d a m Slipinski

sternite VIII in male usually without or with very short median strut (with long strut in Potergus). Tergite IX in male usually emarginate and X well developed and free, sometimes with IX and X fused; sternite IX more or less rounded at base, occasionally angulate, without spiculum gastrale. Aedeagus of trilobate type, symmetrical; phallobase elongate, slightly flattened, with deep emargination; 2 - 5 times as long as parameres, which are individually articulated, lancet-shaped, not laterally curved and without teeth. Penis long and slender, flattened, with long anterior struts. Sternite VIII in female with long, basally fixed, spiculum ventrale. Ovipositor narrowly elongate, lightly sclerotized; paraprocts with baculi, much longer than coxites, each of which has a transverse baculum and terminal apical stylus. Internal female tract with well developed bursa bearing paired sclerites; spermatheca pigmented, attached directly to tract posterad of bursa, with three lobes. [Cobos 1961,1967, 1982; Burakowski 1975,1991; Coffin 1993; Muona 1993,1995; Leseigneur 1995; 1998; Lawrence etal. 1999 b; Johnson 2002.]

Morphology, Larvae (Figs. 4.6.2 D, 3). Moderately elongate, widest at middle, slightly flattened, dorsally curved, evenly lightly pigmented and soft, white to pale yellow, except for portions of head, mandibles and prothoracic rods, which are heavily sclerotized and dark in color. Surfaces smooth and vestiture of sparsely distributed, fine hairs. Head p r o g n a t h o u s or slightly elevated; protracted; not concealed f r o m above by prothorax; clearly narrower t h a n thorax, anteriorly depressed and wedge-like. Posterior edge of head capsule not emarginate. Epicranial suture and median endocarina absent. Median endocarina, dorsal

y) »

4

Fig. 4.6.3. Trixagus dermestoides (Linnaeus), larval head: A, dorsal; B, ventral (from Burakowski 1975; © Polska Akademia Nauk, Warsaw).

Throscidae Laporte, 1840

endocarinae and stemmata absent. Frontoclypeal suture absent; labrum completely fused to head capsule; clypeolabrum anteriorly produced and subtriangular. Antennae very small, less than 0.15 times as long as head width, located on raised articulating membrane and thus well removed from mandibular articulations, 2-segmented; antennomere 1 short and broad, 2 more than twice as long and much narrower; sensorium on preapical antennal segment as long as or longer than antennomere 2, palpiform. Mandibles symmetrical; non-opposable, flattened and more or less triangular; without groove or perforation, more or less straight and apically rounded; incisor edge single; mola and prostheca absent; base of mandible fused to head frame. Ventral mouthparts strongly retracted; cardines not separated from stipites; maxillary bases widely separated; stipes longer than wide; maxillary articulating area absent. Maxilla with galea and lacinia combined to form single subacute lobe but separated by longitudinal suture, surface more or less glabrous; palp 3-segmented. Labium consisting of prementum and postmentum, the latter elongate, about twice as long as prementum and separated by complete sutures from stipites; prementum slightly elongate, ligula slender and about as long as labial palps, which are 1-segmented with an apical sensilla and separated by less than a basal width. Ventral portion of head capsule supported by a heavily sclerotized frame consisting of a pair of thick bars extending from mandibular insertions to base of maxillolabial complex, then forking to form lateral and mesal branch, both of which meet a posterior bar extending along base of head. Hypopharyngeal rods, ventral epicranial ridges and gula absent. Prothorax about equal in length to meso- and metathorax; thoracic terga simple, without sclerotized plates, asperities or carinae. Proventer with a series of sclerotized supporting rods: one oblique or transvere pair on the presternal area, a longitudinal one joining the presternum to the coxa on each side and then forking and delimiting the lateral and mesal edges of the coxal lobe, the mesal fork extending posteriorly to the end of the segment and sometimes forming distinct spur just mesad of coxa; fore legs slightly larger than posterior pairs, with larve, oval coxae separated by about two coxal diameters; meso and metacoxae smaller and separated by almost six coxal diameters; all legs very short, about 0.15 times width of mesothorax, 5-segmented, with claw-like pretarsus bearing 2 setae. Abdomen about 2.5 times as long as thorax; terga simple, without asperites, carinae or lateral processes; segment IX shorter than VIII, tergum IX not or only slightly extending onto ventral surface, usually as lightly pigmented as preceding terga, occasionally with small pigmented area near posterior end; urogomphi very small and tooth-like or absent; sternum IX simple; X distinct; anal region posteroventrally oriented. Spiracles biforous, with closing apparatus, those on thorax slightly larger, those

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on segment VII functional and about the same size as others on abdomen [Boving & Craighead 1931; Burakowski 1975, 1991; Becker 1991; Lawrence etal. 1999 a.] Morphology, Pupa (Fig. 4.6.2 C). [Burakowski 1975.] Phylogeny and Classification. Throscidae traditionally included, not only those taxa treated above, but also genera such as Drapetes Dejean and Lissomus Dalman, which are currently placed in the elaterid subfamily Lissominae (Laporte 1840; Lacordaire 1857; Schenkling 1928; Cobos 1967). Crowson (1955) expanded the limites of the group to include Balgus Fleutiaux and related genera previously placed in Eucnemidae but now forming the elaterid subfamily Thylacosterninae (see 2-4.6). Throscidae is used here in the sense of Burakowsky (1975) and Lawrence (1988), including the four extant genera discussed above plus a fifth genus, Cryptophthalma Cobos (1982), which appears to be based on an autapomorphic Aulonothroscus species. An additional extinct genus Jaira Muona (1993) was described from Baltic amber. Neocrowsonia Kistner & Abdel-Galil (1986), originally placed in Throscidae, does not belong here, and is treated in Chapt. 2 - 4 . 1 8 . In this restricted sense, Throscidae appears to form a clearly monophyletic group, but its relationships to other elateroid families has been disputed. In cladograms based on a range of elateriform taxa utilizing adult and larval characters (Lawrence 1988; Lawrence etal. 1995) or larval characters alone (Beutel 1995), Throscidae (sensu stricto) almost always formed a monophyletic group with Cerophytidae and Eucnemidae. Calder et al. (1993) analyzed a set of 23 elateroid taxa utilizing adult and larval characters and recovered a siimilar clade. In Muona's analyses involving a different set of elateroid taxa, including more throscids and eucnemids, and utilizing mainly adult characters, the four extant throscid genera plus the extinct Jaira formed a monophyletic group with Lissominae, which was sister to Thylacosterninae and well within Elateridae. In cladograms produced by Lawrence et al. (2007) utilizing morphological and COl sequence data, Cerophytum Latreille and Aulonothroscus formed a monophyletic group sister to Brachypsectra LeConte and outside Elateridae, while Thylacosterninae and Lissominae formed two clades within Elateridae. In a more recent molecular phylogenetic study (Bocakova etal. 2007) involving a wide range of Elateriformia, the genus Trixagus formed 1) a monophyletic group with Telegeusis Horn (Telegeusidae) and Drilonius Kiesenwetter (Omethidae) outside Elateroidea (sensu lato), 2) a clade at the base of Elateroidea, excluding Drilonius and Telegeusis, 3) a clade at the base of the family Lycidae or 4) a monophyletic group with Omalisus Geoffroy (Omalisidae) at the base of Eucnemidae, depending on the type of alignment and analysis.

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Acknowledgements CSIRO Australia is acknowledged for supporting the research of J F L and SAS, and CSIRO Australia, Polskie Towarzystwo Entomologiczne, Wroclaw and Polska Akademia Nauk, Warsaw are acknowledged for allowing us to use copyrighted figures. Anne Hastings is gratefully acknowledged for preparation of the plates.

Literature Becker, E. C. (1991): Throscidae (Elateroidea) (= Trixagidae). Pp. 4 1 8 - 4 1 9 in Stehr, F. W. (ed.), Immature Insects. Vol. 2. Kendall/Hunt Publishing Co., Dubuque, Iowa. Beutel, R. G. (1995): Phylogenetic analysis of Elateriformia (Coleoptera: Polyphaga) based on larval characters. - Journal of Zoological Systematics and Evolutionary Research 3 3 : 1 4 5 - 1 7 1 . Bocakova, M., Bocak, L., Hunt, T., Teraväinen, Μ. & Vogler, A. P. (2007): Molecular phylogenetics of Elateriformia (Coleoptera): evolution of bioluminescence and neoteny. - Cladistics 2 3 : 4 7 7 - 4 9 6 . Boving, A. G. & Craighead, F. C. (1931): An illustrated synopsis of the principal larval forms of the order Coleoptera. - Entomologica Americana (N.S.) 11(1930): 1 - 3 5 1 . Burakowski, B. (1975): Development, distribution and habits of Trixagus dermestoides (L.), with notes on the ThroscidaeandLissomidae(Coleoptera,Elateroidea). -Annates Zoologici(Warszawa) 3 2 : 3 7 5 - 4 0 5 . . - (1991): Kluczedo Oznaczania OwadowPolski. CzescXIX. Chrzaszcze - Coleoptera. Zeszyt 35-37. Cerophytidae, Eucnemidae, Throscidae, Lissomidae. 91 pp. Polskie Towarzystwo Entomologiczne, Warsaw, 91 pp. Calder, Α. Α., Lawrence, J. F. & Trueman, J. W. H. (1993):Austrelater, gen. nov. Coleoptera: Elateridae), with a description of the larva and comments on elaterid relationships. - Invertebrate Taxonomy 7: 1349-1394. Cobos, A. (1960): Throscidae de Africa Central (Coloeoptera). Annales du Musee Royal du Congo Beige Tervuren (Belgique). Serie in 8°. Scineces Zoologiques. Volumne 8 6 , 6 7 pp. - (1961): Sobre la posicion sistematica del genero Potergus Bonvouloir y revision de las categorias supragenericas de la familia Throscidae (Coleoptera). - Bulletin de l'Institut Royal des Sciences Naturelles de Belgique 37(35): 1 - 6 . - (1967): Estudios sobre Throscidae, II. - Eos 4 2 (1966): 3 1 1 - 3 5 1 . - (1982): Un notable nuevo genero y especie de Throscini Neotropical (Coleoptera, Throscidae). - Revue Frangaise d'Entomologie (N.S.) 4:54—56. Coffin, J. (1993): "Fosses cephaliques" enigmatiques decouvertes chez les especes du genre Throscus Latreille, 1796 (Coleoptera, Throscidae). -Bulletin de la SocieteEntomologique de France 97(1992): 3 0 9 - 3 1 1 . Crowson, R. A. (1955): The Natural Classification of the Families of Coleoptera. 187 pp. Nathaniel Lloyd, London. - (1960): The Phylogeny of Coleoptera. - Annual Review of Entomology 5:111-134.

Jyrki Muona, John F.Lawrence and Adam Slipinski Johnson, P. J. (2002): 57. Throscidae Laporte 1840. Pp 1 5 8 - 1 5 9 in Arnett, R. H., Jr., Thomas, M. C., Skelley, P. E. & Frank, J. H. (eds.) American Beetles. Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Gainesville, Florida. Kistner, D. H. & Abdel-Galil, F. A. (1986): A new genus and species of termitophilous Throscidae from South Africa (Coleoptera). - Soctobiology 12 (2): 305-314. Lacordaire, T. (1857): HistoireNaturelle deslnsectes. Genera des CoMopteres. Tome4.579 pp., pi. 4 0 - 4 7 . Libraire Encyclopedique de Roret, Paris. Laporte, F. L. (1840): HistoireNaturelle des AnimauxArticules, Annelides, Crustaces, Arachnides, Myriapodes et Insectes. Vol. 2. Histoire Naturelle deslnsectes Coleopteres. Vol. 1.324 pp., 19 pis. P. Dumenil, Paris. Lawrence, J. F. (1988): Rhinorhipidae, a new beetle family from Australia, with comments on the phylogeny of Elateriformia. - Invertebrate Taxonomy 2(1987): 1 - 5 3 . Lawrence, J. F. & Newton, A. F. Jr. (1995): Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names). Pp. 7 7 9 - 1 0 0 6 in Pakaluk, J. & Slipirtski, S. A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: papers Celebrating the 80th Birthday of Roy A. Crowson. XII + 1092 pp. Muzeum i Instytut Zoologii Polska Akademia Nauk, Warsaw. Lawrence, J. F., Hastings, A. M., Dallwitz, Μ. J., Paine, T. A. & Zürcher, Ε. J. (1999 a): Beetle Larvae of the World: Descriptions, Illustrations, Identification, and Information Retrieval for Families and Subfamilies. CDROM, Version 1.1 for MS-Windows. CSIRO Publishing, Melbourne. - (1999 b): Beetles of the World: A Key and Information System for Families and Subfamilies. CD-ROM, Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne. Lawrence, J. F., Muona, J., Teraväinen, Μ., Stahls, G. & Vahtera, V. (2007): Anischia, Perothops and the phylogeny of Elateroidea (Coleoptera: Elateriformia). -Insect Systematics and Evolution 3 8 : 2 0 5 - 2 3 9 . Leseigneur, L. (1995): Statut actuel des genres Trixagus Kugelann, 1794, et Throscus Latreille, 1796. Designation des lectotypes des especes palearctiques de Η. Bonvouloir (Coleoptera, Throscidae). - Bulletin de la SocieteEntomologique de France 1 0 0 : 3 4 7 - 3 5 9 . - (1998): 37. Familie: Throscidae, pp. 2 2 2 - 2 3 1 in W. Lucht & Κ. Β. Klausnitzer (eds.), Die Käfer Mitteleuropas. Band 15.4. Supplement band. Goecke & Evers, Krefeld. Muona, J. (1993): Eucnemidae and Throscidae from Baltic amber (Coleoptera). - Entomologische Blätter 89:15-45. - (1995): The phylogeny of Elateroidea (Coleoptera), or which tree is best today? - Cladistics 1 1 : 3 1 7 - 3 4 1 . Schenkling, S. (1928): Coleopterorum Catalogus. Pars 101. Throscidae, Cerophytidae, Perothopidae. 30 pp. W. Junk, Berlin. Yensen, E. (1975 a): A revision of the North American species of Trixagus Kugelann (Coleoptera: Throscidae). - Transactions of the American Entomological Society 1 0 1 : 1 2 5 - 1 6 6 . - (1975 b): A review of the genus pactopus LeConte (Coleoptera: Throscidae). - The Coleopterists Bulletin 29: 8 7 - 9 1 .

Elateridae Leach, 1815

4.7. Elateridae Leach, 1815 Cleide Costa, John F. Lawrence and Simone Policena Rosa

Distribution. This family, composed of more than 400 genera and almost 10,000 species, is an a b u n d a n t and diverse group in all parts of the world. The four largest subfamilies, Agrypninae, Cardiophorinae, Denticollinae and Elaterinae, are widely distributed. Within the first group, however, the tribe Pyrophorini occurs mainly in the Neotropical region with the exception of two genera f r o m Oceania, while the Agrypnini is represented in all biogeographical regions, the largest n u m b e r of genera and species being Afrotropical and Oriental. The smaller subfamily Negastriinae is also widely distributed, b u t more a b u n d a n t and diverse in the Northern Hemisphere. Pantropical groups include Thylacosterninae, Physodactylinae and Oxynopterinae. Several subfamilies, such as Eudicronychinae, Hemiopinae, Morostominae, Subprotelaterinae occur only in the Eastern Hemisphere, while Campyloxeninae and Semiotinae include only Neotropical genera. Within the subfamily Lissominae, Lissomini are Pantropical, Oestodini (Oestodes LeConte) occurs in northern North America, and Protelaterini occurs in New Zealand and Chile. Cebrioninae are fairly widely distributed, b u t occur primarily in the drier parts of North America, Central and South America, Southern Europe, North Africa, South Africa, Asia Minor and Central Asia. The subfamily Pityobiinae includes one North American genus, one f r o m Chile, one f r o m New Zealand, and several Australian genera. [Schwarz 1906 a, b, 1907 a, b; Hayek 1973; Lawrence 1982; Costa 1975; Costa etal. 1988,1994; Calder 1992,1996,1998; Calder etal. 1993.] Biology a n d Ecology. The external appearance of the elaterids is extremely homogeneous and they are usually recognized by their ability to j u m p into the air while m a k i n g a clicking noise. For that they are k n o w n as "click beetles, skip-jacks, spring beetles, clickers or blacksmiths". The mechanics of this clicking maneuver have been studied by Evans (1972,1973) who stated that the j u m p is normally made f r o m an inverted position and involves a sudden movement of the prosternal process into the mesoventral cavity. The energy needed for the movement is provided by a pair of large prothoracic muscles and muscle tension is built u p by the prosternal process being blocked f r o m moving by a peg located at the opening of the cavity. When sufficient force is applied the prosternal process slides past the peg and into the cavity, and this very rapid movement raises the center of gravity of the beetle fast enough to throw it into the air. Although this maneuver may be used to "right" a beetle which lands on its back, it can be executed from any position and is basically an escape reaction. Generally

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the first reaction of adults when disturbed is to simulate death. Luminous species when disturbed for a long time begin to emit light and are very active. The adults are frequently collected in the vegetation or at lights at night. They are phytophagous consuming plant juices, and in the laboratory many species can be kept alive on a sugar solution. Knowledge of the complete life cycle and i m m a t u r e stages is still quite incomplete, in spite of abund a n t literature on those of economic importance. The life cycle usually takes one to two years, b u t if conditions are unfavorable the development may lengthen f r o m five to eight years. Larvae of Fulgeochlizus bruchi (Candeze) (Pyrophorini) took six years f r o m egg to adult. Stone (1950) reported a longevity for over twelve years of a single larva of Melanactes densus LeConte (Denticollinae). Pupation takes place in an oval chamber that the larva prepares at the end of a gallery in rotten wood, or in the soil. Larvae of Tetralobini (Pseudotetralobus Schwarz f r o m Australia and Tetralobus Lepeletier & Audinet-Serville f r o m Africa) construct an oval cocoon of a thin and brittle, blackish brown secretion. The pupal period usually lasts f r o m two to three weeks and that of the adults one to two months. In temperate geographic regions adults overwinter inside the pupal chamber going, emerging in the spring (Costa etal. 1992). A bibliography containing thousands of international publications on the larvae of Elateridae u p to 1977 was published by Gaedike (1969,1975,1979). Elaterid larvae are quite active and can be found in very distinct biotopes, such as in soil, litter, termite nests or rotten wood. They may be saprophages, phytophages, or predators. Larvae of some species that live in soil (Elaterinae) are commonly k n o w n as "wireworms". These larvae are elongate, more or less cylindrical with a highly chitinized cuticle, and are usually yellow or reddish-brown. Gur'yeva (1969) considered the functional and adaptative significance of the three basic larval types described by Hyslop (1917), correlating t h e m with environmental factors. Larvae of Type I (Hyslop's Pyrophorinae, including Agrypninae, Denticollinae, Pityobiinae, Negastriinae and several other groups), which have a variably sclerotized integument, dorsoventrally flattened body, apically forked tergum IX (paired urogomphi) and falcate mandibles (Fig. 4.7.8 A, B, D), are active burrowers either in heavy soils, or beneath bark and in rotten wood. They are able to move effectively within the tunnels they construct partly with the assistance of attachments devices (spinose legs and urogomphi, anal hooks) (Fig. 4.7.8 A-F). Larvae of Type II (Elaterinae) have a cylindrical or semi-cylindrical body with heavily, more or less uniformly sclerotized integument, a simple tergum IX (without urogomphi) and falcate mandibles (Figs. 4.7.8 H, 4.7.12 C). This type of larva m u s t use ready-made passages and spaces because of their cylindrical bodies and lack of attachment devices. They occur in light soils b u t may also be f o u n d in beneath bark or in rotting timber where

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Cleide Costa,John F.Lawrence and Simone Policena Rosa

Fig. 4.7.1. A-I, Adult habitus (dorsal). Agrypninae: A, Agrypnus molardi Girard; B, A. royae (Girard); C, Dilobitarsus abbreviatus Candeze; D, Elasmosomus testaceus Girard; E, Macromalocera sinuaticollis Blackburn; F, Thoramus laevithorax (White); G, Prodrasterius nimbanus Girard; H, Heteroderesfuscus Latreille; I, H. rufangulus (Gyllenhal). A, B, D, G, after Girard (2003); C, I, after Costa etal. (1988); E, after Calder (1996); F, after Costa (1992); H, after Girard (1971).

Elateridae Leach, 1815

fibers are more or less separated as a result of decomposition. Larvae of Type III (Cardiophorinae) have a soft integument, cylindrical body subdivided into pseudosegments, no u r o g o m p h i and bilobate mandibles (Figs. 4.7.8 G, 4.7.12. Α-B). These larva often occur in light soils and are active burrowers, using the unusual mandibles to break u p soil particles. Several species of wireworms, mainly in the genera Agriotes Eschscholtz, Athous Eschscholtz, Cardiophorus Eschscholtz, Ctenicera Latreille, Conoderus Eschscholtz and Melanotus Eschscholtz, are considered as important pests attacking cereal and forage crops. All larvae are liquid feeders, capable of extra oral digestion. The ventral m o u t h p a r t s are consolidated to form a maxillolabial complex, which is densely clothed above (internally) with short hairs forming a kind of oral filter that keep solid particles out of the gut. It was observed in the Ctenicera aeripennis (Kirby) group (Denticollinae) that the larvae regurgitated a brown liquid, probably ventricular in origin, which contained amylase. To feed the larvae masticate the potato tissue, regurgitate fluid, and then imbibe the plant juices that are liquefied by enzymatic action. The regurgitate of Pyrearinus termitilluminans Costa (Agrypninae: Pyrophorini) contains trypsin and a mixture of carbohydrases (amylase, cellulase, trehalase, and a - and ß-glucosidase), which accomplishes initial, extra-corporeal digestion of the prey. The larvae then ingest preliquefied material, and the intermediate and final digestion take place on the surface of the m i d g u t cells, carried out by aminopeptidase, α-glucosidase, and trehalase. The p H optima of the digestive enzymes range f r o m 6 and 8.5, which agrees with the p H value of regurgitate and m i d g u t contents. Larvae of Dilobitarsus abbreviates Candeze (Agrypninae), which live inside arboreal termite nests ( 6 - 1 2 m of height), and those of Pseudotetralobus spp. (Australia) and Tetralobus spp. (Africa), which live in soil in termite nests or in wood infested by termites, are physiogastric resembling termite queens (Fig. 4.7.8 L). However, larvae of Anchastus brunneofasciatus Schwarz (Elaterinae), which inhabit epigeous termite nests in marshy areas, and those of Oxynopterus mucronatus Olivier (Oxynopterinae), which are predators in the nests of Neotermes Holmgren in Java (Ralshoven 1955), are not physiogastric (Fig. 4.7.8 H). The larva of Tetralobus subsulcatus Guerin-Meneville possesses glands in the dorsal area of abdominal segment VIII the function of which is u n k n o w n . Zacharuk (1962) studied the sensory mechanisms of elaterid larvae and described seven types of sensilla on the larval head: i) thick-walled hair organs; ii) campaniform organs; iii) mandibular pore canal organs; iv) scolopophorous organs; v) peg or thin-walled hair organs; vi) plate organs and vii) antennal sensory appendix. Mandibular pore canal sensilla were studied in more detail by Zacharuk & Albert (1978) who considered that the scolopophorous sensillum has a proprioceptive function, monitoring stresses and deformations in the biting and chewing of

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mandibles. Zacharuk etal. (1977) discovered on the distal maxillary and labial palpomeres of larvae of Ctenicera destructor (Brown) elongate pegs they termed digitiform sensilla. The location of these pegs would provide contact between t h e m and the walls of the larval tunnels in the soil. Since movem e n t of prey or predator in the substrate nearby could generate a vibratory stimulus, the location of the pegs is compatible with directional monitoring of such stimuli for tunneling activity, prey capture or predator evasion. Approximately 200 species of the agrypnine tribe Pyrophorini f r o m Neotropical region and Oceania, and two other Neotropical species in Thylacosterninae (Balgus schnusei Heller) and Campyloxeninae {Campy loxenus pyrothorax Fairmaire) are bioluminescent (Costa 1984 a). Adults of B. schnusei emit a green light f r o m two swellings on the prothorax. Its larvae are u n k n o w n . Adults of Pyrophorini possess a pair of bioluminescent oval vesicles (yellowish when the light is extinct or the insect is dead), located at the base of the posterior angles of the prothorax. There is a third luminescent organ located on the ventral surface of abdominal segment I that is only activated d u r i n g the flight. The Oceanic genus Hifo Candeze has only the abdominal l u m i n o u s organ (Costa 1984 b). The color of the emitted light f r o m the prothoracic vesicles is green and that f r o m the abdomen is orange, yellow or red, b u t b o t h emit continuous light. During flight, the abdominal organ illuminates a circular field below the body, at the same time the thoracic ones remain illuminated. When the beetles are not flying, the thoracic organs may light up. However, the abdominal organ functions only when there is some movement of the wings. Adult bioluminescence may be associated with sexual attraction, b u t the communication system has not been studied. Colepicolo-Neto etal. (1986 b) reported bioluminescence f r o m eggs of Pyrearinus termitilluminans with dark-adapted eyes, and also recorded their spectra. They glow dimly and continuously. Pyrophorine larvae only light u p in the green area of the spectrum. Their luminescence is f o u n d mainly on the p r o n o t u m , and in some species, there are paired, lateral r o u n d organs or transverse dorsoventral zones on each abdominal segment. In the larval phase, bioluminescence may be associated also with defense, b u t detailed studies have not been made. Very impressive are the larvae of P. termitilluminans f r o m Central Brazil, which are found in old nests of Cornitermes cumulans (Kollar) (Isoptera, Rhinotermitidae) a meter or more in height (Costa 1982; Redford 1982). These larvae excavate an intricate network of tunnels in the outer layers of the m o u n d s , with exits to the outside, where they expose their head and greenshining prothorax to attract and catch flying prey, especially termites and ants. Polymer molds of the larval galleries revealed the existence of an intricate superficial mesh of tunnels 0.1 to 0.3cm in diameter, occupying almost the whole surface of the nest, and individual " U " shaped galleries and

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also a small atrium that seems to constitute a place for stockpiling predigested prey. These larvae also illuminate the last abdominal segment intensely. Pre-pupae and pupae generally show luminescence over the entire body, with variations of intensity depending on the species. After 10-14 days of the pupal phase it is easy to perceive the outlines of the adult luminous organs. A newly emerged adult displays pupal luminescence (over the entire body), but as it becomes more sclerotized only the prothorax and abdominal organs emit light (Costa 1970; Casari-Chen & Costa 1986).

nowadays-accepted notion that a higher SOD activity in the tissues of aerobic organisms provides a better anti-oxidant protection against reactive oxygen species. Using in vivo EPR oxymetry with lithium phthalocyanin crystal probes implanted in the prothorax of P. termitilluminans larvae, were found recently that about 20% of the local oxygen (p0 2 ) is promptly consumed, possibly by the light reaction, when the animal is provoked with a termite prey to light up (Timmins etal. 1999). In the other hand, a major difficulty in explaining the evolution of bioluminescence, as a signaling mechanism is to understand how progressive evolution acted upon primal and low-efficient bioluminescent reactions, to select for and optimize such a weak exergonic light emission reaction to a threshold sufficient to be perceived by light-sensing organs. [Dubois 1886; Bianchi 1937; Stone 1950; Costa 1970,1975,1982; Crowson 1981; Sivinski 1981; Redford 1982; Colepicolo-Neto etal. 1986 a; Costa etal. 1988.]

As in many other organisms the mechanism of light production involves a luciferin in the presence of oxygen, the enzyme luciferase, and the cellular energy source ATP (adenosine triphosphate). Noteworthy is the fact that, although the luciferin and oxyluciferin (the bioluminescent reaction emitter) are chemically the same in all bioluminescent beetles, the colors elicited in vitro by Phengodidae (green - > red) and Elateridae (green - > orange) luciferases are not affected by pH changes, whereas those of Lampyridae were shown to shift from yellow-green to red upon lowering the pH from 7.4 to 6.0. This has been interpreted as resulting from subtle pH-dependent changes of the insect luciferase structure, at the microenvironment of the active site, leading to protonation changes of critical amino acid residues (e. g., arginine), rendering the electronically excited product, oxyluciferin, mainly in a lactame form (yellow-green emitter) or in an enolate form (red emitter) (Viviani & Bechara 1995; Viviani et al. 1999). The Jamaican Pyrophorus plagiophthalmus Germar displays a striking light color polymorphism. They vary in the color of their ventral light organs from yellow-green to orange and their dorsal organs from green to yellow-green. It was discovered that the genetic basis for the color variation involves specific amino acid substitutions in the luciferase (Stolz etal. 2003). Several investigators have examined the hypothesis that beetle bioluminescence may have evolved primarily to provide an oxygen detoxifying mechanism auxiliary to those driven by antioxidant enzymes such as superoxide dismutase (SOD) and catalase (Colepicolo-Neto et al. 1986 a; Barros & Bechara 1998; Timmins et al. 1999). Larvae of P. termitilluminans live under conditions close to normoxia into tunnels dug into termite mounds, whereas other elaterid larvae inhabit tunnels into decaying logs, where the local oxygene (p0 2 ) is ~ 2 5%. Not surprisingly, P. termitilluminans larvae have SOD activity 3- and 15-fold, respectively, higher than those found in other luminescent (e. g., Pyrophorus divergens Eschscholtz) and non-luminescent larvae as Chalcolepidius sp., Platycrepidius sp., Conoderus sp. (Agrypninae) and Ischiodontus sp. (Elaterinae) (Colepicolo-Neto etal. 1986 a). Furthermore, the SOD activity is two-fold higher in the brightest segments (prothorax and abdominal segments IX-X) of P. termitilluminans larva than in its dim segments (abdominal segments I-VIII). This is in accordance with the

Morphology, Adult (Figs. 4.7.1 A-I; 4.7.2 A-I; 4.7.3 A-I; 4.7.4 A-I). Length 0.9-75 mm. Body moderately to very elongate, 1.7-5.2 times as long as wide; usually more or less parallel-sided, with elytra slightly expanded basally and tapered apically; prothorax often distinctly curved laterally and narrowest point at pronoto-elytral junction. Somewhat flattened to moderately convex dorsally and ventrally, glabrous or pubescent. Sexual dimorphism usually absent, but some females larger than males, more convex, and with more strongly curved sides, sometimes with differing punctation. Head prognathous to moderately strongly declined, subquadrate to slightly transverse, not abruptly constricted posteriorly. Transverse occipital ridge and median occipital endocarina almost always absent. Frontal region highly variable, not or only slightly declined to vertical or inflexed at apex, forming frontal carina. Eyes entire, often large and protuberant, often smaller in females; finely facetted, without interfacetal setae; ommatidium of exocone type (Caveney 1986). Antennal insertions (Fig. 4.7.5 E, F) moderately to widely separated, exposed or concealed from above, sometimes elevated or placed within large, saucer-like impressions. Subantennal groove present or absent. Frontoclypeal suture absent. Labrum (Fig. 4.7.5 E, F) free, slightly to strongly transverse, moderately or heavily sclerotized. Antennae (Fig. 4.7.6 B) usually 11-segmented (12-segmented in Diplophoenicus Candeze (Morostominae), Euthysanius LeConte (Cebrioninae), Pityobius LeConte (Pityobiinae), Tetralobini (Agrypninae), Warduluipicola Calder (Denticollinae), Odontonychini (Elaterinae), and Eudicronychinae), usually filiform or serrate, sometimes pectinate, bipectinate or flabellate, with rami beginning on antennomere 3 or 4; scape usually much longer than pedicel. Female antennae usually serrate when male antennae are flabellate or pectinate. Mandible (Fig. 4.7.5 G-I) short and broad to long and narrowly falcate, with apex

Elateridae Leach, 1815

79

p i •V, :·.··"

Fig. 4.7.2. A - I , A d u l t h a b i t u s (dorsal). Agrypninae: A, Chalcolepidius zonatus Eschscholtz; B, platycrepidius bicinctus Candeze; C, pyrophorus divergens Eschscholtz. Denticollinae; D, Glypheus villosulus Candeze; E, Rousia dumbrellium Calder; F, Telesus vanderplaetseni Girard; G, Acteniceromorphus nepalensis Öhira & Becker; H , Arachnodima opaca Candeze; I, Gambrinus hiramatsui Öhira. A-C, a f t e r Costa etal (1988); D, Ε, H , after Calder (1996); F, after Girard (2003); G, after Öhira & Becker (1973); I, after Öhira (1976).

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Fig. 4.7.3. A-I, Adult habitus (dorsal). Cebrioninae; A, Cebrio bicolor Fabricius. Cardiophorinae; B, Horistonotus sp. Eudicronychinae; C, Anisomerusprosternalis (Schwarz). Elaterinae; D, propsephus hiekei Girard; E, Anchastus brunneofasciatus Schwarz; F, Dactylosimus dorsalis Fleutiaux. Hemiopinae; G, Hemiopsflava Castelnau. Lissominae; H,Austrelater macphersonensis Calder & Lawrence; I, Lissomus sp. A, after Hyslop (1915); Β, Ε, I, after Costa et al. (1988); C, after Girard (2003); D, after Girard (1986 a); F, after Costa & Casari-Chen (1993); H, after Calder etal. (1993).

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Fig. 4.7.4. A-I, Adult habitus (dorsal). Tetralobini; A, Tetralobus shuckhardi (Hope); B, Tetralobusflabellicornis (Linnaeus). Physodactilinae; C, Physodactylusfleutiauxi Chassain. Subprotelaterinae; D, Subprotelater sp. Semiotinae; E, Semiotus intermedins (Herbst). Pityobiinae; F, Parablaxpadmuri Calder. Thylacosterninae; G, Balgus schnusei (Heller). Oxynopterinae; H, Oxynopterus strachani Hope. Negastriinae; I, Νegastrius pulchellus (Linnaeus). A, after Girard etal. (2007); B, after Costa etal. (1994); C, after Chassain (2005); D, after Calder (1984); E, after Costa etal. (1988); F, after Calder (1996); G, after Chassain (2003); H, after Girard (1971); I, after Stibick (1971).

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strongly curved mesally, bidentate or unidentate; incisor edge usually simple; mola almost always absent (present in some melanotine Elaterinae), mandibular base often with b r u s h of hairs. Maxilla (Fig. 4.7.5 B, C) with galea and lacinia simple and pubescent, occasionally reduced. Labium (Fig. 4.7.5 A, D) partially membranous, without ligula. Apical maxillary and labial palpomeres usually at least slightly expanded apically, sometimes cylindrical or fusiform; last palpomeres greatly elongate in Morostoma palpale Candeze (Morostominae) (Fig. 4.7.5 A, B). Subgenal ridges present or absent. Gular sutures, if present, moderately widely separated. Corpotentorium narrow or absent. Cervical sclerites well-developed. Prothorax (Fig. 4.7.6 A, C) about 0.4-1.35 times as long as wide, usually widest posteriorly; sides usually more or less sinuate, sometimes straight or curved anteriorly, not explanate; lateral carinae usually complete (incomplete or absent in some Cardiophorinae and Cebrioninae and a few Denticollinae); anterior angles rounded, truncate or slightly produced forward; posterior angles acute to narrowly acute and often strongly produced posteriorly or posterolaterally, often carinate, with carinae sometimes extending forward onto disc or rarely forming sublateral carina (e. g., in Tetralobini and Negastriinae); posterior edge usually distinctly sinuate or variously lobed, almost always simple (crenulate in Oistus Candeze and Perissarthron Hyslop, Oxynopterinae); disc usually simple or with broad, shallow sub-basal impressions (with paired carinae in Hemiopinae and some Cardiophorinae). Promesothoracic interlocking mechanism (Fig.

crural impression. Procoxae (Fig. 4.7.6 C) not projecting below prosternum, without or with short concealed lateral extensions; procoxal cavities more or less circular, narrowly to moderately widely separated, externally broadly or narrowly open (postcoxal projection absent or very short to moderately long b u t not meeting prosternal process), almost always without narrow lateral extensions, internally open or closed by slender bar; trochantin almost always reduced and concealed or absent; endopleuron more or less fused to n o t u m at base of notosternal suture. Scutellar shield well developed, abruptly elevated or not, sometimes strongly oblique, almost always anteriorly simple (notched in most Cardiophorinae or crenulate in Oistus), posteriorly acute to broadly rounded or truncate. Elytra usually at least twice as long as combined width and 2 - 6 times as long as pronotum; usually with nine distinct puncture rows or impressed striae and scutellary striole (ten striae in Tetralobini), sometimes irregularly punctate; apices usually conjointly rounded, without internal interlocking tongue, sometimes independently rounded, acute or with apical spines, truncate exposing apical abdominal tergites in females of some Cebrioninae; epipleura complete, incomplete or absent. Mesoventrite (Fig. 4.7.6 A, C) on same plane as or on different plane than metaventrite, separated by complete sutures f r o m mesanepisterna, which are well separated from one another; anteriorly with or without paired, horizontal to strongly oblique or vertical procoxal rests; discrimen absent; mesoventral cavity usually moderately to very large and deep, occasionally shallow, usually with distinct anterior boss on which prosternal process rests prior to "clicking" maneuver. Mesocoxae only rarely projecting. Mesocoxal cavities usually moderately to widely separated, occasionally narrowly separated, rarely contiguous, circular to slightly transverse, usually open laterally (closed by mesepimeron, occasionally with mesanepisternum), closed laterally by ventrites in Cardiophorinae, Negastriinae and some Agrypninae, without joint between meso- and metathoracic portions of coxal cavities. Mesometaventral junction, if present, usually consisting of a straight line, sometimes curved or forming a complex fitting; absent due to fusion of mesoventrite and metaventrite in some genera such as Semiotus Eschscholtz, Oistus (Semiotinae), Campsosternus Latreille (Oxynopterinae) and Chalcolepidius Eschscholtz (Agrypninae). Metaventrite (Fig. 4.7.6 A) usually moderately to very long and slightly to strongly convex; discrimen usually long, occasionally absent; transverse (katepisternal) suture absent; straight and/or strongly recurved postcoxal lines sometimes present in Lissomini; exposed portion of m e t a n e p i s t e r n u m moderately to very elongate. Metacoxae contiguous or narrowly separated, extending laterally to meet elytra; plates present, usually well developed mesally and narrowed laterally; uniformly broad in Thylacosterninae and Lissomini, weakly developed in most Cebrioninae.

4.7.6 C) almost always well developed b u t reduced in some Cebrioninae. Prosternum (Fig. 4.7.6 A, C) in front of coxae almost always much longer than diameter of a coxal cavity, with anterior edge produced forward to form broadly rounded chin piece (shorter than diameter of coxal cavity in some Cebrioninae and with chin piece lacking or truncate in Cebrioninae, as well as in Austrelater Calder & Lawrence [Lissominae] and a few other genera); surface flat to moderately convex, with paired longitudinal carinae in some Drapetes Redtenbacher (Lissominae). Prosternal process complete, parallel-sided or apically narrowed, usually extending well behind procoxae, slightly to strongly curved or abruptly elevated behind them, and fitting into deep cavity in mesoventrite (occasionally shorter or more or less flattened); apex usually acute to narrowly rounded, sometimes broadly rounded or truncate. Notosternal suture usually complete and straight, occasionally curved or sinuate, sometimes anteriorly open (with gap between sternum and hypomeron) or forming deep antennal cavity (forming broad internal antennal cavity above hypomeral wall in Lissomini [Lissominae] and Thylacosterninae). Hypomeron usually simple and flattened; in Subprotelater Fleutiaux (Subprotelaterinae) with deep antennal groove below lateral carina; in some Agrypnini with oblique antennal groove and transverse

Elateridae Leach, 1815

Metacoxae horizontally or slightly obliquely oriented. Metendosternite (Fig. 4.7.6 D) usually with short lateral arms, no laminae, weakly or strongly developed ventrolateral processes, and moderately long anterior process bearing approximate anterior tendons; occasionally with arms longer, anterior process shorter and anterior tendons more widely separated. Hind wing (Fig. 4.7.61, J) almost always well developed; highly reduced or absent in females of a few groups (e. g., some female Cebrioninae and Agrypninae-Pyrophorini); usually 2.5-3.5 times as long as wide (occasionally shorter and broader); radial cell usually well developed and elongate radial cell with inner posterobasal angle right or slightly obtuse and cross-vein r3 moderately long and horizontal; radial cell sometimes shorter and broader or absent (lacking base); apical field usually less than 0.2 times total wing length (approaching or even exceeding 0.3 in at least some Lissominae and Negastriinae), usually with one of three patterns of linear sclerites: a single anterior oblique sclerite, anterior and posterior oblique sclerites meeting or almost meeting basally, or three sclerites forming an epsilon figure (occasionally with no sclerites in apical field); first two apical folds meeting at wing margin; base of RP moderately to very long, sometimes reaching basal fifth of wing; medial field usually with five free veins, no medial binding patch, and a well developed wedge cell with truncate apex from which two subparallel veins (CuA 1+2 and AA3) emerge; free veins reduced to four or three, especially in smaller forms; wedge cell absent in Agrypninae, Subprotelaterinae, Negastriinae, Cardiophorinae and scattered taxa in other subfamilies; anal notch usually absent (present in Negastriinae, a few Cardiophorinae and some Lissominae). Wing reduction may occur in certain Pyrophorini, b u t wing dimorphism is most extreme in some Cebrioninae, where female elytra are greatly shortened and h i n d wings are vestigial. Trochanterofemoral joint transverse to strongly oblique with base of femur separate from coxa; trochanter very long in Lissomini. Metatrochanter and metafemur occasionally greatly enlarged (e. g., Physodactylus Fischer in Physodactylinae, Patriciella Van Zwaluwenburg in Cardiophorinae). Mesotibia usually not strongly widened (widened apically in Physodactylus, Patriciella and some Cebrioninae, subapically in Thylacosterninae), outer edge usually simple, occasionally with teeth or spines, outer apical angle almost always simple or only slightly produced; tibial spurs usually double, occasionally single or absent; tarsi 5-5-5, tarsomeres (Fig. 4.7.6 E - H ) often simple, sometimes with ventral pubescent pads or membranous lamellae on preapical tarsomere (as in Conoderus Eschscholtz) or on more than one tarsomere; pretarsal claws usually simple or toothed in some Lissomini and Protelaterini, cleft in Eudicronychinae, pectinate in Melanotini, Adrastini (Elaterinae), Toxognathus Fairmaire (Physodactylinae) and Aptopus Eschscholtz (Elaterinae), with one or more setae near base in Tetralobini, Morostominae,

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Agrypninae (except Danosoma Thomson and Octocryptus Candeze) and a very few Denticollinae; e m p o d i u m usually well-developed, exposed and bisetose (multisetose in Tetralobini). Abdomen (Fig. 4.7.6 A) almost always with five ventrites (six or seven in some Cebrioninae), the first four of which are usually connate (free in females of some Cebrioninae). Abdominal process almost always acute or narrowly rounded (occasionally broadly rounded or absent). Functional spiracles present on abdominal segment VIII; spiracles on III-VII located in pleural membrane. Anterior edge of sternite VIII in male only rarely with median strut. Anterior edge of sternite IX in male without median strut; tergite IX truncate to deeply emarginate or almost completely divided into two parts. Tergite X usually well developed and free, rarely partly fused to tergite IX or completely membranous. Aedeagus (Fig. 4.7.7 A-F) trilobate, usually symmetrical; parameres almost always individually articulated, with or without apical-lateral hooks. Penis with ventral lobe absent, reduced (rod-like) or developed (Pyrophorini), almost always with paired anterior struts. Sternite VIII in female with long anterior, median strut (spiculum ventrale) and pair of shorter anterolateral struts. Ovipositor (Fig. 4.7.7 G-I) long and slender, lightly to heavily sclerotized, depressed; paraprocts m u c h longer than gonocoxites, which may be divided or undivided; styli usually present and subapical, sometimes absent. Internal female tract (Fig. 4.7.7 G-I) often highly complex; vagina usually somewhat enlarged anteriorly, usually without sclerites, b u t often with paired globular colleterial glands near gonopore; anterior bursa long and tubular to large and sac-like, often with sclerites; one or more spermathecae entering bursa independently of a duct leading to a large, multilobed spermathecal gland. [Lacordaire 1857; Candeze 1857, 1859, 1860, 1863, 1874; Crowson 1955, 1961; Zacharuk 1958; Laurent 1961; Stibick 1971, 1979; Gur'yeva 1974; Costa 1975; Dolin 1975, 1978 b, 2000 a, b; Calder 1992, 1996; Calder et al. 1993; Costa et al. 1994; Lawrence et al. 1999 b.]

Morphology, Larva. Length: 10-60 m m (Fig. 4.7.8 A-M). Body almost always elongate, more or less parallel-sided, flattened to more or less cylindrical (broader and grub-like in Tetralobini and Thylacosterninae). Dorsal and ventral surfaces similarly darkly or lightly pigmented, surfaces lightly pigmented except for head capsule and t e r g u m IX, dorsal surfaces more darkly pigmented than ventral ones, or dorsal surfaces with dark and light pattern. Cuticle generally smooth, b u t sometimes with tubercles, transverse carinae or patches of specialized setae. Vestiture usually consisting of fine hairs or setae only (dense vestiture of long hairs in Tetralobini). Head usually prognathous or slightly declined (strongly declined in Cebrioninae), usually more or less flattened, b u t globular in Cebrioninae;

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Fig. 4.7.5. A—I, Adult structures. Labium and maxillae: A, B, Morostomapalpate Candeze (Morostominae). Maxillae and labium; C, D, Vesperelater occidentalis (Champion) (Agrypninae). Labrum; E, Piezophyllus macrocerus (Castelnau) (Agrypninae); F, Neotetralobusafiicanus Girard (Agrypninae). Mandible (dorsal, lateral, ventral); G - I , Tetralobusflabellicornis (Linnaeus) (Agrypninae). E - I , after Costa etal. (1994).

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Fig. 4.7.6. A-J, Adult structures. Habitus (ventral): A, Opselaterpyrophanus (Illiger) (Agrypninae). Antenna; B, O. pyrophanus. Prothorax and mesothorax (lateral); C, Achrestus venustus Champion (Agrypninae). Metendosternite; D, Opselater costae Rosa. Tarsomeres; E, Dactylosimus dorsalis Fleutiaux (Elaterinae) (lateral); F - H , Anathesis laconoides Candeze (Agrypninae) (dorsal, lateral). Hind wing; I, D. dorsalis; J, Lingana illita (Candeze) (Elaterinae). B, D, after Rosa (2004); C, after Costa & Casari-Chen (1990); E - I , after Costa & Casari-Chen (1993); J, after Calder (1996).

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Cleide Costa,John F.Lawrence and Simone Policena Rosa

Fig. 4.7.7. A-H, Adult structures. Aedeagus (dorsal, ventral): A, B, Chalcolepidiuszonatus Eschscholtz (Agrypninae); C, D, platycrepidiuswapleri Salle (Agrypninae); E, F, Opselaterhelvolus (Germar) (Agrypninae). Female genitalia; G, P. wapleri; H, Achrestus venustus Champion (Agrypninae); I, Dactylosimus dorsalis Fleutiaux (Elaterinae). A, B, by C. E. Simonka; C, D, G, I, after Costa & Casari-Chen (1993). E, F, after Rosa (2004); H, after Costa & Casari-Chen (1990).

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87

a t

Of mm

κ

Fig. 4.7.8. A-M, Larval habitus (dorsal). Agrypninae: A, Dilobitarsus abbreviates Candeze; B, Chalcolepidius zonatus Eschscholtz; C, Platycrepidius bicinctus Salle; D, Pyrophorus divergens Eschscholtz; E, Thoramus laevithorax (White); F, Heteroderes rufangulus (Gyllenhal); L, Tetralobus shuckhardi (Hope). Cardiophorinae; G, Horistonotus sp. Elaterinae; H , Anchastus brunneofasciatus Schwarz; I, Physorhinus distigma Candeze. Lissominae; J, Austrelater macphersonensis Calder;

K, Lissomus sp.. Semiotinae: M, Semiotus intermedins (Herbst). Α-D, F-I, Κ, M, after Costa et al. (1988); Ε, after Costa

(1992); J, after Calder et al. (1993); L, after Girard etal. (2007).

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Fig. 4.7.9. A-I, Larval structures. Nasale: A, Thoramus laevithorax (White) (Agrypninae); B, Dilobitarsus abbreviates Cande (Agrypninae); C, Pyrophorus divergens Eschscholtz (Agrypninae); D, Lissomus sp. (Lissominae); E, Physorhinus distigma Candeze (Elaterinae); F, Semiotus ligneus (Linnaeus) (Semiotinae); G, Horistonotus sp. (Cardiophorinae). Antennae; Η, I, P. divergens (first instar and mature larva). A, after Costa (1992); B - G , I, after Costa etal. (1988); H, after Casari-Chen & Costa (1986).

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Fig. 4.7.10. A-N, Larval structures. Ventral mouthparts (maxillolabial complex): A, Τhoramus laevithorax (White) (Agrypninae); B, Physorhinus distigma Candeze (Elaterinae); C, Dilobitarsus abreviatus Candeze (Agrypninae); D, Lissomus sp. (Lissominae); E, Horistonotus sp. (Cardiophorinae); F, Semiotus ligneus (Linnaeus) (Semiotinae). Mandibles (dorsal, ventral); G, H, S. intermedius (Herbst); I, J, T. laevithorax; K, L, Pyrearinus termitilluminans Costa (Agrypninae); Μ, N, Anchastus brunneofasciatus Schwarz (Elaterinae). A, I, J after Costa (1992); B-Η, K-N, after Costa etal. (1988); K-M, after Costa etal. (1994).

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phragmotic in Tetralobini. Posterior edge not to distinctly emarginate. Epicranial stem usually absent or very short (moderately to very long in Cebrioninae); frontal arms (Fig. 4.7.9 A-G) absent (Tetralobini), lyriform, V-shaped or U-shaped, distinctly separated or contiguous at base, joined anteriorly by transverse line in Cardiophorinae. Median endocarina and paired endocarinae absent. Stemmata often absent, sometimes with a single stemma (eye spot) on each side. Labrum completely fused to anterior edge of frontoclypeal plate, often produced to form a medial nasale (Fig. 4.7.9 A-G) consisting of one to several teeth and a pair of adnasalia (Fig. 4.7.9 D, F), which may be longer than the nasale between them. Antennae (Fig. 4.7.9 H - I ) moderately long, 3-segmented, with conical, palpiform or dome-like sensorium on preapical antennomere (occasionally with compound or multiple sensoria). Mandibles (Fig. 4.7.10 G-N) symmetrical, usually either narrow and falcate, broad at base and narrow at apex, or stout and wedge-like (in Cardiophorinae elongate and apically cleft with teeth facing laterally and mesally), unidentate to tridentate, without accessory ventral process; incisor edge simple or with one or two teeth or heavily sclerotized retinacula. Mesal surface of mandibular base simple or with brush of hairs or spines; mola absent. Ventral mouthparts (Fig. 4.7.10 A-F) strongly retracted, almost always forming maxillolabial complex; maxillary articulating area absent. Cardines undivided, divided by internal ridge or externally divided; sometimes approximate or fused together at midline; maxillary apex formed by single, articulated mala or articulated galea and fixed lacinia; palp 4-segmented. Labium consisting of p r e m e n t u m and postmentum, almost always connate with maxillae to form maxillolabial complex (partially free in Cebrioninae); postmentum posteriorly truncate (e. g., in Elaterinae and Denticollinae) to narrowly rounded or acute (e. g., Agrypninae, Semiotinae, Pityobiinae), occasionally extending posteriorly beyond edges of stipites (e. g., Lissominae); ligula absent or shorter than labial palps, which are 2-segmented and narrowly to widely separated. Hypopharyngeal sclerome absent or consisting of transverse bar or mola-like sclerite. Hypostomal rods absent; ventral epicranial ridges present. Gular sutures separate or partly or completely fused. Cervical region in Cebrioninae with large eversible sac.

setae or short, stout spines; usually 5-segmented, including pretarsus, which is claw-like, usually with two setae lying side by side or obliquely situated (reduced and 4-segmented without claw setae in Thylacosterninae); coxae usually separated by less than a basal coxal diameter. Abdominal terga often simple with vestiture of fine hairs. Anterior part of tergum with transverse carina abruptly curved laterally on each side to form longitudinal carina in many Denticollinae; transverse slender striated impressions on anterolateral portions of terga in many Elaterinae; patches of short, stout setae on several terga in most Lissominae and on both terga and sterna in Senodoniini (Denticollinae). Abdominal segment IX excluding appendages usually shorter than to slightly longer than segment VIII (much longer in Cebrioninae), not forming articulated plate, always extending onto ventral surface, so that segment X (Figs.

Prothorax not longer than meso- and metathorax combined (except in Cebrioninae). Thoracic segments with or w i t h o u t tergal plates; usually without patches or rows of asperities. Prothoracic venter usually with large subtriangular presternal plate sometimes subdivided at middle, rarely with additional small posterior sclerite; sometimes with subtriangular precoxal plates. Meso- and metathorax often w i t h o u t distinct ventral plates; sometimes with reduced presternal plate, precoxalia, laterotergites (anterior one bearing spiracle on mesothorax) and small intersternites. Legs usually well developed, slender or stout, clothed with fine

4.7.11 B, D, F, H, 4.7.12 C, D, F, H) is ventrally or posteroventrally oriented; t e r g u m IX (Figs. 4.7.11 A, C, E, G, 4.7.12 A, E, G) highly variable, often with paired bifurcate urogomphi; apical prongs on each u r o g o m p h u s posteriorly oriented and slightly diverging in Agrypninae or with inner p r o n g mesally oriented and outer one projecting dorsally in most Lissominae and Denticollinae; u r o g o m p h i occasionally simple and converging (Negastriinae) or m i n u t e and hook-like (Thylacosterninae); terg u m IX simple and rounded to acute or slightly flattened with scalloped or tridentate apex in Elaterinae, Cebrioninae and Cardiophorinae; forming concave plate surrounded by serrations in Octocryptus; surface of t e r g u m IX sometimes with dorsal or lateral tubercles, longitudinal grooves, rugosities or other armature. Sternum IX almost always partly or entirely exposed (concealed in Cebrioninae), simple or sometimes U-shaped with apicolateral horns or teeth; not enclosed by s t e r n u m VIII. Segment X often forming short, cylindrical pygopod, which is flanked by paired teeth or hooks in many Agrypninae; with a pair of moderately long pygopods (Fig. 4.7.12 Α-B) in Cardiophorinae. Spiracles biforous (bilabiate), without closing apparatus; located on laterotergites, with air tubes projecting posteriorly, with anterior spiracular scar. In general it is easy to identify elaterid larvae to the genus, b u t in many cases specific identification is achieved only by rearing larvae to the adult stage. The most important larval characters include mouthparts (nasale, mandible, labial and maxillary palps, galeashape), shape of the postmentum, antennal sensoria, modifications of abdominal segment IX, the separation, subdivision or reduction of laterotergites, and spiracle location. [Hyslop 1915, 1917,1923; Emden 1932,1945; Gardner 1936; Glen et al 1943; Öhira 1962; Zacharuk 1962; Gur'yeva 1969; Costa 1970, 1972, 1982, 1992; Burakowski 1973, 1975, 1976; Dolin 1975, 1978 a, 2000 a, b; Stibick 1979; Casari-Chen & Costa 1986; Ulrich 1987; Costa et al 1988, 1992; Calder et al 1993; Öhira & Fukaishi 1999; Lawrence etal 1999,2007.]

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Morphology, P u p a (Fig. 4.7.13 A-I). Adecticous and exarate. Cream or cream-white. Body smooth or with short or long hairs or setae; sometimes with microspines. Head more or less covered by p r o n o t u m in dorsal view. Eyes small partially convered by antennae in ventral view (Fig. 4.7.13 H). Mouthparts visible in ventral view. Antennae lying parallel to ventral sides of protorax. P r o n o t u m large of variable forms, with or without pair of prominences and setae usually at the anterior margin and/or near the middle or at the h i n d angles apices. Pterothecae extending u p to third or f o u r t h ventrite. Legs free, visible in ventral view; metathoracic legs almost completely covered by the wing pads except for distal segments of tarsi, which usually extend past the third or f o u r t h ventrites. Abdomen with nine visible tergites and seven or eight visible sternites. Tergite IX with or without paired appendages similar to larval u r o g o m p h i or variable ornamentations. Lateral gin-traps (Fig. 4.7.13 C-D) present, in variable n u m b e r on abdom e n of some species of Agrypninae. Spiracles I-VII in the pleural membrane, in some species visible f r o m dorsal view. [Casari-Chen & Costa 1986; Costa etal. 1988.] P h y l o g e n y a n d T a x o n o m y . The family Elateridae has been considered to be a natural group since the earliest classifications of Coleoptera, first as a tribe of the Sternoxi (Latreille 1804,1825) and then as a family (Lacordaire 1857). The vast majority of elaterid genera appear to form a monophyletic group, b u t the outer limits of Elateridae have still not been conclusively defined, despite the efforts of various authors to establish a higher classification for the family. Several morphologically-based cladistic analyses including Elateridae were carried out by Lawrence (1988), Calder etal. (1993), Muona (1995), Lawrence et al. (1995) and Beutel (1995), and although none of these analyses were specifically directed to test the monophyly of the family or define its limits, they did lead to important conclusions which have influenced the current classification of Elateridae. Lawrence (1988) analyzed phylogenetic relationships among the families of Elateriformia and based on larval characters, as well as those of the adult prothorax, proposed the inclusion of Cebrionidae within Elateridae as a subfamily. Three of the four cladograms presented suggest Elateridae + Cebrionidae as a sister-clade of (Eucnemidae (Cerophytidae + Throscidae)). Beutel (1995) also asserted the close relationship between cebrionids and elaterids based on a larval synapomorphy: thorax and abdomen strongly sclerotized with pleural areas partially or completely concealed. In the preferred cladogram of Calder et al. (1993), Elateridae includes Oestodes (Lissominae), Protelater Sharp, Sphaenelater Schwarz (Protelaterinae) and Austrelater, together with Lissomus Dalman and Drapetes, f o r m i n g the clade Lissominae, which

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is sister group of Cussolenis Fleutiaux (Thylacosterninae). Cebrio Olivier (Cebrioninae) represents the basal clade to all remaining elaterids, and the clade ((Cerophytidae + Throscidae) Eucnemidae) is the sister group of Elateridae including Cebrioninae. In a strict consensus tree resulting f r o m Muona's analysis (1995), Elateridae, as usually delimited, is paraphyletic, in that Throscidae (sensu Crowson 1955) ((Thylacosterninae, Lissominae) Throscinae) is a monophyletic group within the elaterid clade. Cebrio was not included in the analysis, and Anischia Fleutiaux (Eucnemidae) is one of four basal taxa within the elaterid clade. Anischia was described originally in Eucnemidae and this position is confirmed by a recent cladistic analysis based on morphological and molecular data (Lawrence et al. 2007). The relationship between Elateridae and Eucnemidae was confirmed by Lawrence et al. (1995) in an analysis of 34 exemplar genera of Elateriformia (sensu lato, including Scirtoidea). Ampedus Dejean (Elaterinae) turned out as sister to Perothops Eschscholtz (Eucnemidae). Such disagreements on the position of Elateridae have been partly due to methodological differences underlying the phylogenetic analyses, b u t more importantly to bias in the nature of the data: Lawrence (1988) utilized 87 adult characters and 25 larval, Calder et al. (1993) 62 adult, 32 larval, Muona (1995) 63 adult, 6 larval, Lawrence et al. (1995) 99 adult, 23 larval, and Beutel (1995) larval characters only. It is possible that there is a great incongruence between data based on larvae and adults, producing conflicting hypotheses, depending on the proportion a m o n g those data in the analysis. In addition, not all taxa possess k n o w n larvae, and a large a m o u n t of missing data may influence the result. Muona (1995) considered the larval data the largest source of homoplasies, due to repeated re-invasions of soil or rotten wood in several lineages within Elateroidea. In opposition, Calder etal. (1993) consider the adult characters more homoplastic due to the morphological uniformity related to the evolution of a single defense mechanism (the click mechanism) in the elateroids. Sagegami-Oba et al. (2007), in the first molecular study based on partial sequences of nuclear 28S ribosomal DNA, utilized 77 elaterid species in 57 genera, primarily f r o m Japan, and 15 outgroups f r o m Throscidae, Eucnemidae, Lampyridae, Lycidae, Omethidae, Artematopodidae, Dascillidae and Carabidae. Their analysis resulted in the monophyly of the family Elateridae. However Lissominae and Thylacosterninae were not included. The elaterid subfamilies were originally defined on superficial adult characteristics, especially the structure of the antennae, mandibles, prosternum, pterothorax and tarsi (Lacordaire 1857; Candeze 1857, 1859, 1860, 1863, 1874; Schwarz 1906 a, b, 1907 a, b; Fleutiaux 1947). Hyslop (1917, 1923) first made use of larval features to define elaterid subfamilies, and other more recent workers (see above) have described and illustrated the larval

Fig. 4.7.11. A-H, Larval structures. Terminal abdominal segments (dorsal, ventral) (Agrypninae): A, B, Heteroderes rufangulus (Gyllenhal); C, D, Pyrophorus divergens Eschscholtz (first instar); E, F, P. divergens (mature larvae); G, Η, Platycrepidius bicinctus Salle. A-H, after Costa et al. (1988).

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Fig. 4.7.12. A-H, Larval structures. Terminal abdominal segments: A, B, Horistonotus sp. (Cardiophorinae) (dorsal, ventral); C, Anchastus brunneofasciatus Schwarz (Elaterinae) (ventral); D, Physorhinus distigma Candeze (Elaterinae) (ventral); E, F, Semiotus intermedius (Herbst) (Semiotinae) (dorsal, ventral); G, H, Lissomus sp. (Lissominae) (dorsal, ventral). A-H, after Costa etal. (1988).

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Fig. 4.7.13. A-I, Pupal habitus. Agrypninae: A, Dilobitarsus abbreviates Candeze (Idorsal); B, Platycrepidus bicinctus Salle (dorsal); C, Chalcolepidius zonatus Eschscholtz (dorsal); D, Pyrophorus divergens Eschscholtz (dorsal); E, Thoramus laevithorax (White) (dorsal). Elaterinae; F, Anchastus brunneofasciatus Schwarz (dorsal); G, Tetralobus shuckhardi (Hope) (dorsal); H, Tetralobus shuckhardi (dorsal, ventral). Semiotinae; I, Semiotus intermedius (Herbst) (dorsal). Α-D, F, I after Costa etal. (1988); E, after Costa (1992); G, H, after Girard etal. (2007).

Elateridae Leach, 18Ί 5

morphology in many groups. Classifications based on specific adult character systems were produced by Gur'yeva 1974 (prothorax and mesothorax) and Dolin 1975 (wing venation) or on fossil forms (Dolin 1978 b). Classifications of both Dolin and Gur'yeva included 10 subfamilies: Agrypninae, Pityobiinae, Oxynopterinae, Diminae, Tetralobinae, Negastriinae, Denticollinae (as Athoinae), Oestodinae, Elaterinae and Cardiophorinae. According to Dolin (1978 b), the Protoagrypninae originated in the Lower Jurassic but were extinct by the beginning of the Cenozoic, when four new lineages arose: 1) Agrypninae, which gave rise to Pityobiinae and Oxynopterinae; 2) Diminae, which produced the Tetralobinae; 3) Negastriinae, from which evolved the Denticollinae and Oestodinae; and 4) Cardiophorinae, which produced the Elaterinae. The classification in Lawrence & Newton (1995) is a synthesis of the above works and others produced over the past seven decades (e. g., Emden 1932; Fleutiaux 1947; Nakane & Kishii 1956; Crowson 1961; Laurent 1961, 1966; Öhira 1962; Burakowski 1973, 1975; Stibick 1979; Costa & Casari-Chen 1993; Calder et al. 1993; Muona 1995). In that work, Elateridae was divided into 18 subfamilies: Cebrioninae, Tetralobinae, Thylacosterninae, Lissominae, Semiotinae, Pityobiinae, Oxynopterinae, Agrypninae, Denticollinae, Negastriinae, Diminae, Elaterinae, Cardiophorinae, Hemiopinae and Physodactylinae, Eudicronychinae, Anischiinae and Subprotelaterinae, the last three of which were of uncertain position. These authors were unaware of an additional subfamily, Campyloxeninae, which was proposed by Costa (1975). Monophyly of some of the above subfamilies was corroborated by Calder et al. (1993), but their analysis was based on only one or two exemplar genera for each group. In a strict consensus cladogram of nine elaterid subfamilies, the clade (((Pityobiinae + Agrypninae) (Cardiophorinae + Elaterinae)) + Denticollinae) formed part of a polytomy with Lissominae, Thylacosterninae, Semiotinae, Tetralobinae, Cebrioninae and (Eucnemidae (Cerophytidae + Throscidae). Sagegami-Oba et al. (2007) in their molecular phylogeny included representative species of the following subfamilies (as delimited below): Agrypninae (14), Oxynopterinae (2, one of which, Pectocera fortunei Candeze, was listed as Pityobiinae), Denticollinae (22, including four Hypnoidini), Elaterinae (27, including five Melanotini), Negastriinae (6), Cardiophorinae (6). Their resulting cladograms were compared with those produced by Stibick (1979) and Öhira (1999). Trees resulting from neighbor-joining, maximum likelihood and parsimony agreed in general topology with four distinct clades: D (Elaterinae including Melanotini) basal to the remaining members of the family, and A (Agrypninae) basal to C (Negastriinae + Cardiophorinae) and Β (Denticollinae, including Hypnoidini, Campsosternus and Pectocera Hope). A similar topology was arrived at by Oba (2007) using Bayesian analysis. In the recent

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analysis of Lawrence et al. (2007), based on both morphological and molecular data but with the exemplar taxastrongly skewed in favor of Eucnemidae, Thylacosterninae and Lissominae consistently formed a monophyletic group within Elateridae. However only three other elaterid taxa (Danosoma, Ampedus and Selatosomus Stephens) were included and their interrelationships were unresolved. The following is an alphabetical list of the subfamilies recognized in this work, with notes on their constitution and characteristics and lists of recognized tribes. The oldest family group names have been used in all but four cases. Oophorini, Dendrometrinae, Prosternini and Synaptini proposed by Gistel (1856) and used in some recent works (Sanchez-Ruiz 1996; Johnson 2 0 0 2 a, 2000 b) have been replaced by the junior names Monocrepidiini, Denticollinae, Ctenicerini and Adrastini, respectively, pending a decision by the ICZN on the suppression of these senior synonyms (Cate 2007). Agrypninae Candeze. This cosmopolitan subfamily containing about 120 genera and 2500 species was once referred to as Pyrophorinae and includes a number of formerly recognized subfamilies, such as Chalcolepidiinae, Hemirhipinae, Octocryptinae and Tetralobinae. Adults of all taxa share two synapomorphies: presence of one or more stout setae arising from the outer side of each pretarsal claw and absence of a wedge cell in the hind wing (Crowson 1961; Calder et al. 1993, Calder 1998). Larvae may be recognized by a combination of a triangular, posteriorly acute postmentum, simple mandibles (without retinaculum), and lightly sclerotized trunk segments with a notched tergum IX (Hayek 1973, 1979). The group includes eight tribes: Agrypnini Candeze, Anaissini Golbach, Cleidecostini Johnson (= Heligmini Costa), Hemirhipini Lacordaire, Monocrepidiini Candeze (= Conoderini Fleutiaux, Oophorini Gistel), Platycrepidiini Costa and Casari-Chen, Pseudomelanactini Arnett, Pyrophorini Candeze and Tetralobini Laporte. Fleutiaux's name Conoderini is a junior homonym of Conoderini Schoenherr which is currently used in Curculionidae (AlonzoZarazaga & Lyal 1999). Cleidecostini was proposed as a replacement for Heligmini Costa because Heligmus Candeze is preoccupied by a nematode genus (Johnson 2002 b). Many workers recognize Tetralobini at the subfamily level, although Stibick (1979), Costa et al. (1992, 1994), Calder (1998) and Girard et al. (2007) considered it as a tribe of Agrypninae. The tribe Hemirhipini was redefined by CasariChen (1993) and a phylogenetic hypothesis of the included genera (Casari-Chen 1994) indicated that some genera were not monophyletic, requiring specific revisions (Casari 2002). Campyloxeninae Costa. This subfamily contains the single Neotropical species Campyloxenus pyrothorax, which was placed among the agrypnines in older classifications based mainly on the

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prothoracic light organs in the adult. Costa (1975) removed it from that subfamily because of the lack of claw setae and presence of a wedge cell in the hind wing and placed it in a separate subfamily. Larvae are unknown.

those of Elaterinae in having a posteriorly truncate postmentum, but tergum IX in Denticollinae is always notched, so that paired urogomphi are formed. Included tribes are Ctenicerini Fleutiaux, Crepidomenini Candeze, Denticollini Stein & Weise, Dimini Candeze, Hypnoidini Schwarz, Pleonomini Semenov & Pjatakova, and Senodoniini Schenkling. The last tribe is considered by Schimmel (1996) to be sister to or embedded within Dimini. However the larva of Senodonia Laporte, illustrated by Dolin (2000 b) and examined by one of us (JL), is of a very different type than those of either Ditna Carpentier (Dolin 1978 a) or Venia Laporte (Dolin 1990), resembling those of various Lissominae in the structure of the mandible, postmentum and abdominal apex and in having movable spines of the lissomine type on abdominal terga and sterna (Dolin 2000 a).

Cardiophorinae Candeze. This subfamily, which contains about 35 genera and 800 species (Calder 1998), is easily recognizable based on the unique larvae (Fig. 4.7.8 G), which are predators in soil or rotten wood. Adults are usually distinguished from those of all other groups by a combination of laterally closed mesocoxal cavities, short, truncate prosternal process, cordate scutellar shield and lateral pronotal carinae which are incomplete or absent. The listing of Cardiorhinus Eschscholtz and Hypodesis Germar under this subfamily by Lawrence & Newton (1995) was a lapsus. The Australian genus Patriciella was moved from Physodactylini to this subfamily by Calder (1996). An unpublished phylogenetic analysis based on adult morphology of 27 cardiophorine genera and outgroups from Negastriinae and Hypnoidini indicates that major changes are indicated in the constitution and tribal level classification of this group (Douglas & Peck 2005). Cebrioninae Latreille. This group includes those genera normally placed in the family Cebrionidae (e. g., Anakstesa Leach, Cebrio, Cebriorhipis Chevrolat, Musopsis Chevrolat, Scaptoknus LeConte, Selonodon Latreille, Stenocebrio Solervicens), plus those comprising the tribe Aplastini of Stibick (1979) (= Plastoceridae of authors, not Crowson) (e. g., Aplastus LeConte, Euthysanius, Octinodes Candeze). Paulusiella Mandl, placed in the family Karumiidae by Paulus (1972) and Mandl (1974), also belongs to this group. Adults are usually of moderate size and often relatively soft-bodied, with projecting falcate mandibles and a short, broad labrum. The antennae may be filiform, serrate or pectinate, the prosternum truncate, without a chin-piece, the legs sometimes fossorial and often with enlarged spurs, and the abdomen with five or six ventrites. Sexual dimorphism is often well marked, the females usually having shorter antennae, often lacking hind wings and occasionally with very short elytra exposing several abdominal segments. The larvae are easily recognized by the long prothorax and the enlarged and eversible cervical membrane (Hyslop 1923; Becker 1991). [Chevrolat 1874; Schwarz 1907 b; Van Dyke 1932; Solervicens 1988; Galley 1999.] Denticollinae Stein & Weise. This cosmopolitan subfamily, with about 150 genera and 1500 species (Calder 1998), includes Diminae and Hypnoidinae, as well as the tribe Pleonomini, which was elevated to subfamily rank by Laurent (1966) but placed in Aplastinae by Stibick (1979). Adults are not easily separable from members of the Elaterinae, but in general the head is more flattened and prognathous than in the latter group. Larvae resemble

Elaterinae Leach. This group comprises some 200 genera and 3500 species worldwide (Calder 1998). Elaterine adults, in general, have a more globular and deflexed head than those of Denticollinae, but exceptions occur. Larval Elaterinae (Figs. 4.7.8 H - I , 12 C-D) never have a notch on tergum IX, which may be rounded, conical, apically acute or (in Melanotini) concave with one or more teeth at the apex. Casari (2008) carried out a phylogenetic analysis of the 31 genera of Dicrepidiini (considered to be a subtribe of Ampedini), concluding that the group appears to be a sister group of Physorhinini and that the Chilean genus Ovipalpus Solier should be removed and probably placed in Denticollinae. Included tribes are Adrastini Candeze, Agriotini Champion, Ampedini Gistel, Dicrepidiini Candeze, Elaterini Leach, Melanotini Candeze, Odontonychini Girard, Physorhinini Candeze and Pomachiliini Candeze. [Girard 1972; Hayek 1990.] Eudicronychinae Girard. Three Afrotropical genera, Eudicronychus Mequignon (= Dicronychus Brulle, not Laporte), Anisomerus Schwarz and Tarsalgus Candeze, were placed in a separate family Dicronychidae by Schwarz (1907 c) based in part on the unusual form of the aedeagus, and this placement was followed by Girard (1986 b, 1991). Dolin (1975) considered the group to be "outside the system" but pointed out that the hind wing venation is similar to that in the elaterine tribe Dicrepidiini. The taxon was treated as a synonym of Cardiophorinae by Jeannel (1955), Arnett (1960), Leseigneur (1972) and Stibick (1979), but that was based on confusion between Dicronychus Brulle (1832), an Oriental cardiophorine genus, and Dicronychus Laporte (1840), which was subsequently renamed Eudicronychus by Mequignon (1931). Larvae are unknown and the affinities of the group are still uncertain. Hemiopinae Fleutiaux. This subfamily includes several genera related to Hemiops Laporte and Parhemiops Candeze. The position of this group is

Elateridae Leach, 1815

uncertain, and Stibick (1979) placed Hemiops in the tribe Oestodini. Larvae are u n k n o w n . Lissominae Laporte. This group, which is found in all Zoogeographie regions, comprises about 10 genera and 150 species. It is used here in a broad sense to include not only Lissomini, b u t also Oestodini and Protelaterini, plus the Australian genus Austrelater (Calder et al. 1993; Calder 1998). The genus Oestodes is usually considered to form a monophyletic group with the Notogean Protelaterini, b u t the similarities of the latter with Lissomini are based mainly on larval characters, such as the emarginate clypeolabrum, large, complex retinacul u m and the presence of specialized movable spines on the dorsal surface (except in Oestodes). Crowson (1961) suggested that the lissomine genus Drapetes be moved to Elateridae, b u t retained Lissomus in Throscidae, while Burakowski (1975) placed b o t h genera in the family Lissomidae. Austrelater, which shares the unusual type of larva, differs f r o m b o t h groups in adult features, superficially resembling a cebrionine. Costa et al. (1988) described the larva of Lissomus and suggested a close relationship of that genus with Semiotus based on the deeply emarginate clypeolabrum and shape of the anal region. Excluded f r o m this subfamily (and f r o m Elateridae) is the highly modified, termitophilous beetle Neocrowsonia viatoricus Kistner & Abdel-Galil (1986). The species was placed in Throscidae by its describers, following a suggestion by R. A. Crowson, b u t as pointed out by Lawrence & Newton (1995), the suggestion was probably based on the presence of well developed tarsal lamellae resembling those in Lissomini. The genus is treated briefly in 2-4.18. M o r o s t o m i n a e Dolin. This subfamily was proposed by Dolin (2000 a, 2000 b) for Morostoma Candeze (originally placed in Allotriites (= Senodoniini) by Candeze 1879), plus Diplophoenicus and several other Malagassy genera which Fleutiaux (1910, 1929) had placed together as a group within the Plastoceridae (Dolin & Girard 2003). The genus Morostoma has very elongate labial and maxillary palps (Fig. 4.7.5 A, B) and Diplophoenicus has long, strongly flabellate, 12-segmented antennae. Negastrinae Nakane & Kishii. This small group includes about 26 genera and 300 species worldwide which are most commonly found in riparian situations, on sand banks of rivers and streams (Calder 1998). These small beetles have laterally closed mesocoxal cavities as in Cardiophorinae, b u t they differ from that group in having a normally long prosternal process and a non-cordate scutellar shield. Larvae have a posteriorly truncate postmentum and notched tergum IX, as in Denticollinae, b u t the galea is 1-segmented (Stibick 1971). Keys and descriptions to world genera were provided by Stibick (1971), a characterization of the subfamily and key to European genera by Leseigneur (1970), and a species revision for eastern North America by Stibick (1990).

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Oxynopterinae. As used here, Oxynopterinae includes about ten genera and 50 species occurring mainly in the Oriental region (e. g., Oxynopterus Hope, Campsosternus, Pectocera, Ceropectus Fleutiaux, and Rostricephalus Fleutiaux) and three New World genera (e. g., Oistus, Perissarthron and Mehmetes LeConte) containing about 20 species (Johnson 2002 a). The genus Semiotus, included here by Gur'yeva (1974), Dolin (1975) and Stibick (1979) is placed in a separate subfamily. The predaceous oxynopterine larvae have a triangular postment u m , simple mandibles and a complex antennal sensorium (Öhira 1962; Öhira & Fukaishi 1999), and at least one species of Oxynopterus is a predator in termite nests (Ralshoven 1955). Physodactylinae Lacordaire. Eight genera f r o m South America, Southeast Asia and Africa have been placed in this group based on falciform mandibles (at least in one sex) and digging h i n d legs (Fleutiaux 1892), b u t larvae are u n k n o w n and the group may not be monophyletic. The Australian genera Antoligostethus Blackburn and Nullarborica Blackburn were transferred to Elaterinae by Calder (1996). A key to all genera is given by Schwarz (1906 b) and a key to some South American species may be f o u n d in Fleutiaux (1892). Pityobiinae Hyslop. This subfamily consists of eight genera and 20 species. The concept of Pityobiinae used here corresponds to that of Gur'yeva (1974), Dolin (1975), Ulrich (1987) and Calder (1992,1996), including not only the North American PityoUus, b u t also the New Zealand Metablax Candeze, the South American Tibionema Solier and the Australian genera Parablax Schwarz, Parasaphes Candeze, Tasmanelater Calder, Xuthelater Calder and Wynarka Calder. Larvae are predaceous and occur in rotten wood. The genus Pectocera, included here by Stibick (1979), is placed in Oxynopterinae. S e m i o t i n a e Jacobson. As here defined, this group includes two Neotropical genera: Semiotus, with 75 species and Semiotinus Pjatakowa with 17 (Wells 2003; Parreira & Casari 2004). The genus Oistus, placed here by Golbach (1970), is included in Oxynopterinae. Semiotus was originally placed near the agrypnine genus Chalcolepidius and later moved to Oxynopterinae, based partly on the solid fusion of the meso- and metaventrites, a feature which has evolved at least three times in the family. In addition, adult semiotines are generally brightly colored and subglabrous with a complete frontal carina, sometimes with projecting spines, mandibles are bifid at the apex or at least equipped with a small, subapical tooth, elytral apices are sharp, often bifid, and tarsi have two or three ventral lobes. The predaceous, wood-inhabiting larvae of Semiotus are of an unusual type, with the frontal arms incomplete, adnasalia projecting well in f r o n t of the truncate or bilobed nasale, antennal sensorium simple, mandibles grooved dorsally, and sternum IX more or less

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U-shaped with a tooth at the end of each arm (Costa 1972; Costa etal. 1988).

Universidade de Säo Paulo and also for permission to use several of his illustrations. Thanks are also given to Conselho Nacional de Desenvolvimento Cientifico e Tecnologico (CNPq) for the Research Grant 302721/2007-0 to C. Costa. CSIRO Australia is acknowledged for support of research (JFL) and for allowing us to use copyrighted images.

Subprotelaterinae Fleutiaux. This subfamily

includes the single genus Subprotelater. Cobos (1959) transferred the genus from Eucnemidae to Elateridae based on the original description and this was confirmed by Muona (1987) based on a study of the holotype of S. bakeri Fleutiaux. The lateral antennal grooves on the pronotal hypomera resemble those in some Eucnemidae, but the hind wing lacks a wedge cell and resembles those of Agrypninae. Thylacosterninae Fleutiaux. This group, which occurs in South America, Africa, Asia and northern Australia, includes about 45 species in five genera: Thylacosternus Bonvouloir, Balgus Fleutiaux, Pterotarsus Guerin-Meneville, Cussolenis Fleutiaux and Lumumbaia Muona & Vahtera (Vahtera et al. 2009). The thylacosternines were placed in the family Eucnemidae by Fleutiaux (1902, 1920), although Lameere (1900) had previously noted their similarity to Elateridae based on adult features, and both Emden (1932) and Gardner (1936) supported this view based on characters of the larvae. Crowson (1955, 1961) treated thylacosternines as a subfamily of Throscidae, while in cladograms produced by Calder etal. (1993), Cussolenis was placed either at the base of a clade containing Cerophytidae, Throscidae and Eucnemidae or at the base of a clade containing all groups comprising Lissominae, as delimited above. In cladograms produced by Lawrence et al. (2007), Cussolenis and Pterotarsus formed a monophyletic group within an elaterid clade also including Austrelater, Drapetes and Lissomus. Although adults of Balgus schnusei emit green light from a pair of luminous spots on the prothorax (Costa 1984 a), bioluminescence has not been reported for the other three species in this Neotropical genus.

Acknowledgements We are very indebted to Carlos Esteväo Simonka (MZUSP, Säo Paulo, Brazil) for the excellent graphic project, which included the electronic treatment of the figures, the layout and the final art of the plates. The following individuals are also gratefully acknowledged: Etelvino Jose Henriques Bechara (Instituto de Quu'mica, USP, Säo Paulo) and Sergio Ide (Instituto Biologico, Säo Paulo) for critical revision of the manuscript and useful suggestions, Claude Girard (MNHN, Paris, France), Pierre Rasmont (editor Annales de la Societe Entomologique de France, Paris, France), Jyrki Muona, Paul Johnson and Hume Douglas for advice and information, Andrew A. Calder (CSIRO, Canberra, Australia) and Sönia A. Casari (MZUSP) for permission to use several of their illustrations, and Jacques Chassain (Thomery, France) for the donation of specimens of Morostoma palpale to the Museu de Zoologia,

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(1975): Bibliographie der Elateridenlarven-Literat u r d e W e l t [to 1968-1972].-Beiträge zur Entomologie 25: 85-98. - (1979): Bibliographie der Elateridenlarven-Literat u r d e W e l t [to 197 3-1977}.-Beiträge zur Entomologie 29:307-317. Galley, Κ. Ε. Μ. (1999): Revision of the genus Selonodon Latreille (Coleoptera: Cebrionidae). - Occasional Papers of the Florida State Collection of Arthropods 10, v + 49pp. Gardner, J. C. M. (1936): A larva of the subfamily Balginae (Col., Elateridae). - Transactions of the Royal Entomological Society ofLondon 5 (1): 3 0 - 3 2 . Girard, C. (1971): Les coleopteres Elateridae de Lamto (Cote d'lvoire). - Bulletin de l'Institut Fundamental d'Afrique Noire 33 (A 3): 5 4 9 - 650. - (1972): Contribution a l'etude des Elateridae de l'Afrique noire (Col.). - Bulletin de la Societe Entomologique de France 1 7 : 2 7 6 - 2 8 3 . - (1986 a): Sept nouvellles especes de Dicrepidinae d u genre propsephus Hyslop (Coleoptera, Elateridae). - Νouvelle Revue d'Entomologie (N. S.) 3 (4): 473-483. - (1986 b): Les especes d u groupe Anisomerus prosternalis Schwarz (Coleoptera Dicronychidae). - Revue ZoologiqueAfricaine99:313-320. - (1991): Sept nouvelles especes afrotropicales de Dicronychidae d u genre Eudicronychus Mequignon (Coleoptera). - Bulletin de la Societe Entomologique de France 96 (2): 145-154. - (2003): Etude des peuplements d'Elateridae de la region d u Mt Nimba et descriptions de taxons nouveaux, in Lamotte, M. & Roy, R. Le peuplement animal d u Mont Nimba (Guinee, Cöte d'lvoire, Liberia). - Mdmoires du Museum national d'Histoire naturelle 1 9 0 : 3 9 3 - 5 4 9 . Girard, C., Costa, C. & Rosa, S. P. (2007): Presence insolite des larves de Tetralobus dans les Termitieres de Μ acrotermes: bionomic donnees et morphologie des larves et des nymphes de trois especes (Coleoptera, Elateridae, Tetralobinae). -Annales de la SocieteEntomologique de France (n. s.) 43 (1): 4 9 - 5 6 . Glen, R., King, Κ. M. & Arnason, A. P. (1943): The identification of wireworms of economic importance in Canada. - Canadian Journal ofResearch 2 1 : 3 5 8 - 3 8 7 . Golbach, R. (1970): Semiotinae, nueva subfamilia de Elateridae (Col.). - Acta Zoologica Lilloana 25 (24): 319-322. Gur'yeva, Ye. L. (1969): Some trends in the evolution of click beetles (Coleoptera, Elateridae). - Entomologicheskoye Obozreniye 48: 2 6 3 - 2 7 2 (in Russian; translation in Entomological Review, Washington 48: 154-159). - (1974): Thoracic structure of click beetles (Coleoptera, Elateridae) and the significance of the structural characters for the system of the family. - Entomologicheskoye Obozreniye 5 3 : 9 6 - 1 1 3 . (in Russian; translation in Entomological Review, Washington 53: 67-79). Hayek, C. M. F. von (1973): A reclassification of the subfamily Agrypninae (Coleoptera: Elateridae). -Bulletin of the British Museum (Natural History) Entomology Supplement 2 0 : 1 - 3 0 9 . - (1979): Additions and corrections to Ά reclassification of the subfamily Agrypninae (Coleoptera:

Elateridae Leach, 1815

Elateridae)'. - Bulletin of the British Museum of Natural History (Entomology) 3 8 : 1 8 3 - 2 6 1 . - (1990): A reclassification of the Melanotus group of genera (Coleoptera: Elateridae). - Bulletin of the British Museum ofNatural History (Entomology) 59 (1): 37-115. Hyslop, J. A. (1915): Wireworms attacking cereal and forage crops. - Bulletin of the United States Department ofAgriculture 1 5 6 : 1 - 3 4 . - (1917): The phylogeny of the Elateridae based on larval characters. -Annals of the Entomological Society ofAmerica 1 0 : 2 4 1 - 2 6 3 . - (1923): The present status of the coleopterous family Plastoceridae. - Proceedings of the Entomological Society of Washington 2 5 : 1 5 6 - 1 6 0 . Jeannel, R. (1955): L'edeage. Initiation aux recherches sur la systematique des coleopteres. - Publication du Museum National d'Histoire Naturelle de Paris 16, 155 p. Johnson, P. J. (2002 a): 58. Elateridae Leach 1815. Pp. 160-173 in Arnett, R. H., Jr., Thomas, M. C., Skelley, P. E. & Frank, J. H. (eds.) American Beetles. Volume 2. polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Gainesville, Florida. - (2002 b): Lectotype designations for Elateridae (Coleoptera) described by George C. Champion in the Biologia Centrali-Americana. - Dugesiana 9 (1): 15-46. Kaishoven, L. E. G. (1955): Additional note on the giant elaterid, Oxynopterus mucronatus Ol., a predator on termites in Java. -Entomologische Berichten 15 (13): 273-278. Kistner, D. Η. & Abdel-Galil, F. Α. (1986): A new genus and species of termitophilous Throscidae f r o m South Africa (Coleoptera).-Sociobiology 12(2): 3 0 5 314. Lacordaire, J. T. (1857): Histoire Naturelle des Insectes. Genera des Coleopteres ou expose methodique et critique de tous le genres proposes jusqu'ici dans cet ordre d'insectes, Vol. 4, 579 pp. Librairie Encyclopedique de Roret, Paris. Lameere, A. (1900): Notes pour la classification des Coleopteres. - Annales de la Societe Entomologique de Belgique 4 4 : 3 5 5 - 3 7 7 . Laporte, F. L. (1840): Histoire Naturelle des Animaux Articules, Annelides, Crustaces, Arachnides, Myriapodes et Insectes. Vol. 2. Histoire Naturelle des Insectes, Coleopteres. Vol. 1.324 pp. Dumeril, Paris. Latreille, P. A. (1804): Histoire Naturelle, Generale etParticuliere des Crustaces et des Insectes. Families Naturelles des Genres. F. Dufart, Paris. - (1825): Families Naturelles du Regne Animal, Exposees Succinctemente et dans un Ordre Analytique, avec Vindication de leurs Genres. Bailliere, Paris. Laurent, L. (1961): Sur la suture meso-metasternale chez les Elateridae (Coleoptera). - Bulletin de la Sociite Royale des Sciences de Liege 3 0 : 6 8 - 7 1 . - (1966): Denticollinae, Pleonominae, Athoomorphinae de la region aethiopienne (Coleoptera, Elateridae). - Bulletin de la Societe Royale des Sciences de Liege 3 5 : 8 0 1 - 8 2 1 . Lawrence, J. F. (1982): Coleoptera. Pp. 4 8 2 - 5 5 3 in Parker, S. P. (ed.) Synopsis and Classification of Living Organisms, Vol. 2. McGraw-Hill, New York.

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(1988): Rhinorhipidae, a new beetle family f r o m Australia, with comments on the phylogeny of the Elateriformia. - Invertebrate Taxonomy 2 (1987): 1-53. Lawrence, J. F. & Newton, A. F., Jr. (1995): Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names). Pp. 7 7 9 - 1 0 0 6 in Pakaluk, J. & Slipiriski, S. A. (eds) Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson, Vol. 2. M u z e u m i Instytut Zoologii PAN, Warszawa. Lawrence, J. F., Nikitsky, Ν. B. & Kirejtshuk, A. G. (1995): Phylogenetic position of Decliniidae (Coleoptera: Scirtoidea) and comments on the classification of Elateriformia (sensu lato). Pp. 3 7 5 - 4 1 0 in Pakaluk, J. & Slipiriski, S. A. (eds) Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson, Vol. 1. M u z e u m i Instytut Zoologii PAN, Warszawa. Lawrence, J. F., Hastings, A. M., Dallwitz, Μ. J., Paine, T. A. & Zürcher, Ε. J. (1999 a): BeetleLarvae of the World: Descriptions, Illustrations, Identification, and Information Retrieval for Families and Subfamilies." CD-ROM, Version 1.1 for MS-Windows. CSIRO Publishing, Melbourne. - (1999 b): Beetles of the World: A Key and Information System for Families and Subfamilies. CD-ROM, Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne. Lawrence, J. F., Muona, J., Teräväinen, Μ., Stahls, G. & Vahtera V. (2007): Anischia, Perothops and the phylogeny of Elateroidea (Coleoptera: Elateriformia). - Insect Systematics and Evolution 3 8 : 2 0 5 - 2 3 9 . Leseigneur, L. (1970): Revision des Zorochorus Europeens. - Bulletin Mensuel de la Societe Linneenne de Lyon 39 (1): 19-44. - (1972): Coleopteres Elateridae de la faune de France continentale et de Corse. - Bulletin Mensuel de la Societe Linneenne de Lyon 41 (supplement), 379 pp. Löbl, I. (2007): Elateridae: Subprotelaterinae, p. 207 in Löbl, I. & Smetana, A. (eds) Catalogue of Ρalaearctic Coleoptera. Volume 4. Elateroidea - Derodontoidea - Bostrichopidea - Lymexyloidea - Cleroidea - Cucujoidea. Apollo Books, Stenstrup. Mandl, Κ. (1974): Eine neue Gattung, drei neue Arten u n d eine neue Unterart aus der Familie Karumiidae (Dascilloidea, Col.). - Verhandlungen der Naturforschenden Gesellschaft in Basel 84: 635-645. Mequignon, A. (1931): Notes synonimiques sur quelques Elaterides (Col.) 5 e m e note. - Bulletin de la Societe Entomologique de France 1931: 207-208. Muona, J. (1987): The generic names of the beetle family Eucnemidae (Coleoptera). - Entomologica Scandinavica 1 8 : 7 9 - 9 2 . - (1995): The phylogeny of Elateroidea (Coleoptera), or which tree is the best? - Cladistics 9 : 4 1 3 - 4 2 6 . Nakane, T. & Kishii, T. (1956): On the subfamilies of Elateridae f r o m Japan (Coleoptera). - Kontyü 24: 201-206, pis. 2 2 - 2 3 . Oba, Y. (2007): Molecular phylogenetic analysis of click beetles (Coleoptera: Elateridae). - Coleopterists' News 1 5 7 : 7 - 1 0 (in Japanese; English summary).

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Öhira, Η. (1962): Morphological and Taxonomic study on the larvae ofElateridae of Japan (Coleoptera), iv + 179 pp., 61 pis. H. Ohira, Okasaki (in Japanese; English summary). - (1976): New or little-known Elateridae (Coleoptera) from Japan, XX. - Transactions of the Shikoku Entomological Society 1 3 : 4 7 - 5 0 . - (1999): The systematics of subfamilies in Elateridae from Japan. Booklet for a lecture in the 17th annual meeting of the Japanese Society of Coleopterology. 4 pp. Öhira, Η. & Becker, Ε. C. (1973): Elateridae (Coleoptera) from the Canadian Nepal expedition (1967) 6. Description of two new species of Acteniceromorphus of the subfamily Ctenicerinae. - Oriental Insects 7 (4): 563-566. Öhira, Η. & Fukaishi, Τ. (1999): The larva of Campsosternus matsumurae (Coleoptera, Elateridae) from Ishigaki-jima Is. of the Ryukyu Islands, Japan. -Elytra, Tokyo 27(1): 159-165. Parreira, Μ. H. & Casari, S. A. (2004): Morphology of three Brazilian species of Semiotus (Coleoptera, Elateridae, Semiotinae). - Iheringia, Series Zoologia, Porto Alegre 94 (3): 285-294. Paulus, Η. F. (1972): Die systematische und phylogenetische Stellung der Karumiidae mit einer Beschreibung von Escalerina serraticornis n. sp. aus S-Persien. -SenckenbergianaBiologica 5 3 : 3 7 - 5 4 . Redford, Κ. H. (1982): Prey attraction as a possible function of bioluminescence in the larvae of Pyrearinus termitilluminans (Coleoptera: Elateridae). - Revista Brasileira de Zoologia 1 , 3 1 - 3 4 . Rosa, S. P. (2004): Revisäo do genero Opselater Costa (Coleoptera, Elateridae, Agrypninae). - Revista Brasileira deEntomologia 48 (2): 203-219. Sagegami-Oba, R., Oba, Y. & Öhira, Η. (2007): Phylogenetic relationships of click beetles (Coleoptera: Elateridae) inferred from 28S ribosomal DNA: Insights into the evolution of bioluminescence in Elateridae. - Molecular Phylogenetics and Evolution 42 (2): 410-421. Sänchez-Ruiz, A. (1996): Catälogo bibliogräfico de las especies de la familia Elateridae (Coleoptera) de la Peninsula Iberica e Isias Baleares. - Documentos FaunalbericaZ: 1-265. Schwarz, O. (1906 a): Genera Insectorum deP. Wytsman. Fase. 46A. Coleoptera. Farn. Elateridae. Pp. 1 - 1 1 2 . P. Wytsman, Brussels. - (1906 b): Genera Insectorum de P. Wytsman. Fase. 46B. Coleoptera. Farn. Elateridae. Pp. 113-224. P. Wytsman, Brussels. - (1907 a): Genera Insectorum de P. Wytsman. Fase. 46C. Coleoptera. Farn. Elateridae. Pp. 225-370, pis. 1 - 6 . P. Wytsman, Brussels. - (1907 b): Genera Insectorum de PI. Wytsman. Fase. 50. Coleoptera. Fam. Plastoceridae. 10 pp., 1 pl. P. Wytsman, Brussels. - (1907 c): Genera Insectorum de P. Wytsman. Fase. 51. Coleoptera. Fam. Dicronychidae. 5 pp., 1 pl. P. Wytsman, Brussels. Sivinsky, J. (1981): The nature and possible functions of luminescence in Coleoptera larvae. - The ColeopteristsBulletin 35 (2): 167-179. Smetana, A. (2007): Elateridae: Lisominae, pp. 46, 207-208 in Löbl, I. & Smetana, A. (eds), Catalogue of

Palaearctic Coleoptera. Volume 4. Elateroidea - Derodontoidea - Bostrichopidea - Lymexyloidea - Cleroidea - Cucujoidea. Apollo Books, Stenstrup. Solervicens, J. (1988): Stenocebrio coquimbensis (Coleoptera: Cebrionidae), nuevo genero y especie y primera cita de esta familia para Chile. - Revista Chilena deEntomologia 1 6 : 1 5 - 2 1 . Stibick, J. N. L. (1971): The generic classification of the Negastriinae (Coleoptera, Elateridae). - Pacific Insects 13 (2): 371-390. - (1979): Classification of the Elateridae (Coleoptera). Relationships and classification of the subfamilies and tribes.-PacificInsects 20 (1): 145-186. - (1990): North American Negastriinae (Coleoptera, Elateridae): The Negastriinae of the eastern United States and adjacent Canada. Insecta Mundi 4: 99-131. Stolz, U., Velez, S., Wood, Κ. V., Wood, M. & Feder, J. L. (2003): Darwinian natural selection for orange bioluminescent color in a Jamaican click beetle. - Proceedings of the National Academy of Sciences 100: 14955-14959. Stone, M. W. (1950): An unusual record of longevity for an elaterid larva.-Journal of Kansas Entomological Society 23 (4): 126-128. Timmins, G. S., Penati, C. Α. Α., Bechara, E. J. H. & Swartz, Η. M. (1999): Measurement of oxygen partial pressure, its control during hypoxia and hyperoxia, and its effect upon light emission in a bioluminescent elaterid larva. - Journal of Experimental Biology 202: 2631-2638. Timmins, G. S., Jackson, S. K., Swartz, Η. M. (2001): The evolution of bioluminescent oxygen consumption as an ancient oxygen detoxification mechanism. -Journal of Molecular Evolution 5 2 : 3 2 1 - 3 3 2 . Ulrich, G. W. (1987): The Phylogeny of the Pityobiinae based upon larval morphology (Elateridae: Coleoptera). 169 pp. University of California (PhD Thesis), Berkeley. Vahtera, V., Muona, J. & Lawrence, J. F. (2009): Phylogeny of the Thylacosterninae (Coleoptera, Elateridae). - Cladistics 25:147-160. Van Dyke, E. C. (1932): Miscellaneous studies in the Elateridae and related families of Coleoptera. - Proceedings of the California Academy ofSciences 20:291-465. Viviani, V. R. & Bechara, E. J. H. (1993): Biophysical and biochemical aspects of phengodid (railroadworm bioluminescence. - Photochemistry and Photobiology 58 (4): 615-622. - (1995): Bioluminescence of Brazilian fireflies (Coleoptera: Lampyridae): Spectral distribution and pH effect on luciferase-elicited colors. Comparison with elaterid and phengodid luciferases. - Photochemistry andPhotobiology 6 2 : 4 9 0 - 4 9 5 . - (1997): Bioluminescence and biological aspects of Brazilian railroad-worms (Coleoptera: Phengodidae). - Annals of the Entomological Society of America 90: 389-398. Viviani, V. R., Bechara, E. J. H. & Ohmyia, Y. (1999): Cloning, sequence analysis and expression of active Phrixothrix railroad-worms luciferases: relationship between bioluminescence spectra and primary structures. -Biochemistry 3 8 : 8 2 7 1 - 8 2 7 9 . Wells, S. A. (2003). A revision of the neotropical click beetle genus Semiotinus Pjatakowa (Coleoptera:

Plastoceridae Crowson, 1972

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Elateridae). - Proceedings of the Entomological Society of Washington 105 (3): 647-663. Zacharuk, R. Y. (1958): Structures and functions of the reproductive systems of the prairie grain wireworm, Ctenicera aeripennis destructor (Brown) (Coleoptera: Elateridae). - The Canadian Entomologist36: 725-751. - (1962): Some new larval characters for the classification of Elateridae (Coleoptera) into major groups. Proceedings of the Royal Entomological Society ofLondon (B) 31 (3/4): 29-32. Zacharuk, R. Y. & Albert, P. J. (1978): Ultrastructure and function of scolopophorus sensilla in the mandible of an elaterid larva (Coleoptera). - Canadian Journal ofZoology 56:246-259. Zacharuk, R. Y., Albert, P. J. & Bellamy, F. W. (1977): Ultrastructure and function of digitiform sensilla on the labial palp of a larval elaterid (Coleoptera). - Canadian Journal ofZoology 55:569-578.

4.8. Plastoceridae Crowson, 1972 Marc A. Branham Distribution. This family is composed of a single genus Plastocerus (= Ceroplastus) with two species. Plastocerus angulosus (Germar) is known from Turkey and Asia Minor, while P. thoracicus Fleutiaux is known from Southeast Asia (Lawrence 1991; Lawrence & Newton 1995). Biology and Ecology. Nothing is known of the biology or ecology of this family and the females and immature stages are unknown. Morphology, Adult Males (Figs. 4.8.1 A, B). Length 11-18 mm. Body elongate and more or less parallel-sided. Dorsal surfaces clothed with moderately long setae, especially on head and prothorax. Head prognathous, subquadrate, posteriorly biemarginate, without transverse occipital ridge or median endocarina. Frontoclypeal region strongly, abruptly declined. Eyes entire, moderately large and strongly protuberant; finely facetted, without interfacetal setae. Antennal insertions widely separated, slightly raised, concealed or barely exposed from above. Frontoclypeal suture absent. Strongly transverse labrum free, visible, and well sclerotized. Antennae 11-segmented, unipectinate, with flattened rami extending from distal portions of antennomeres 3-10; scape elongate, pedicel short. Short mandible broader at base, strongly curved apically, with unidentate apex; mola and prostheca absent; long setae present on dorsal surface of mandibular base. Maxilla with galea and lacinia short, lightly sclerotized and pubescent; palp 4-segmented, with apical palpomere somewhat elongate and flattened, Labium with highly reduced, membranous ligula;

Fig. 4.8.1. Plastocerus angulosus (Germar), male. A, dorsal, line = 2.5mm; B. P. angulosus, prothorax, ventral view. (Α, Β © Μ. Branham) palp 3-segmented, with apical palpomere slightly securiform. Gular sutures well separated; gula longer than wide. Corpotentorium absent. Cervical sclerites well-developed. Prothorax about 0.65-0.85 times as long as wide; widest at middle or posteriorly; sides sometimes slightly explanate; lateral carinae complete, simple, without raised margin; anterior angles not produced forward; posterior angles strongly acute;

104 posterior edge trisinuate. Promesothoracic interlocking m e c h a n i s m weakly developed. P r o s t e r n u m in f r o n t of coxae longer t h a n shortest diameter of a coxal cavity, slightly convex, w i t h o u t paired carinae; anteriorly truncate, w i t h o u t chin piece. Prosternal process complete, parallel-sided, slightly overlapp i n g mesoventrite, apically acute. Procoxae projecting well below p r o s t e r n u m , w i t h o u t concealed lateral extensions. Trochantins broadly exposed a n d n o t fused to n o t u m . Procoxal cavities transverse, very narrowly separated, externally broadly open, w i t h o u t n a r r o w lateral extensions; internally open. Scutellar shield well developed, n o t abruptly elevated, posteriorly truncate. Elytra 2 . 6 - 2 . 9 times as l o n g as combined w i d t h a n d 3 . 7 - 4 . 9 times as long as p r o n o t u m , w i t h o u t distinct p u n c t u r e rows or reticulate sculpturing; apices conjointly r o u n d e d ; epipleura very narrow, wider at base a n d apex. Mesoventrite separated by complete sutures f r o m mesanepisterna, which are well separated f r o m one another; anterior edge on same plane as metaventrite, w i t h o u t paired procoxal rests; mesoventral cavity small a n d shallow. Mesocoxae projecting. Trochantins exposed. Mesocoxal cavities contiguous, weakly defined, o p e n laterally (partly closed by mesepimeron, and mesanepisternum). Metaventrite slightly convex, w i t h l o n g discrimen and no transverse suture; exposed portion of metanepistern u m moderately elongate. Metacoxae contiguous, e x t e n d i n g laterally to meet elytra; plates n a r r o w b u t m o r e or less complete. Metendosternite w i t h lateral arms short, laminae absent, anterior process moderately l o n g a n d anterior t e n d o n s a p p r o x i m a t e . H i n d w i n g elongate w i t h very s h o r t apical field cont a i n i n g a single anterior oblique sclerite; radial cell elongate w i t h inner posterobasal angle m o r e or less right; cross-vein r3 well developed a n d very slightly oblique; basal portion of RP very long; medial field w i t h five free veins; MP 3 + 4 w i t h basal cross-vein and s p u r , j o i n e d by C u A j before forking; wedge cell well developed, apically truncate; anal lobe well developed b u t w i t h o u t distinct e m b a y m e n t . Legs long a n d simple; tibial spurs paired on all legs; tarsi 5-5-5, tarsomeres simple, w i t h o u t pubescent pads or m e m b r a n o u s lamellae; pretarsal claws simple, w i t h o u t setae near base. A b d o m e n w i t h seven ventrites (sternites III-IX), the first three of which are connate. Ventrite 1 not m u c h longer t h a n 2, w i t h o u t intercoxal process. Spiracles located in pleural m e m b r a n e , those on segm e n t VIII functional. Anterior edge of sternite VIII w i t h o u t m e d i a n strut. Anterior edge of sternite IX w i t h o u t spiculum gastrale. Tergite IX deeply emarginated. Tergite X well developed and free. Aedeagus trilobate, symmetrical; parameres individually articulated, w i t h slight o u t w a r d hooks; penis w i t h paired anterior struts. [Lawrence etal. 1999] P h y l o g e n y a n d T a x o n o m y . T h e family Plastoceridae was proposed by Crowson (1972) for P. angulosus (Germar) a n d was placed in Cantharoidea. Lawrence a n d N e w t o n (1995) transferred t h e family

Ladislav Bocak, Marc A. Branham and Robin Kundrata to Elateroidea along w i t h the other " c a n t h a r o i d " families. T h e Plastocerus concept used by Crowson (1972) a n d Lawrence & N e w t o n (1995) is based u p o n the original concept of Schaum (1852), a n d does n o t include t h e N o r t h American species once placed by LeConte (1861) in Plastocerus, w h i c h n o w b e l o n g to Octinodes (Elateridae: Cebrioninae). Also excluded f r o m the family are the diverse genera placed in h e r e by Schwarz (1907) a n d currently included in Eucnemidae: Phyllocerinae (Phyllocerus Serville a n d Cephalodendron Latreille) or in various subfamilies of Elateridae (Aphricus LeConte, Aplastus LeConte, Diplophoenicus Candeze, Dodecacius Schwarz, Enisonyx H o r n , Euplastius Schwarz a n d Euthysanius LeConte).

Literature Cited Crowson, R.A. (1972): A review of the classification of Cantharoidea (Coleoptera), with the definition of two new families, Cneoglossidae and Omethidae. -Revistade la Universidad de Madrid 21 (82): 35-77. Lawrence, J. F. (1991): Plastoceridae (Cantharoidea). P. 422 in Stehr, F. W. (ed.) Immature Insects. Volume 2. Kendall/Hunt, Dubuque, Iowa. Lawrence, J. F. & Newton, A. F. Jr. (1995): Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names). Pp. 855 in Pakaluk, J. & Slipiriski, S. A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warsaw. Lawrence, J. F., Hastings, A. M., Dallwitz, Μ. J., Paine, T. A. & Zürcher, Ε. J. (1999): Beetles of the World: A Key and Information System for Families and Subfamilies. CD-ROM, Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne. LeConte, J. L. (1861): Classification of the Coleoptera of North America. Prepared for the Smithsonian Institution. Part 1. - Smithsonian Miscellaneous Collections 3: i-xxv + 1 - 2 0 8 . Schaum, Η. R. (1852): Catalogus Coleopterorum Europae. Vierte Auflage. Entomologischer Verein in Stettin (ed.), Berlin, ν + 96 + 1 + 12pp. Schwarz, O. (1907): Genera Insectorum dirigesparp. Wytsman. Fascicule 50. Coleoptera. Fam. Plastoceridae. 10 pp., 1 pi. Pp. 225-370, pis. 1 - 6 . P. Wytsman, Brussels.

4.9. Family Drilidae Blanchard, 1845 Ladislav Bocak, Marc A. Branham and Robin Kundrata

D i s t r i b u t i o n . Drilidae is a small elateroid family w i t h approximately 100 species classified in only six genera: Drilus Olivier (25 spp.), Euanoma Reitter (8 spp.), Malacogaster Bassi (10 spp.), Paradrilus Kiesenwetter (1 sp.), Pseudeuanoma Pic (3 spp.), a n d

Family Drilidae Blanchard, 1845

Selasia Laporte (about 50 spp.) (Kundrata & Bocak 2007). Most genera and species occur in the Mediterranean region, usually with restricted ranges corresponding to Pleistocene refugia in the Iberian Peninsula (Paradrilus), or Greece, Turkey, and the Causassus (Euanoma and Pseudeuanoma). The genera Drilus and Malacogaster are widespread in the Mediterranean, b u t only two species of Drilus occur n o r t h of the Alps. Selasia is k n o w n f r o m the Afrotropical Region, Arab Peninsula, and Suqutra. [Bahillo de la Puebla & Lopez Colon 2005; Kundrata & Bocak 2007; Kleine 1942; Wittmer 1944.] Biology a n d Ecology. Drilidae occur in various habitats with populations of terrestrial mollusks u p o n which their larvae feed (Cros 1930; Gittenberger 1999; Orstan 1999; Mesher & Welter-Schultes 2001; Symondson 2004). There are no records of feeding by adult males (Crowson 1972). Drilidae are most common in lowland or lower m o u n t a i n forest habitats in central Europe and f r o m costal marshes to high altitude habitats in the Mediterranean. Adult males of Mediterranean species are diurnal, b u t they show m a x i m u m activity in the evening hours before sunset when there is h i g h h u m i d i t y and lower temperature. They f r e q u e n t the vegetation of herbal and lower s h r u b strata, where they can be collected by sweeping f r o m late March in low elevations in southern Europe and till the end of July in the mountains. Although they are capable of slow, reluctant flight, their vagility is generally very low and species ranges appear very restricted. All k n o w n females of the genera Drilus, Malacogaster and Selasia are wingless, with a larviform morphology of the thorax and abdomen and they can be found along with larvae in empty mollusk shells or foraging on the soil surface. The larvae of Drilus and Malacogaster are t h o u g h t to feed on one individual land snail d u r i n g each instar. Each instar is spent within the snail shell, feeding at first, then using the shell for shelter and ultimately p u p a t i n g within the shell. In early instars, multiple larvae can be found within a single snail shell (Balduf 1935). The larval biology of the genenParadrilus, Euanoma, and Pseudeuanoma is u n k n o w n , b u t similar life history is supposed also in these groups (Bahillo de la Puebla & Lopez Colon 2005; Kundrata & Bocak 2007). Drilidae are mostly k n o w n only f r o m males and due to their rareness they are poorly represented even in major European collections. The last record of the monotypic genus Paradrilus was at the end of the 19 t h century and the species may be extinct. The Afrotropical genus Selasia occurs commonly in forest and s h r u b habitats and their males are often attracted to light (Bourgeois 1882).

Morphology. Adult Males (Figs. 4.9.1-2). Moderately sclerotized beetles. Body form elongate, dorsoventrally flattened, parallel-sided to slightly widened posteriorly (Fig. 4.9.1 A). Length 2 . 4 10.8 m m . Coloration either dark brown to black (Pseudeuanoma, Paradrilus, most species of Drilus,

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some Selasia) or light brown to yellow (most species of Selasia), seldom with bright red body parts (Malacogaster, some Euanoma). Vestiture sparse, often long, moderately erect. P r o n o t u m and elytra sometimes shiny, with fine to rough, dense sculpture. Head hypognathous, small, only slightly retracted into prothorax. Transverse, width including eyes almost as wide as p r o n o t u m at frontal margin; prolonged anteriorly in some genera. Antennal insertions widely separated, anterolaterally oriented and barely visible from above; subantennal grooves shallow. Eyes (Fig. 4.9.1 B) lateral, hemispherically prominent, their frontal interocular distance 1.15-2.60 times eye diameter. Antennae stout, 11segmented, reaching one third of elytral length, narrowed to apex; filliform in Euanoma, Pseudeuanoma and Paradrilus, strongly serrate to flabellate in Drilus and Malacogaster, flabellate with very long lamellae in Selasia; scapus robust, pear-shaped, slightly asymmetrical; length of antennomeres 2 - 4 variable, antennomere 2 shorter than 3 in Drilus, Malacogaster and Selasia, subequal in Euanoma, Pseudeuanoma and Paradrilus (Figs. 4.9.1 F-J); following antennomeres gradually shortened, except terminal one; all antennomeres with moderately dense, erect pubescence. Frontoclypeal suture absent; anterior edge of frontoclypeus concave. Labrum membranous anteriorly, with long setae at apex. Mandibles long, slender, considerably curved, incisor margin simple or with teeth in middle (Fig. 4.9.1 L). Maxilla with fused galea and lacinia(Fig. 4.9.1K). Maxillary palpi small, 4-segmented; basal palpomere very short, transverse; palpomere 2 3.5 times longer than palpomere 1, widened to apex; palpomere 3 short, transverse; palpomere 4 almost as long as palpomere 2, parallelsided to gradually narrowed to apex, with obliquely truncate, membranous apex. Labium transverse, without ligula. Labial palpi small, 3-segmented; apical palpomere similar in shape to apical maxillary palpomere (Fig. 4.9.1 E). P r o n o t u m flat, 1.22-1.45 times wider than length at midline, widest in anterior third, seldom widest at base (Fig. 4.9.1 C); disc surface sometimes shiny, margins always finely to coarsely punctured; punctures at margins dense, gradually more sparse medially or roughly punctured across entire surface; shallow longitudinal depression along midline present in some species; entire disc with sparse, erected, long setae. Posterior angles sharp, p r o m i n e n t to rectangular; posterior edge bisinuate or widely rounded; lateral edges sinuate or convex; anterior edge straight or slightly concave. Prosternum long (e. g., Pseudeuanoma, Euanoma) or more transverse (Drilus, Selasia), without apparent prosternal process (Fig. 4.9.1 D). Scutellum small, simply rounded at apex. Mesoventrite with frontal margin straight (Euanoma), shallowly emarginate (Drilus) or deeply emarginate (Pseudeuanoma). Elytra flattened, almost parallel-sided to slightly widened posteriorly, widest in apical third, 2.20-2.90 times longer than width at humeri; shiny, with sparse to dense, fine, erected pubescence; longitudinal costae

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Fig. 4.9.1. A, Euanoma starcki Reitter, general appearance, line = 1 mm; B, Pseudeuanoma sp., head ventrally, line = 0.5 mm; C, Pseudeuanoma ionica Pic, pronotum, line = 0.5 mm; D, Pseudeuanoma sp., prothorax, ventrally, line = 0.5 mm; E, Drilus concolor Olivier, labium, line = 0.25 mm; F, Drilusflavescens Olivier, basal antennomeres, line = 0.5 mm; G, Selasiapulchra Pascoe, basal antennomeres, line = 0.5 mm; H, Malacogaster sp., basal antennomeres, line = 0.5 mm; I, Pseudeuanoma sp., basal antennomeres, line = 0.25 mm; J, Paradrilus opacus Kiesenwetter, basal antennomeres, line = 0.25 mm; K, Drilus concolor Olivier, maxilla, line = 0.5 mm; L, Drilus concolor Olivier, mandible, line = 0.25 mm.

absent, at most with very weak, irregular, longitudinal rows of punctures. Hind wings (Fig. 4.9.2 A) fully developed in all males (never brachypterous or apterous); apical field very short, bearing one or two anterior oblique sclerites and another posterior one; radial cell large with inner posterobasal angle slightly acute; cross-vein r3 very long and oblique; medial field with two or three free veins and no wedge cell; anal lobe well developed, without embayment. Legs slender, slightly flattened; coxae well separated; trochanters robust; femur attached apically; tibia without apical spurs; tarsi with five slender tarsomeres. Claws simple. Abdomen with seven ventrites. Terminal male segments as in Fig. 4 . 9 . 2 1 - K . Aedeagus of trilobate type, with fine sickle-shaped phallobase, regularly with processes at base; parameres fused basally; phallus strong, usually longer than parameres, widest at base, sometimes considerably curved (Figs. 4.9.2 B-G); internal sac membranous or slightly sclerotized, rod-shaped.

Morphology, Adult Females (Fig. 4.9.3 A-D).

Length 1 3 - 3 5 mm. Larviform, with elongate body (Drilus, Malacogaster) or robust (Selasia), slightly flattened dorsoventrally, parallel-sided (Drilus, Fig. 4.9.3 A) to considerably widened posteriorly (Selasia). Cuticle soft, weakly sclerotized (Drilus) or moderately sclerotized (Selasia). Vestiture usually dense, often long, erect. Coloration yellowish brown to dark brown. Head small, prognathous, slightly transverse, half the width of the prothorax. Small compound eyes lateral, flat, their frontal distance about 3 times eye diameter in lateral view. Antennal sockets inconspicuous, widely separated. Antennae robust, very short, as long as width of head, filliform, 10-segmented. Scapus robust, pear-shaped, slightly asymmetrical; antennomere 2 shorter; apical antennomere very small and narrow in Drilus (Fig. 4.9.3 B). Mandibles long, slender, distinctly curved towards apex; incisor margin with teeth. Both palpi small, with basal palpomeres very slender,

Family Drilidae Blanchard, 1845

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Fig. 4.9.2. A, Pseudeuanoma sp., wing, line = 2 mm; Β, Selasia sp., male genitalia, lateral view, line = 0.5 mm; C, D, Euanoma curvata Kundrata & Bocak, male genitalia, lateral and ventral view, line = 0.25 mm; E, Selasia sp., male genitalia, ventral view, line = 0.5 mm; F, Euanoma svihlai Kundrata & Bocak, male genitalia, ventral view, line = 0.25 mm; G, Drilus concolor Olivier, male genitalia, ventral view, line = 0.25 mm; H, Pseudeuanoma sp., hind leg, line = 0.5 mm; I, Euanoma starcki Reitter, apical abdominal segments, line = 0.25 mm; J, Pseudeuanoma reitteri Pic, apical abdominal segments, line = 0.25 mm; K, Selasia sp., apical abdominal segments, line = 0.5 m m .

subapical p a l p o m e r e transverse, a n d apical palpomere pointed. Thoracic s e g m e n t s larviform, slightly narrower t h a n those of a b d o m e n . Elytra a n d wings absent. Legs similar to those of males, b u t shorter, w i t h widely separated coxae; claws paired (Fig. 4.9.3 C). A b d o m e n w i t h n i n e visible segments. Widest in posterior third; apical s e g m e n t IX m u c h narrower a n d smaller. Ovipositor w i t h coxites plate-like, f u s e d w i t h paraproctal baculi (Figs. 4.9.3 D). Styli

small, articulate, connected to coxites by extensive m e m b r a n e . Vagina simple, sac-like, m e m b r a n o u s , w i t h u n p a i r e d extensive glands. [Kundrata & Bocak 2007; Cicero 1988; W i t t m e r 1989.] M o r p h o l o g y , Larvae (internal characters only verified for Drilus; Beutel 1995) (Figs. 4.9.4 A-C). Larvae elongate, s o m e t i m e s slightly twisted dorsally into a C-shape, slightly flattened. All b u t last instar w i t h a conspicuous vestiture of l o n g setae, especially on

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A

Fig. 4.9.3. Α-D, Drilusflavescens Olivier, female, A, general appearance, line = 5 mm; B, antenna, line = 0.5 mm; C, hind leg, line = 0.5 mm; D, female genitalia, line = 1 mm.

dorsal surface of abdomen; with heavily sclerotized and pigmented tergites and slightly less heavily sclerotized ventrites (Malacogaster and Drilus) (Fig. 4 . 9 . 4 C). Prognathous head sclerotized, short, partly retracted into prothorax. Epicranial sutures absent. Single stemma present on either side. Labrum fused to head capsule. Antennae 3-segmented; sensorium present on preapical segment; shorter than apical segment. Mandibles narrow and falcate, widely separated at base, slightly up-turned at tips; sucking channels present; mola and prostheca absent;. Ventral mouthparts form maxillolabial complex; exposed articulatory membrane absent. Cardo indistinct or absent; stipes subparallel, almost as long as head capsule; galea 1-segmented; maxillary palp 4-segmented. Mentum (or postmentum) elongated; labial palp 2-segmented. Sclerotized gula present but strongly shortened. Tentorium vestigial. Antennal muscles with cranial origin; tentoriocardinal and tentoriostipital muscles almost vertical; lateral tentoriohypopharyngeal muscle absent. Dense, hairy preoral filter present; closed prepharyngeal tube elongated. Cerebrum located in the postoccipital region.

Spiracles I-VIII biforous, positioned between terga and pleura at the ends of spiracular tubes. Tergite IX with a pair of stout, fixed urogomphi with numerous well developed setae and a slender pointed apex. Segment X forming sucker-like pygopod. Luminous organs absent. [Boving & Craighead 1931; Lawrence 1991; Beutel 1 9 9 5 (Drilus).]

Prothorax subparallel, slightly widening posteriorly, nearly as long as meso- and metathorax combined. Legs 5-segmented, pretarsus with 2 spatulate setae. Mesothorax with biforous spiracles at ends of spiracular tubes.

P h y l o g e n y a n d T a x o n o m y . Drilids were first described from western and southern Europe (Olivier 1790; Bassi 1834). Laporte (1836) described the Afrotropical genus Selasia, while other species were recognized from the eastern Mediterranean, northern Africa, and the Caucasus. Historically, only Drilus, Malacogaster and Selasia were originally assigned to Drilidae. The three remaining genera that are

Abdominal segments with well developed lateral tergal and pleural processes bearing long setae; lateral lobes more than twice as long as tergal lobes.

Last larval instar strongly twisted dorsally into a C-shape, narrowing abruptly from posterior to anterior end. Setae restricted to dorsal surface of abdominal segments I I - X (Malacogaster and Drilus); cuticle otherwise smooth, unsclerotized and light in color. Moderately sclerotized only in the dorsoposterior region of abdomen; abdominal segments bearing greatly reduced lateral tergal and pleural processes, both bearing short setae; lateral lobes more pronounced than tergal lobes (Fig. 4 . 9 . 4 A B). Mandibles widely separated, greatly reduced. Legs reduced and shortened, less sclerotized than in earlier instars and lacking claws. Tergite IX bearing two pairs of fixed, well sclerotized urogomphi; inner pair of setae twice as long as external pair. Segment X somewhat reduced. [Lawrence 1991; Balduf 1935; Crawshay 1903.]

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Family Drilidae Blanchard, 1845

were added to Drilidae in the early 20th century (e. g., Pic 1901). Recently the Afrotropical genus Selasia was studied by Wittmer (1989) and Geisthardt (2003) and the genera Euanoma and Pseudeuanoma were investigated by Kundrata & Bocak (2007). The family has been historically classified near Lampyridae, Omalisidae and other families of soft bodied beetles in the superfamily Cantharoidea (sensu Crowson, 1955). In his revision of the classification of Cantharoidea, Crowson (1972), restricted Drilidae to the Palaearctic genera Drilus and Malacogaster, and the Afrotropical Selasia, and he provisionally transferred Pseudeuanoma from Omalisidae to Drilidae, while also mentioning the possibility that several less-diverse genera which were not available for his study may belong to Drilidae, such as Euanoma, a genus close to Pseudeuanoma and Paradrilus, which was moved to Drilidae by Winkler (1925). The Oriental and east Palaearctic genera formerly classified in Drilidae (Olivier 1910; Wittmer 1944) were transferred to Lampyridae, Rhagophthalmidae, Omethidae, and Lycidae (Crowson 1972). The above listed genera were considered to compose Drilidae by Lawrence & Newton (1995). The monophyly of Drilidae in the present sense is weakly supported, without a single synapomorphy defining the family that is present in all genera (Kundrata & Bocak 2007). The position of Drilidae remains contentious. Beutel (1995) hypothesized their placement with Omalisidae, Lycidae and Lampyridae, while Branham & Wenzel (2003) hypothesized Drilidae to be a sister-group of all cantharoid families in their analysis. Molecular data (Bocakova et dl. 2007) indicate a close relationship between Omalisidae, Drilidae and Elateridae. Consequently, Drilidae are suggested to be only distantly related to the remaining soft-bodied elateroid families such as Lycidae, Lampyridae and Cantharidae.

Acknowledgements The Ministry of Education of the Czech Republic is acknowledged for the support of the studies on Drilidae (MSM6198959212). We thank Jennifer Zaspel for her assistance in producing the figure of drilid larvae.

C Fig. 4.9.4. A, Malacogaster sp. larvae, last instar larva, dorsolateral view, line = 5 mm; B, Malacogaster sp., last instar larva beside an earlier instar larva protruding from a broken snail shell, ventrolateral view; C, Drilus sp., larva, dorsal view, line = 5mm (A-C © M. Branham). included within the modern concept of the family were originally assigned to Omalisidae (Bourgeois 1882; Kleine 1933). The French entomologist Maurice Pic described almost all Palaearctic species that

Literature Balduf, W. V. (1935): The Bionomics of Entomophagous Cokoptera. 220 pp. St. Louis, John S. Swift. Bassi, C. A. (1833): Description du genre Malacogaster. In Guerin-Meneville, F. C. Magasin de Zoologie, d'Anatomie comparee et de Paleontologie, Insecta. Nr. 9 9 , 1 col. pi. Arthur Bertrand, Paris. Bahillo de la Puebla, P. & Lopez Colon, J. I. (2005): Los Drilidae Lacordaire, 1857 de la Peninsula Iberica e Isias Baleares (Coleoptera). - Boletin Sociedad Entomolögica Aragonesa 37:119-128. Beutel, R. G. (1995): Phylogenetic analysis of Elateriformia (Coleoptera: Polyphaga) based on larval

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characters. -Journal ofZoological Systematics and Evolutionary Research 33:145-171. Bocakova, M., Bocak, L., Hunt, T., Teraväinen, Μ. & Vogler, A. P. (2007): Molecular phylogenetics of Elateriformia (Coleoptera): evolution of bioluminescence and neoteny. - Cladistics 23:477-496. Bourgeois, J. (1882): Monographie des Lycides de l'ancien-monde.-Abeille 20:1-118. Boving, A. G. and Craighead, F. C. (1931): An illustrated synopsis of the principal larval forms of the order Coleoptera. - Entomologica Americana (N.S.) 11(1930): 1-351. Branham, M. A. & Wenzel, J. W. (2003): The origin of photic behavior and the evolution of sexual communication in fireflies (Coleoptera: Lampyridae) Cladistics 19:1-22. Crawshay, L. R. (1903): On the life history of Drilus flavescens Rossi. - Transactions of the Entomological Society ofLondon 51:39-51. Cicero, J. M. (1988): Ontophylogenetics of cantharoid larviforms (Coleoptera: Cantharoidea). -TheColeopterologists Bulletin 42 (2): 105-151. Cros, A. (1930) Malacogaster passerini: moeurs, evolution. - Bulletin de la Societe d'Histoire Naturelle d'AfriqueduNord 21:133-160. Crowson, R. A. (1955): The Natural Classification of the Families of Coleoptera. 187 pp. Nathaniel Lloyd, London. - (1972): A review of the classification of Cantharoidea (Coleoptera), with definition of two new families, Cneoglossidae and Omethidae. - Revista de la Universidad de Madrid 21 (82): 35-77. Geisthardt, Μ. (2003): Zwei neue Arten der Gattung Selasia Castelnau, 1836 aus Jemen. - Mitteilungen des Internationalen Entomologischen Vereins 28 (3/4): 99-109. Gittenberger, E. (1999) Predatory bore-holes in shells of terrestrial snails: Roth has priority. - Basteria 63: 164. Kleine, R. (1933): Coleopterorum Catalogus auspiciis et auxilio W. Junk editus S. Schenkling. Pars 128: Lycidae. 145 pp. W. Junk, Berlin. - (1942): Bestimmungtabellen der Lycidae. Bestimmungtabellen der europäischen Coleopteren. 123. Heft, 90 pp. Troppau. Kundrata, R. & Bocak, L. (2007): A revision of genera Euanoma and Pseudeuanoma (Coleoptera: Drilidae). -AnnalesZoologici57·. 427-441. Laporte, F. L. (1836): Etudes Entomologiques, ou descriptions d'insectes nouveaux et observations sur la synonymie. - Revue Entomologique 4:5-60. Lawrence, J. F. (1991): Drilidae (Cantharoidea). P. 424 in Stehr, F. W. (ed.) Immature Insects. Vol. 2. Kendall/ Hunt Publishing Co., Dubuque, Iowa. Lawrence, J. F. & Newton, A. F. Jr. (1995): Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names). Pp. 779-1006 in Pakaluk, J. & Slipiriski, S. A. (eds.) Biology, Phylogeny, and Classification ofColeoptera: papers Celebrating the 80th Birthday ofRoy A. Crowson. Muzeum i Instytut Zoologii PAN, Warsaw. Mesher, C. S. & Welter-Schultes, F. W. (2001): Morphological properties of Albinaria species on the island of Dia, north of Crete (Gastropoda: Clausiliidae). - Biologia Gallo Hellenica 26 (2): 87-110.

Ladislav Bocakand Milada Bocakova Olivier, G. A. (1790): Entomologie, ou Histoire Naturelle des Insectes, avec leurs caracteres generiques et specifiques, leur description, leur synonymie et leurfigure enluminee. Vol. 2. No 23, pp. 1-4, No 24, pp 1-4. Baudonin, Paris. Olivier, E. (1910): Coleopterorum Catalogus auspiciis et auxilio W. Junk editus S. Schenkling. Pars 10: Rhagophthalmidae, Drilidae. 10 pp. W. Junk, Berlin. Orstan, A. (1999): Drill holes in land snail shells from western Turkey. - Schriften zur Malakozoologie aus dem Haus der Natur Cismar 13:31-36. Pic, M. (1901): Synopsis des Euanoma Reitt. etPseudeuanoma Pic. - L'Echange, Revue Linneenne 202:74-75. Symondson, W. O. C. (2004): Coleoptera (Carabidae, Staphylinidae, Lampyridae, Drilidae and Silphidae) as predators of terrestrial gastropods. Pp. 37-84 in Barker, G.M. (ed.). Natural enemies of terrestrial molluscs, χ + 644 pp. CABI Publishing, Wallingford and Cambridge. Winkler, A. (1925): Drilidae. Catalogus Coleopterorum regionis palaearcticae. Pars 5, pp. 497-624. A. Winkler, Wien. Wittmer, W. (1944): Catalogue des Drilidae E. Oliv. (Coleoptera - Malacodermata). - Revista de la Sociedad Entomologica Argentina 12(3): 203-221. - (1989): Die Familie Drilidae (Coleoptera) in Südafrika, sowie Beschreibung von neuen Arten aus dem Südlichen Afrika. - Entomologica Basilensia 13: 187-205.

4.10. Family Omalisidae Lacordaire, 1857 Ladislav Bocak and Milada Bocakova

Distribution. Three genera are contained in Omalisidae: Omalisus Geoffroy (5 species), Thilmanus Gemminger (2 species) and Phaeopterus Costa, 1857 (1 species). Omalisus fontisbellaquei Geoffroy is widely distributed (western, southeastern and central Europe) and all other species are known from a small number of specimens from localities in southern Europe. Note that the generic name Omalisus Geoffroy was conserved by Opinion 1754 (ICZN, 1994). Biology and Ecology. Omalisidae live in forest or shrub habitats. Adult males can be collected by general sweeping. Larvae of O. fontisbellaquei prey on millipedes (Glomeris sp.). This species occurs in lower montane deciduous forests and is locally abundant. Males are capable of flight, but they usually sit motionless on leaves in the lowest forest stratum. Females are wingless (Bertkau 1891) and remain in the surface soil and few specimens have been collected. Larval luminosity was reported incorrectly by Bertkau (1891) for O. fontisbellaquei (see also Beutel 1995) and refuted by Burakowski

Family Omalisidae Lacordaire, 1857

(1988), who collected larvae of this species in high numbers. Other species of Omalisidae have been collected only in the adult stage and their biology is unknown. Most species are extremely rare and even the largest collections house few specimens collected after the 19th century. Although Thilmanus obscurus Baudi is usually the second best represented species in most collections, we have not found any collected in the 20th century even from areas of intensive collecting activity. Morphology, Adult Males (Figs. 4.10.1-2). Length 3 - 9 mm. Moderately sclerotized beetles. In

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Thilmanus cuticle on elytra and abdomen softer, flexible. Body form elongate, dorsoventrally flattened, parallel-sided. Some species concolorous and dark (Phaeopterus unicolor Costa, Thilmanus spp.), some with brightly colored elytra (several south European species with reddish brown elytra) or with light brown lateral elytral margins (O. fontisbellaquaef). Vestiture sparse, long, moderately erect. At least some parts of the body shiny in most species; otherwise cuticle with rough sculpture. Head small, much narrower than prothorax, prolonged anteriorly, exposed from above; prognathous, with fully developed mouthparts. Antennal

Fig. 4.10.1. Omalisus fontisbellaquaei Fourcroy adult. A, general appearance (artist: © M. Brlik), line = 1 mm; B, cranium, dorsally, line = 0.1 mm; C, labium, line = 0.1 mm; D, maxilla, line = 0.1 mm; E, labium, line = 0.1 mm; F, mandible, line = 0.1 mm; G, pronotum, line = 0.5 mm; H, prothorax, ventrally, line = 0.5 mm; I, wing, line = 1 mm; J, hind leg, line = 0.5 mm. (B-J © L. Bocak)

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Fig. 4.10.2. Omalisus fontisbellaquaei Fourcroy (A-E) and Thilmanus obscurus Baudi (F) adult features. A, scutellar plate, line = 0.1 mm; B, meso- and metaventrite, line = 0.5 mm; C, metanotum, line = 0.1 mm; D, male genitalia, line 0.1 mm; E, male terminal abdominal segments, line 0.1 mm. F, male genitalia, line 0.1 mm. (A-F © L. Bocak)

sockets large, with long, deep groove between them; antennal insertions widely separated; located on lateral frontal region. Eyes inserted laterally, strongly hemispherical and prominent (Omalisus) or somewhat flattened (Thilmanus); with coarse pattern of ommatidial corneae; eye diameter distinctly less than the distance between them. Frontoclypeal suture absent; anterior edge of frontoclypeus concave. Labrum membranous apically (Fig. 4.10.1 E), with long densely arranged setae at anterior margin. Antennae 11-segmented, cylindrical in cross section; scapus robust, pear shaped, slightly asymmetrical; pedicellus and antennomere 3 subequal, shorter than antennomere 4; following antennomeres gradually shortening except terminal one; all antennomeres with moderately dense, erect pubescence. Mandibles long, robust, distinctly curved; mesal edge simple, without teeth (Fig. 4.10.1 F). Maxilla with galea and lacinia fused (Fig. 4.10.1 D). Maxillary palpi 4-segmented; apical palpomere parallelsided, obliquely truncate at apex, with short dense pubescence. Labium small, slender, without ligula; labial palpi minute, 3-segmented (Fig. 4.10.1 C). P r o n o t u m apparently narrower than elytra, flat, with rough sculpture; median part shiny in

Omalisus (Fig. 4.10.1 G), entire p r o n o t u m uniformly sculptured in Thilmanus. Anterior edge almost straight; anterior angles distinct, slightly rounded, laterally bisinuate; posterior angles projecting, acute; posterior edge simple, straight. Scutellar shield triangular to parallel-sided, simply rounded at apex. Prosternum robust, with very slender prosternal process (Fig. 4.10.1 H). Mesoventrite transverse. Metaventrite with strong keelshaped anterior process. Metacoxae small, inserted in deep excavations laterally. Metendosternite with simple, robust stalk; arms absent. Elytra parallel-sided, flattened. Each elytron with nine longitudinal costae; rounded, partly irregular cells in simple rows present between costae {Omalisus); cells inconspicuous in Thilmanus; costa 6 m u c h stronger than others, elevated, forming humeral edge. Legs slender, moderately flattened, slightly sclerotized, densely pubescent. Coxae well separated, slightly elongate; trochanters slender, obliquely attached to femora; femora slender, long; tarsi with five slender tarsomeres (Fig. 4.10.1 J). Claws simple. Abdomen with eight free ventrites, terminal male segments as in Fig. 4.10.2 E. Aedeagus of trilobate type, with separate articulated parameres

Family Omalisidae Lacordaire, 1857

(Figs. 4.10.2 D, F); internal sac inconspicuous, membranous; in a few species with slender sclerotized rod. Females. Body less sclerotized. Mouthparts similar to those of adult males. Antennae and legs shortened, but with normal number of segments. Elytra vestigial and hind wings absent. Meso- and metathorax morphologically similar. Abdomen physogastrous. Morphology, Larvae (Fig. 4.10.3). Length 10-13 mm. Body slender, about 2 mm wide. Tergites dark brown, well sclerotized, with rough surface. Sternites pale, weakly sclerotized. Head wider than long (Fig. 4.10.3 A); anterior edge prolonged, forming a slender lobe which is sclerotized basally, almost membranous distally, and slightly emarginate at its apex. Mandibles slightly longer than clypeolabrum, slender, moderately curved at apex. Maxillolabial complex with well developed maxilla; stipes robust, elongate; galea small, pointed anteriorly; lacinia elongate; palpi with four palpomeres. Labium elongate, with well developed, transverse submentum, elongate mentum and subtrapezoidal prementum; palpi with two palpomeres. Antennae slender, 3-segmented (Fig. 4.10.3 C); antennomeres 1 and 2 cylindrical, robust; apical segment minute. Pronotum almost rectangular, with both anterior and posterior angles rounded. Remaining tergites except terminal one similar in shape, with rounded, finely dentate lateral margins. Thoracic and abdominal terga I-VIII with longitudinal keel and oval, shiny elevations on each side. Tergum IX rounded laterally, without keel. Tergum X elongate and conical, with retractile, membranous, multilobed grooming organ. Legs relatively long and densely setose. (Bertkau 1891; Burakowski 1988.) Phylogeny and Taxonomy. Omalisidae were originally included in Lycidae as a subfamily, and until recently, were considered to be their closest

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relatives. Molecular data showed that they do not belong to the Lycidae + Lampyridae + Cantharidae clade and preliminary results suggest a close relationship with Elateridae (Bocakova et al. 2007). The possibility of such a relationship was discussed by Bourgeois (1882), who considered Omalisidae to be a part of Lycidae as a tribe "Lycides" in the family "Malacodermes." They were subdivided in two subtribes "Homalisides" and "Lycides vrais". Bourgeois (1882) included three genera in Omalisidae: Omalisus, Thilmanus and Paradrilus Kiesen wetter, the last of which is now classified in Drilidae. Almost all species of Omalisidae were described during the 18th and 19th century, the only species described more recently being Omalisus nicaeensis Lesne, 1921. Kleine (1933) followed the traditional view on the position of Omalisidae and listed them as a subfamily Homalisinae within Lycidae in his world catalogue. He listed four genera in the group, Homalisus Illiger (including the subgenus Phaeopterus Costa), Thilmanus, Euanoma Reitter, and Pseudeuanoma Pic. Later (Kleine 1942) produced a key for their identification. Omalisidae were recognized as a family for the first time by Crowson (1955). Later, Crowson (1972) tentatively transferred Thilmanus to Lycidae and Euanoma and Pseudeuanoma to Drilidae. Thilmanus was returned to Omalisidae by Bocak & Bocakova (1990) but it was placed in Drilidae by Medvedev & Kazantsev (1992). Lawrence & Newton (1995) listed the genera Omalisus and Thilmanus, which represent eight species after critical revision of their status (Bocak and Brlik 2008). The position of Thilmanus has been unclear and Crowson (1972) excluded it from the family based on the shape of the prosternal process and the absence of puncture rows on the elytra. Bocak & Bocakova (1990) returned Thilmanus to Omalisidae on the basis of the male genital morphology. Kazantsev (2005) described the subfamily Thilmaninae and placed it in Lycidae close to those groups having neotenous larvae. In spite of this decision, Kazantsev (2005) listed several characters present in Thilmanus but missing in lycids (i. e., seven abdominal segments and conspicuous epipleura).

Fig. 4.10.3. Omalisus fontisbellaquaei Fourcroy, larva. A, head, dorsal; B, general appearance, lateral; C, antenna. (A-C, from Burakowski 1988; © Polish Entomological Society)

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His classification and analysis are strongly affected by the presence of neoteny in several clades and it is in deep conflict with molecular data (Bocak et al. 2008). Neoteny and the lack of larval characters were discussed by Crowson (1955). Moreover, some characters may be modified due to the small, weakly sclerotized body (e. g., wing venation, rows of punctures on elytra) and unfortunately, all Thilmanus specimens known in collections were collected before 1900 and are not useful for DNA studies. As the problem cannot be solved at the moment, most authors retain the traditional position of Thilmanus in Omalisidae (Bocak and Brlik 2008).

Acknowledgements Institutions cited in the captions are gratefully acknowledged for allowing us to use copyrighted figures. The Grant Agency of the Czech Republic is acknowledged for the support of our studies on Omalisidae (206/06/1392). Finishing work was supported from the grant of the Ministry of Education of the Czech Republic MSM6198959212.

Literature Bertkau, P. (1891): Beschreibung der Larve and des Weibchen von Homalisus suturalis. - Deutsche Entomologische Zeitschrift 1891: 37-42. Beutel, R. G. (1995): Phylogenetic analysis of Elateriformia (Coleoptera: Polyphaga) based on larval characters. -Journal ofZoological Systematics and Evolutionary Research 33:145-171. Bocak, L. & Bocakova, M. (1990): Revision of the supergeneric classification of the family Lycidae (Coleoptera). -PolskiePismoEntomologiczne 59:623-676. Bocak, L. & Brlik, M. (2008): Revision of the family Omalisidae (Coleoptera, Elateroidea). - Insect Systematics and Evolution 46:189-212. Bocak, L., Bocakova, M., Hunt, T. & Vogler, A. P. (2008): Multiple ancient origins of neoteny in Lycidae (Coleoptera): consequences for ecology and macroevolution. - Proceedings of the Royal Society B, 275: 2015-2023. Bocakova, M., Bocak, L., Hunt, T., Teravainen, M. & Vogler, A. P. (2007): Molecular phylogenetics of Elateriformia (Coleoptera) and the evolution of bioluminiscence and neoteny. - Cladistics 23:477-496. Bourgeois, J. (1882): Monographie des Lycides de l'ancien-monde. -A&aY/e 20:1-118. Burakowski, B. (1988): Observations on the larval morphology and biology of Omalisus fontibellaquei Fourcroy (Coleoptera, Homalisidae). - Polskie Pismo Entomologiczne 58: 571-574. Crowson, R. A. (1955): The Natural Classification of the Families of Coleoptera. 187 pp. Nathaniel Lloyd, London. Crowson, R. A. (1972): A review of the classification of Cantharoidea (Coleoptera), with definition of two new families, Cneoglossidae and Omethidae. - Revista de la Universidad de Madrid 21(82): 35-77.

Ladislav Bocakand Milada Bocakova Kazantsev, S. V. (2005): Morphology of Lycidae with some considerations on evolution of Coleoptera. -Elytron 19:49-226. Kleine, R. (1933): Coleopterorum Catalogus auspiciis et auxilio W. Junk editus S. Schenkling. Pars 128: Lycidae. 145 pp. W. Junk, Berlin. Kleine, S. V. (1942): Bestimmungstabellen der Lycidae. Bestimmungstabellen der europäischen Coleopteren. 123. Heft, 90 pp. Edmund Reitter's Nachf. Emmerich Reitter, Troppau. Lacordaire, J. T. (1857): Histoire Naturelle des Insectes. Genera des Coleopteres ou expose methodique at critique de tout les genres proposes jusqu'ici dans cet ordre des insects. Tome quatrieme contenant les families des buprestides, troscides, eucnemides, elaterides, cebrionides, cerophytides, rhipicerides, dascyllides, malacodermes, clerides, lymexylones, cupesides, ptiniores, bostrichides et cissides. Librairie Encyclopedique de Roret, Paris. 579 pp. ICZN (1994): Opinion 1754. Histoire abregee des insectes qui se trouvent aux environs de Paris (Geoffroy, 1762): some generic names conserved (Crustacea, Insecta). - Bulletin of Zoological Nomenclature 51(1): 58-69. Lawrence, J. F. & Newton, A. F. Jr. (1995): Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names). Pp. 779-1006 in Pakaluk, J. & Slipiriski, S. A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: papers Celebrating the 80th Birthday ofRoy A. Crowson. Muzeum i Instytut Zoologii PAN, Warszawa. Lesne, P. (1921): Un Lycide nouveau de la faune frangaise. - Bulletin de la Societe Entomologique de France 1921:180-182. Medvedev, L. N. & Kazantsev, S. V. (1992): A new subfamily and a new genus oflycid beetles (Coleoptera, Lycidae) from Southeast Asia. Pp. 55-60 in Medvedev L. N. (Ed.) Insect Systematics and Ecology of Vietnam. Nauka Publishing House, Moscow.

4.11. Lycidae Laporte, 1836 Ladislav Bocakand Milada Bocakova Distribution. Lycidae occur throughout the World on all continents except Antarctica and New Zealand (which has an introduced Australian species) with the highest diversity and abundance in the humid tropics. There are about 4600 described species classified in about 160 genera. Only the Palaearctic and Nearctic lycid faunas are relatively well known. The fauna of the western part of the Palaearctic region comprises about 20 species, most of them from the genera Dictyoptera Latreille, Benibotarus Köno, Pyropterus Mulsant, Platycis Thomson, Lopheros LeConte, and Lygistopterus Mulsant. Some genera occur in Northern Africa and in the Middle East (Lycostomus Motschoulsky and Adoceta Bourgeois). The Japanese fauna of Lycidae is well known and there are about 100 species from 20 genera. In addition to genera widely distributed in the Holarctic region, Metriorrhynchini are represented

Lycidae Laporte, 1836

by Cautires e and Xylobanus Waterhouse, and Platerodini by Plateros Bourgeois. The Nearctic fauna resembles that of Japan in generic structure, with Metriorrhynchini absent and Calopterini widely distributed (Calopteron Laporte, Leptoceletes Green, Caenia Newman). The large Metriorrhynchini clade has a Gondwanan distribution and with about 1400 described species it is the most diverse group of Lycidae. They have the center of generic diversity in the northernmost part of the Australian region and the h u m i d parts of the Oriental region. Platerodini are the second largest and widespread tribe; they are absent only from Europe and Australia except northernmost Queensland. There are two branches within the lycine complex: the Lycini are mostly Afrotropical with a very diverse genus Lycus Fabricius in sub-Saharan Africa and the closely related Lycostomus in Asia, northern Africa and North America; the largely Neotropical Calopterini are represented by 24 genera and about 500 species. A few species of the group are known from the Nearctic Region. Calochromini are the only group of Lycidae with a cosmopolitan distribution, although they are not common. The largest genera are Calochromus Guerin-Meneville, the closely related Lygistopterus, and Adoceta. Erotini and Dictyopterinae are Holarctic, and poor in numbers of species. The distribution of taxa with neotenous females is generally limited. Leptolycini are known from the West Indies and South America, Dexorinae from h u m i d areas of the Afrotropical region (e. g., Cameroon, Kenya, Tanzania, Sierra Leone), Ateliinae and Lyropaeinae from the Oriental region. A characteristic feature of lycid distribution is a high degree of endemism at the species, genus, and tribe levels. At the species level the typical range is limited to a mountain range or a single island while most genera occur in a part of a single geographic region, few of them in two regions. Only Plateros, Lycus, and Calochromus are widespread. [Bocak 2002; Bocak & Bocakova 1987,1988,1990, 2008; Bocakova 2001, 2003; Green 1949,1950,1951,1952,1953; Kleine, 1933,1942; Nakane 1969.]

Biology and Ecology. Lycidae are terrestrial beetles with a strong preference for forest or shrub habitats. They are not present in deserts or above upper tree limit in high mountains. Adults of most species avoid sunny places and remain under canopy, where they sit on leaves or decaying wood. Many species are usually flying early in the morning and shortly before sun set. The slow and cumbersome flight is characteristic for all lycids. Few genera visit flowers, b u t some species may aggregate in high numbers in flowering trees. This is characteristic for Lycus and Lycostomus in Africa and Asia, respectively, Calopteron and Thonalmus Bourgeois, 1883 in the Americas, Calochromus worldwide and Leptotrichalus Kleine, 1925 in South East Asia. Although many species are nocturnal, they are attracted to light only in moderate n u m bers. Flight-intercept traps yield high numbers of

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species in shaded places under the canopy. Most adult lycids are probably short lived and do not feed in the adult stage, though nectar may be consumed by some. Adults of lycids are capable of reflex bleeding. Hemolymph appears on elytral margins, antennae, and on femorotibial joints after disturbance. It is malodorous and protects lycids from predators. Although birds are the main potential predators mentioned in the literature (Moore & Brown 1981), recent observations indicate that spiders are also important (Eisner etal. 2008). Aposematic coloration occurs frequently. Lycids are commonly black and red or black and yellow, sometimes with complicated reticulate color patterns formed by brightly coloured elytral costae and dark bottoms of cells or with tricolored elytra. Color patterns may be geographically limited with one or a few patterns dominant locally. Lycids may form aggregations of u p to 15 species and non-lycids are sometimes involved in these mimicry rings (i. e., Coleoptera, Lepidoptera, Heteroptera). Most lycid species are winged in both sexes b u t several genera have neotenous larviform females. Female neoteny was proved by observing copulating males and females of Duliticola Mjöberg in South East Asian rain forests (Mjöberg 1925; Wong 1996) and Lyropaeus Waterhouse (Gravely 1915). Males and females of some Platerodrilus Pic and Macrolibnetis Pic species were identified by molecular markers (Levkanicova and Bocak, 2009). Miller (1991) described a larviform female of Leptolycus heterocornis Leng & Mutchler from the West Indies. We can expect this phenomenon in other genera elsewhere when male specimens are present in museum collections in high numbers and females are unknown. This pattern exists for Dexoris Waterhouse (Dexorinae) in the Afrotropical region, Melius Waterhouse, and Scarelus Waterhouse (Ateliinae), Pendola Bocak, Skrivania Bocakova & Bocak, Microlyropaeus Pic, and Antennolycus Bocakova & Bocak (Lyropaeinae) in the Oriental region (Bocakova 2006). All known Duliticola and Platerodrilus females remain larviform and do not pupate before sexual maturation. The only morphological change is the opening of the sexual ducts after shedding the last exuviae (Wong 1996). Females of Duliticola and Platerodrilus are up to 15 times longer than males. Lycid larvae occur in dead wood in various stages of decomposition, in forest litter and soil with high content of organic material. They often remain on the surface of moist wood or under loose bark. Inside rotten trunks or branches they do not form tunnels b u t they pass through soft decayed parts or through crevices. Lycid larvae often live in wood in close contact with the soil. Moist, shaded microhabitats that promote microbial growth are typical for Lycidae and only members of a few genera are able to colonize tree trunks in sunny places (Lycostomus spp., Calochromus spp.). Lycid larvae are commonly found aggregated, especially surface active species that are often aposematically colored (Lycostomus, Calopteron, Calochromus, Macrolibnetis). Larvae of lycids were suggested

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Fig. 4.11.1. Adults (line = 0.5 mm unless otherwise noted). A, Lycostomus kraatzi Bourgeois, habitus; B, Platycisschneidert Kiesenwetter, head, dorsal view; C, platerosplanatus Waterhouse, head capsule, ventral view; D, Prometanoeus ochraceus Kleine, maxilla, line = 0.1 mm; E, Metriorrhynchus parallelus (Guerin Meneville), labium, line = 0.1 mm; F, M. parallelus, mandible, line = 0.1 mm; G, platerosplanatus Waterhouse, labrum and hypopharynx, line = 0.25 mm; H, Cladophorus elegantulus Kleine, antennomeres 1 - 7 , line = 0.1 mm; I, Platycis minutus (Fabricius), antennomeres 1 - 5 ; J, Dictyoptera aurora (Herbst), pronotum; K, Platycis cosnardi (Chevrolat), pronotum; L, Microlycus minutus Pic, pronotum; M, Libnetispiceovittatus Pic, prothorax, ventral view; N, Calolycus calanticatus Gorham, wing; O, Plateros planatus Waterhouse, metathorax, dorsal view; P, Libnetis piceovittatus Pic, meso- and metathorax, ventral view. (Α, Β, I, J, Κ © Richard Jass, D, E, F, Η from Bocak 2002, © Institute of Entomology, CAS; C, G, L, M - P from Bocakova 2001, © Institute of Entomology, CAS).

to prey on larvae of other beetles, molluscs or on larvae of Diptera (Perris 1846; Shelford 1916). Miller (1988) observed foraging on small molluscs in North America. Many authors reported feeding on fermenting juices and this is now considered to be the principal diet (see Bocak δε Matsuda 2003). The exact number of larval instars and the length of larval development are unknown. So far, no lycid has been reared from the egg to the last larval instar

but the authors' observations show that lycid larvae require several years to become fully-grown. [Bocak & Bocakova 2 0 0 8 ; Bocak & Matsuda 2 0 0 3 ; Burakowski 1988; Hayashi 1 9 8 6 ; Miller 1988, 1 9 9 7 ; Wong 1996.]

Morphology, Adults. Length 2-28 mm. Soft bodied beetles with cuticle lightly sclerotized and flexible, especially on elytra and abdomen. Body form

Lycidae Laporte, 1836

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Fig. 4.11.2. Adults (line = 0.5 mm). A, Benibotarus taygetanus (Pic), male genitalia; B, Cautires sinensis (Pic), male genitalia; C, Dihammatus cribripennis Waterhouse, male genitalia; D, Cavoplateros dentatus Kleine, male genitalia; E, Bulenides obsoletus Waterhouse, female genitalia; F, Platerosparticularis (Pic), female genitalia; G, Libnetispiceovittatus Pic, female genitalia; H, Teroplasfusculus Gorham, terminal abdominal segments of male; I, Dihammatus sp., female terminal sternum; J, Cautires sinensis (Pic), elytral reticulation; K, Malacolycus paululus Kleine, elytral reticulation. (A © Richard Jass, Β, E, J, Κ from Bocak 2002, © Institute of Entomology, CAS; C, D, F, G-I from Bocakova 2001, © Institute of Entomology, CAS). elongate, dorsoventrally flattened, more or less parallel-sided (Fig. 4.11.1 A), with head partially covered by p r o n o t u m . Some species concolorous and dark, b u t many brightly colored or aposematically patterned in yellow and black or red and black. Vestiture dense, short, and suberect in most species; few species with shiny body parts (Calochromus, Metriorrhynchus Gemminger & Harold, Trichalus Waterhouse). Head small (Figs. 4.11.1 A-C), m u c h narrower than prothorax. Prognathous in most species with fully developed mouthparts; head f o r m i n g a long

rostrum in floricolous groups (Lygistopterus, Lycostomus; see Fig. 4.11.1 A, Leptotrichalus, Porrostoma Laporte); m o u t h parts reduced to m i n u t e vestiges in several genera (e. g., Dexoris, Lyropaeus). Eyes lateral, hemispherically prominent, with fine facets; eye diameter usually about their frontal distance; males of some nocturnal species with much larger eyes (some Metriorrhynchini and Platerodini). Frontoclypeal suture absent; anterior edge of frontoclypeus concave. Labrum rounded, small (Fig. 4.11.1 G). Antennal insertions widely separated. Antennae almost always with eleven antennomeres

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(ten in Coloberos Bourgeois and Lyropaeus). Scapus robust, pear shaped, slightly asymmetrical; pedicellus small, often transverse (Figs. 4.11.1 I, H); following antennomeres slightly serrate to flabellate (males of some species). All antennomeres densely pubescent. Mandibles long, robust, seld o m slender (Duliticola), or short, reduced (floricolous species); incisor edge simple, without any teeth (Fig. 4.11.1 F). Maxilla with small triangular cardo, stipes simple, parallel-sided to triangular, with fused galea and lacinia. Maxillary palpi foursegmented;apicalpalpomereusuallyparallel-sided, obliquely truncate at apex (acute in lyropaeines, widely dilated in some Calochromus). Labium small, p r e m e n t u m variable in shape, mostly slender, seld o m longitudinaly divided in two parts, m e n t u m tiny, transverse, without ligula; labial palpi threesegmented (Fig. 4.11.1 E), seldom reduced. Pronotum slightly narrower than elytra, flat, sometimes with elevated lateral edges (Lycus, Lycostomus). Anterior edge usually projecting forward; anterior angles obtuse, lateral edges simple, usually concave; seldom with protuberances on posterior third (Metriorrhynchini: Trichalina). Posterior edgesprojecting,moreorless acute; posteriormargin bilobed (Figs. 4.11.1 J-K). Prosternum transverse (Fig. 4.11.1M), Scutellar shield triangular to parallelsided, usually emarginate at apex. Mesoventrite emarginate anteriorly (Fig. 4.11.1 P). Metaventrite broad, with short discrimen (Fig. 4.11.1P). Metendosternite with simple, robust stalk; arms absent. Elytra subparallel, distinctly widened posteriorly (Lycostomus), flattened and widely rounded (Lycus, especially subgenus Chlamydolycus Bourgeois), or globular (Broxylus Waterhouse); usually with longitudinal and often with transverse costae (Figs. 4.11.2 J-K) in the following patterns: (1) nine longitudinal costae, four of them stronger, called primary costae and five weaker secondary costae with regular transverse costae (e. g., Dictyopterinae, Lycinae: Erotini, some Metriorrhynchini and Platerodini); (2) secondary costae absent and only four primary costae and transverse costae present (e. g., some Metriorrhynchini), intercostal spaces with irregular veins (Lycini), or with irregular punctures (Lyropaeus, Lil·netis Waterhouse, Macrolycus Waterhouse, Dilophotes Waterhouse); (3) longitudinal costae weak, often incomplete, intercostal spaces without any structure, glabrous (Calochromini); (4) elytra without any costae (Dexoris, Antennolycus). Hind wings present in all taxa with developed elytra, never shortened or vestigial. Apical field very short, wedge cell absent, AA4 inconspicuous or absent (Fig. 4.11.1 N); wing venation reduced in small species. Legs slender, flattened, weakly sclerotized, flexible, densely pubescent. Coxae separate, globular to slightly elongate, reaching lateral margin of metathorax; trochanters slender, obliquely attached to femora; femora slender, long, except males of few species; tarsi with five tarsomeres; tarsomeres 2 - 4 often with membranous pads. Claws simple in most species, bifid at apex in Macrolycus and Dilophotes.

Ladislav Bocakand Milada Bocakova

Abdomen with eight (males) or seven (females) visible sternites. Sclerites soft, connected by membranes. Abdomen usually shorter and much narrower than elytra. Male terminal abdominal segments as in Fig. 4.11.2 H; spiculum gastrale absent. Last visible female sternite with very long spiculum ventrale in Calochromus and some Dictyopterinae, reduced or absent in other groups (Fig. 4.11.2 I). Aedeagus (Fig. 4.11.2 A) of the trilobate type with articulated parameres, which are absent in some groups (Platerodini, Metriorrhynchini, Figs. 4.11.2 B, D), considerably reduced in others (Lycini). Internal sac inconspicuous, membranous in most species; often with thorns in Metriorrhynchini. Ovipositor with coxites platelike, separate, freely attached to rod-like paraproctal baculi (Figs. 4.11.2 E-G); paraprocts sometimes fused. Styli short, freely movable, connected to coxites by extensive membrane. Vagina simple, sack-like, membranous, with two lateral accessory glands attached distally; seldom vagina strongly sclerotized (some Metriorrhynchus). Spermathecal duct short to very long (Dilophotes); usually spirally curved. Spermatheca simple, moderately sclerotized, globular, apically bearing y-shaped gland. [Bocak & Bocakova 1990, 2008; Kazantsev 2005; Lawrence etal. 1999 b.] Morphology, Larvae. Lycid larvae may be separated in two homogenous groups according to their general form. Female neotenous larvae of genera Duliticola, Lyropaeus, and Platerodrilus represent the first group. The general appearance of these "trilobite larvae" is shown in Figs. 4.11.3 D, Ε (for detailed description see Bocak & Matsuda 2003, Levkanicova & Bocak 2009). The male i m m a t u r e stages are u n k n o w n , as are those of the neotenous groups, Ateliinae and Dexorinae. The second group includes 98% of the described species of Lycidae. Both, males and females of t h e m undergo pupation and the adults are winged. The larvae are characterized as follows; body campodeiform, cylindrical to strongly flattened, usually subparallel; middle part of abdomen sometimes distinctly wider (Lycostomus, Porrostoma); larvae usually moderately sclerotized; sclerites variable in size (Figs. 4.11.3 A-F). Head prognathous, small, partly retracted into p r o n o t u m ; flattened dorsoventrally, sometimes transverse (Figs. 4.11.4 A-C), rarely elongate (Lycostomus), with slightly marked nasale, or simply rounded at anterior head margin. Stemmata absent, or one small or large stemma present on each side (Figs. 4.11.4 D-F). Frons, clypeus and labrum fused; sutures absent. Sides of head capsule slightly less sclerotized than dorsal side or membranous (Figs. 4.11.4 D-F). Antennae 2-segmented, with large basal articulatory membrane; often deeply retracted (Fig. 4.11.4 C); antennomere 1 strongly transverse wider than long, ring-like, sometimes with three small processes on posterior margin; antennomere 2 cylindrical, with membranous apex; sclerotized part with several digitiform

Lycidae Laporte, 1836

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Fig. 4.11.3. Larvae, anterior part of body (line = 1 mm). Α-B, Lycostomusferrugineus (Fabricius), dorsal and ventral view; C, Plateros sp., dorsal view; D, E, Duliticola sp., dorsal and ventral view; F, Cautiresyuasai Nakane, dorsal view. (A-F from Bocak & Matsuda 2003, © Taylor and Francis Inc., Basingtoke).

pegs or one dorsal peg (Figs. 4.11.4 Α-B). Mandibles falciform, strongly curved to almost straight, without mola; bases approximate medially; apical parts diverging, entire mandibles cleft longitudinally, consisting of three parts (designated as premandible, labral lobe and stiletto by Kazantsev [2005]); inner parts fits into groove of outer part, with entire incisor margin. Ventral m o u t h p a r t s retracted, forming maxillolabial complex, without lateral movability of maxilla. Maxillae with fused galea and lacinia f o r m i n g slender, falciform mala, which may be shortened or vestigial. Mala and palpifer partly fused, separated f r o m ventral plate by m e m b r a n o u s area; palpi 3-segmented, mostly slender, elongate, attached to segment-like, elongate palpifer. Posterior part of labium formed by undivided p o s t m e n t u m ; entirely fused with stipites; p r e m e n t u m small, more or less transverse, sometimes divided into two segment-like sclerites; ligula absent; palpi 2-segmented.

Thorax cylindrical to strongly flattened. General shape of meso- and metathorax sometimes different f r o m prothorax, b u t similar to abdominal segments I—VIII. Thoracic terga consisting of one large undivided plate or two separate plates divided by a membrane, or p r o n o t u m undivided and following thoracic segments with medially divided plates; terga tripartite in Plateros (Figs. 4.11.3 C, 4.H). Lateral portions of meso- and metathoracic venter usually uniform, consisting of anterior and posterior laterotergites; anterior mesothoracic laterotergite bears spiraclular opening (termed spiracular plate); non-functional spiracle sometimes present on anterior metathoracic laterotergite. Ventrolateral part of thoracic pleura consisting of larger episternum and smaller, less sclerotized or reduced epimeron; ventral apex of pleural suture articulates with coxae at a single point. Sternum consists of single sclerite without any sutures or divisions, sometimes reduced or lacking sternal

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Ladislav Bocakand Milada Bocakova

Fig. 4.11.4. Larvae (line = 0.5 mm). A, Duliticola sp., ventral view; B, Xylobanellus erythropterus (Baudi), head, ventral view; C, Lyponia sp., head, ventral view; D, Metanoeus pendleburyi Kleine, head, lateral view; E, Plateros sp., head, lateral view; F, Lyponia sp., head, lateral view; G, Lyponia sp., hind leg; H, Plateros sp., terminal abdominal segments, dorsal view; I, Cautires yuasai Nakane, terminal abdominal segments, dorsal view; J, Lopheros lineatus (Gorham), terminal abdominal segments, lateral view. (A-J from Bocak & Matsuda 2003, © Taylor and Francis Inc., Basingtoke.) pits. Prosternum triangular or slightly elongate; precoxalia separated from prosternum or fused with it. Legs 5-segmented, moderately long to fairly short. Pretarsus simple, slender (Fig. 4.11.4 B). Abdomen 10-segmented, with uniform segments I—VIII. Tergites I-VIII usually consisting of a large median plate, sometimes divided longitudinally into two smaller plates; two large sclerites closely attached to smaller median sclerite in Plateros (Fig. 4.11.4 H). Two laterotergites inserted between tergite and sternite upper laterotergite bearing spiracular opening either in excavate area or in lateral part of sclerite; fixed or movable finger-like processes present on some laterotergites (Fig. 4.11.4 I). Segment IX smaller; shield-like tergite IX, usually not divided, but rarely divided longitudinally as in preceding segments. Segment IX with paired, short, fixed urogomphi, with stick-like urogomphi, or urogomphi absent (Figs. 4.11.4 I, H). Segment X forming short tubular pygopodium (Fig. 4.11.4 J). Spiracles biforous, subequal in size; in or at ventral margin of upper pleurite, rarely on apex

of more or less long, slender tubercles of segments VII and VIII. [Bocak & Matsuda 2003; Costa et al. 1988; Lawrence 1991; Lawrence & Britton 1991; Lawrence et al. 1999 a; Wong 1995,1998.] Morphology, Pupae. Exarate, slender to moderately robust. Pronotum often with small triangular marginal tubercles each bearing a long seta, or with long slender tubercles, or with numerous minute marginal tubercles, or with single small tubercle at each posterior pronotal angle, or tubercle absent. Abdomen with similar tubercles at posterior margin of terga and on sides. Abdominal tubercles bear apical or subapical setae. Pupal urogomphi present in several species. Phylogeny and Taxonomy. Lycidae are generally accepted as a monophyletic group within Elateroidea (sensu Lawrence & Newton 1995; Elateroidea plus Cantharoidea of Crowson 1955). The latest molecular analyses support the monophyly of Lycidae, indicating a close relationship of the family

Lycidae Laporte, 1836

with Lampyridae and Cantharidae (Bocakova et al. 2007), and a more distant relationship with Omalisidae, a group that was previously classified within Lycidae. Kleine (1933) placed lycids and omalisids in one family, and divided his subfamily Lycinae in 15 tribes; h e did not include Duliticola. Bocalt & Bocakova (1990) reviewed lycid classification (without including Duliticola) and grouped the previously described tribes into six subfamilies: Leptolycinae, Lycinae, Erotinae, Metriorrhynchinae, Ateliinae and Calochrominae; several new family group taxa were described within the redefined subfamilies. Medvedev & Kazantsev (1990) described Paralycinae, which is a junior synonym of Lyropaeinae. Miller (1997) elevated the tribe Platerodini to subfamily level based on larval characters and Bocalt & Matsuda (2003) evaluated these features as well as the position of Duliticola, b u t no formal changes were proposed. Kazantsev (2002) introduced Duliticolinae for Duliticola and Platerodrilus, b u t the name was invalid as he designated Platerodrilus as its type and considered Duliticola to be a junior synonym of this genus without studying the type. Type material was examined earlier by Wong (1998) who defended the validity of both Platerodrilus and Duliticola. Later, Kazantsev (2005) proposed a new classification of Lycidae and defined seven subfamilies: Lyropaeinae, Leptolycinae, Ateliinae, Thilmaninae (see Bocak & Brlik, 2008 and 2 - 4 . 1 0 on Omalisidae for f u r t h e r information), Miniduliticolinae (= Duliticolinae), Lycinae and Calochrominae. Miniduliticolinae were elevated to the subfamily status as replacement of the invalid Duliticolinae. Although a formal cladistic evaluation was undertaken by Kazantsev (2005), the final proposed phylogeny is in conflict with the results of his analyses. Kazantsev (2005) did not acknowledge the terminal position of Dexorinae inside Lycidae (Kazantsev 2005: Figs. 595-598). Instead he placed this taxon as sister group of the remaining Lycidae in the final hand-drawn tree (Kazantsev 2005: fig. 599) and elevated it to family rank (Dexoridae). Unfortunately, the analyses were based on an incomplete data set due to the lack of females of neotenous taxa and the poorly k n o w n larval morphology. The impact of presumably affected male adult morphology on phylogenetic hypothesis when females are neotenic was discussed by Mjöberg (1925), Miller (1991), and Bocak & Bocakova (2008). Kazantsev (2005) hypothesized a basal position of Lycidae in Polyphaga and compared larval lycid mandibles with those of Chilopoda and basal insect orders. Kazantsev (2006) postulated a polyphyletic origin of beetles and a basal position of Polyphaga within Neoptera, implying multiple origins of dicondylous mandibles, insect wings, and holometaboly on the basis of the morphology of the larval mandibles of Lycidae (see Beutel et al. 2007). The classifications suggested by Kazantsev (2005, 2006) are in contradiction with phylogenetic patterns based on other morphological structures and molecular data.

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The analyses of sequences of four genes 18S, 28S, and 16S rDNA of Elateriformia demonstrated that Lycidae are deeply nested within Elateriformia (Bocakova et al. 2007). Extensive analyses of molecular data (the above mentioned + ND5 and CYTB mtDNA; (Bocak etal. 2008; Bocak & Bocakova 2008) showed multiple origin of neoteny in Lycidae and the terminal position of neotenic lineages as a consequence. The structural affinities shared by neotenous taxa (and their dissimilarity to other lycids) is due to convergence and not to c o m m o n ancestry. Bocak & Bocakova (2008) obtained five basal lineages based on analyses of molecular data: Libnetinae, Dictyopterinae, Lyropaeinae (including Miniduliticolinae), Ateliinae, and Lycinae. Most k n o w n genera and species are contained in the last group, and most previously recognized subfamilies and/or tribes were given tribal rank within this lineage (e. g., Platerodini, Metriorrhynchini, Calochromini, Erotini, Lycini, Calopterini). The dexorines were given subfamilial status, at least tentatively, based on morphology alone. Available molecular data indicate a very low reliability of traditionally used external characters such as the structure of elytral costae or pronotal carinae.

Acknowledgements Institutions cited in the captions are gratefully acknowledged for allowing us to use copyrighted figures. The Grant Agency of the Czech Republic is acknowledged for the support of our studies on Lycidae (206/02/0982). Finishing work was supported f r o m the grant of the Ministry of Education of the Czech Republic MSM6198959212.

Literature Beutel, R. G., Bocak, L. & Bocakova, M. (2007): Are Polyphaga (Coleoptera) really a basal neopteran lineage - a reply to Kazantsev. - Acta Zoologica 88: 153-158. Bocak, L. (2002): Generic revision and phylogenetic analysis of the Metriorrhynchinae (Coleoptera: Lycidae). - European Journal of Entomology 99: 315-351. Bocak, L. & Bocakova, M. (1987): Notes on the taxonomy of some European species of the family Lycidze.-ActaEntomologicaBohemoslovaca 84:111-121. - (1988): Revision of the genus Dexoris C. O. Waterhouse (Coleoptera, Lycidae). - Acta Entomologica Bohemoslovaca 85:194-204. - (1990): Revision of the supergeneric classification of the family Lycidae (Coleoptera). - Polskie Pismo Entomologiczne 59:623-676. - (2008): Phylogeny and classification of the family Lycidae (Insecta: Coleoptera). - Annales Zoologici 58: 695-720. Bocak, L., Bocakova, M., Hunt, T. & Vogler, A. P. (2008): Multiple ancient origins of neoteny in Lycidae (Coleoptera): consequences for ecology and

Ladislav Bocakand Milada Bocakova

122 macroevolution. Proceedings of the Royal Society Β Biological Sciences 275: 2015-2023. Bocak, L. & Brlik, M. (2008): Revision of the family Omalisidae (Coleoptera, Elateroidea). - Insect Systematics and Evolution 4 6 : 1 8 9 - 2 1 2 . Bocak, L. & Matsuda, K. (2003): Review of i m m a t u r e stages of the family Lycidae (Insecta: Coleoptera). -Journal ofNatural History 37:1463-1507. Bocakova, M. (2001): Revision and phylogenetic analysis of the subfamily Platerodinae (Coleoptera: Lycidae). -European Journal ofEntomology 9 8 : 5 3 - 8 5 . - (2003): Revision of the tribe Calopterini (Coleoptera: Lycidae). - Studies on Neotropical Fauna and Environment 38: 207-234. - (2006): Review of the tribe Lyropaeini (Coleoptera: Lycidae).-EuropeanJournal of Entomology 103:127-136. Bocakova, M., Bocak, L., H u n t , T., Teravainen, M. & Vogler, A. P. (2007): Molecular phylogenetics of Elateriformia (Coleoptera) and the evolution of bioluminescence and neoteny. - Cladistics 23:477-496. Burakowski, B. (1988): Notes on the biology of Xylobanellus erythropterus (Baudi a Selve) (Coleoptera, Lycidae), with description of the i m m a t u r e stages. - Polskie Pismo Entomologiczne 58: 575-585. Costa, C., Vanin, S. A. & Casari-Chen, S. A. (1988): Larvas de Coleoptera do Brasil. 282 pp., 165 pis. Museu de Zoologia da Universidade de Säo Paulo, Säo Paulo. Crowson, R. A. (1955): The Natural Classification of the Families of Coleoptera. 187 pp. Nathaniel Lloyd, London. Eisner, T., Schroeder, F. C., Snyder, N., Grant, J. B., Aneshansley, D. J., Utterback, D., Meinwald, J. & Eisner, M. (2008) Defensive chemistry of lycid beetles and of mimetic cerambycid beetles that feed on them. - Chemoecology 1 8 : 1 0 9 - 1 1 9 . Gravely, F. H. (1915): The larvae and pupae of some beetles f r o m Cochin. - Records of the Indian Museum 11(20): 353-366. Green, J. W. (1949): The Lycidae of the United States and Canada. I. The Tribe Lycini (Coleoptera). - Transactions of the American Entomological Society 75: 53-70. - (1950): The Lycidae of the United States and Canada. II. The Tribe Lygistopterini (Coleoptera). - Transactions of the American Entomological Society 76: 13-25. - (1951): The Lycidae of the United States and Canada. III. The Tribe Platerodini (in part) (Coleoptera). - Transactions of the American Entomological Society 77:1-20. - (1952): The Lycidae of the United States and Canada. IV. The Tribe Calopterini (Coleoptera). - Transactions of the American Entomological Society 78: 1-19. - (1953): The Lycidae of the United States and Canada. V. Plateros (Coleoptera). - Transactions of the American Entomological Society 7 8 : 1 4 9 - 1 8 1 . Hayashi, N. (1986): Key to families of Coleoptera based on the larval characters. Pp. 202-218 in Morimoto, K. & Hayashi, Η. N. (eds.): The Coleoptera ofJapan in Color. Volume I, pp. 202-218. Hoykusha Publishing Co., Osaka. Kazantsev, S. V. (2002): A generic review of Duliticolinae, new subfamily (Coleoptera, Lycidae). - Elytron, 16,5-21.

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(2005): Morphology of Lycidae with some considerations on evolution of Coleoptera. - Elytron 19: 49-226. - (2006): Comparative morphology of mandibular syrtuctures in lycid larvae and its phylogenetic implications (Polyphaga, Hexapoda). - ActaZoologica 87: 229-238. Kleine, R. (1933): Coleopterorum Catalogus auspiciis et auxilio W. J u n k editus S. Schenkling. Pars 128: Lycidae. 145 pp. W. Junk, Berlin. - (1942): Bestimmungtabellen der Lycidae. Bestimmungtabellen der europäischen Coleopteren. 123. Heft, 90 pp. E d m u n d Reiner's Nachf. Emmerich Reitter, Troppau. Laporte, F. L. (1836): Etudes Entomologiques, ou descriptions d'insectes nouveaux et observations sur la synonymie. - Revue Entomologique 4 : 5 - 6 0 . Lawrence, J. F. (1991): Lycidae (Cantharoidea). Pp. 4 2 3 - 4 2 4 in Stehr, F. W. (ed.) Immature Insects. Vol. 2. Kendall/Hunt Publishing Co., Dubuque, Iowa. Lawrence, J. F. &Britton, Ε. B. (1991): Coleoptera (Beetles). Pp. 5 4 3 - 6 8 3 in CSIRO Division of Entomology (ed.) Insects of Australia: a Textbookfor Students and Research Workers, Second Edition. Vol. 2. Melbourne University Press, Carlton, Victoria. Lawrence, J. F. & Newton, A. F. Jr. (1995): Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names). Pp. 779-1006 in Pakaluk, J. & Slipinski, S. A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. M u z e u m i Instytut Zoologii PAN, Warsaw. Lawrence, J. F., Hastings, A. M., Dallwitz, Μ. J., Paine, T. A. & Zürcher, Ε. J. (1999 a): "Beetle Larvae of the World: Descriptions, Illustrations, Identification, and Information Retrieval for Families and Subfamilies." CD-ROM, Version 1.1 for MS-Windows. CSIRO Publishing, Melbourne. - (1999 b): "Beetles of the World: A Key and Information System for Families and Subfamilies." CD-ROM, Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne. Levkanicova, Z. & Bocak, L. (2009) Identification of net-winged beetle larvae (Coleoptera: Lycidae) using three mtDNA fragments: a comparison of their utility. Systematic Entomology 3 4 : 2 1 0 - 2 2 1 . Medvedev, L. N. & Kazantsev, S. V. (1990): A new subfamily and a new genus of Lycidae f r o m South East Asia. Pp 5 5 - 6 0 in Medvedev, L. N. (ed.) Systematics and Ecology of Insects of Vietnam. Nauka, Moscow. Miller, R. S. (1988): Behavior of Calopteron reticulatum (F.) larvae (Coleoptera: Lycidae). - Ohio Journal of Science 8 8 : 1 1 9 - 1 2 0 . Miller, R. S. (1991): A revision of the Leptolycini (Coleoptera: Lycidae) with a discussion ofpaedomorphosis. 403 pp. The Ohio State University (Dissertation.), Columbus, Ohio. Miller, R. S. (1997): I m m a t u r e stages of Plateros floralis (Melsheimer) and discussion of phylogenetic relationships (Coleoptera: Lycidae). - The Coleopterists Bulletin 5 1 : 1 - 1 2 . Mjöberg, Ε. (1925): The mystery of so called 'trilobite larvae' or Perty's larvae' definitely solved. - Psyche 32:119-157.

Telegeusidae Leng, 1920

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Moore, Β. P. & Brown, W. V. (1981): Identification of warning odour components, bitter principles and antifeedants in an aposematic beetle - Metriorrhynchus rhipidium (Coleoptera: Lycidae). - Insect Biochemistry 15:493-499. Nakane, T. (1969): Fauna Japonica, Lycidae (Insecta: Coleoptera). 224 pp. Academic Press of Japan, Tokyo. Perris, E. (1846): Note pour servir a l'histoire du Lygistopterus sanguineus, Dej., Lycus sanguineus, Fabr., Dictyopterus sanguineus, Latr. - Annales de la Societe Entomologique de France Deuxieme Serie, Vol. 4, pp. 343-346. Shelford, R. (1916): A Naturalist in Borneo. 331 pp. Fisher Unwin, Ltd, London (Reprinted by Oxford University Press, 1985). Wong, A. T. C. (1996): A new species of neotenous beetle, Duliticola hoiseni (Insecta: Coleoptera: Cantharoidea: Lycidae) from Peninsular Malaysia and Singapore.-Raffles Bulletin ofZoology 44:173-187. - (1998): A revision of the neotenous "trilobite" larvae of the genera Duliticola and Platerodrilus (Coleoptera: Cantharoidea: Lycidae). 167 pp. National University of Singapore (MSc. Thesis), Singapore.

4.12. Telegeusidae Leng, 1920 John. F. Lawrence Distribution. Telegeusis Horn contains seven species distributed from the southwestern United States to Panama, species ofPseudotelegeusis Wittmer are known from Trinidad and Ecuador, Pseudokarumia angustata Pic was described from Costa Rica, and undescribed species of Pseudokarumia Pic have been seen from Costa Rica to Panama (Ivie 2002). Biology and Ecology. Telegeusid males are often collected at lights or in flight intercept traps, but nothing is known of their habits and females and larvae are unknown. Morphology, Adult males (Fig. 4.12.1). Length about 2 . 5 - 8 mm. Body narrowly elongate, more or less parallel-sided and distinctly flattened. Moderately densely clothed with erect or suberect hairs. Head large, longer than wide, distinctly longer than pronotum, not abruptly constricted posteriorly. Posterior edge of head capsule with two widely separated emarginations. Occipital region long, without transverse ridge. Frontal region abruptly declined anteriorly. Eyes protuberant, widely separated, entire; finely facetted, without interfacetal setae. Antennal insertions slightly elevated, widely separated and exposed from above; subantennal grooves absent. Frontoclypeal suture absent; anterior edge of clypeus straight. Labrum free, strongly transverse, anteriorly trilobed. Antennae usually moniliform (strongly serrate or pectinate

Fig. 4.12.1. Telegeusis nubifer Martin, male, dorsal (© CSIRO Australia), length = 3.5 mm.

from antennomere 3 to 10 in Pseudotelegeusis), 11segmented, with pedicel globular or transverse and much shorter than scape. Mandible long, slender, strongly curved and acute, strongly overlapping when in closed position, with open channel mesally; base somewhat broadened but without mola or prostheca. Maxilla with reduced, setose galea and lacinia; palp 4-segmented with apical palpomere longer than basal palpomeres combined (several times as long in Telegeusis), somewhat flattened and densely clothed with specialized forked hairs. Ligula absent; labial palps 1- to 3-segmented; apical palpomere in Telegeusis very long and similar

124 to that of maxilla. Gular sutures moderately to very narrowly separated. Corpotentorium absent. Cervical sclerites present. Pronotum 0.7-0.75 times as long as wide, slightly wider at middle; sides straight; base not or slightly narrower than elytral bases. Lateral pronotal carinae complete, simple, with or without raised margin; anterior and posterior angles rounded or right; posterior edge simple; disc with two or more submarginal foveae on each side. Prosternum in front of coxae consisting of slender strip of cuticle, much shorter than shortest diameter of procoxal cavity. Prosternal process very short, incomplete and apically acute or absent. Notosternal sutures very short. Procoxae projecting well below prosternum, without concealed lateral extensions. Trochantins exposed, large and subtriangular, setose. Procoxal cavities weakly impressed, strongly transverse, contiguous, externally broadly open, without narrow lateral extensions; internally open. Scutellar shield not abruptly elevated, anteriorly simple, posteriorly broadly rounded or truncate. Elytra 1.4-2.3 times as long as wide and 2.2-3.5 times as long as pronotum, irregularly punctate, without scutellary striole; apices independently rounded and separated by broad gap, exposing several abdominal tergites; epipleuron very short and narrow or absent. Mesoventrite separated by complete sutures from mesanepisterna, which are distinctly separated at midline; anterior edge on same plane as metaventrite, without paired procoxal rests. Mesoventral cavity absent. Mesocoxae conical and projecting, with exposed, setose trochantins. Mesocoxal cavities weakly impressed or virtually absent, contiguous, open laterally (closed by both mesanepisternum and mesepimeron); mesometaventral junction absent. Metaventrite moderately to strongly convex, with straight sides; discrimen very long; postcoxal lines absent; exposed portion of metanepisternum elongate. Metacoxae transverse b u t with mesal portions subconical and strongly projecting, contiguous, horoizontally oriented, extending laterally to meet elytra; plates absent. Metendosternite with lateral arms very short, laminae absent, anterior process moderately long and anterior tendons very close together. H i n d wing with entire surface and margins densely clothed with short hairs; apical field long, without sclerotizations; radial cell incomplete basally; crossveins r3 and r4 absent; R-M loop forming narrowly acute angle; base of RP very short or absent; medial spur straight, extending to wing margin; medial field with two free veins unconnected at base; anal lobe somewhat reduced; anal embayment shallow. Femoral attachment of mid trochanter strongly oblique with base of femur separate from coxa. Legs slender; tibial spurs well developed, pubescent; tarsi 5-5-5; tarsomeres simple; pretarsal claws more or less serrate or pectinate. Abdomen flattened, with eight free ventrites (sternites II-IX). Ventrite 1 shorter t h a n 2, w i t h o u t postcoxal lines; metacoxal cavities weakly developed and intercoxal process absent. Functional

John F.Lawrence

spiracles on abdominal segment VIII present. Spiracles located on pleural membrane. Luminous organs absent. Anterior edge of sternite VIII in male without median strut. Anterior edge of sternite IX in male broadly rounded, w i t h o u t spiculum gastrale; tergite IX truncate. Tergite X well developed and free. Aedeagus of modified trilobate type with phallobase broadly overlapping bases of parameres, which are fused together basally and fused to base of penis; phallobase symmetrical, without anterior strut; parameres not outwardly hooked; anterior edge of penis with short struts. [Horn 1895; Barber 1952; Allen & H u t t o n 1969; Zaragoza Caballero 1975,1990; Wittmer 1976 b; Fleenor & Taber 2001; Ivie 2002.] Phylogeny and Taxonomy. Telegeusis deUlis Horn was placed in the family Drilidae by its describer (Horn 1895) and remained there in the Junk Catalogue (Olivier 1910). The genus was made the type of a new family by Leng (1920) who placed it with some doubt in a superfamily containing both Lymexylidae and Micromalthidae. This move may have been based on comments made by Muir and cited by Barber (1913) on the similarities in the aedeagi of Telegeusis and the lymexylidAtractocerus brevicornis (Linnaeus) (as A. africanus Boheman). Barber (1952) illustrated the aedeagus of Telegeusis nubifer Martin and placed Telegeusis (but not Micromalthus LeConte) in the family Lymexylidae. Crowson (1955,1972) considered Telegeusis to be "unmistakably Cantharoid in the form of its tentorium, met-endosternite and aedeagus", noting also that it differed from Lymexylidae in the presence of functional spiracles on segment VIII and in the form of the mandibles and abdominal apex. Although the placement of Telegeusidae in Elateroidea (including Cantharoidea) has been followed in most recent works (Zaragoza Caballero 1990; Lawrence & Newton 1995; Lawrence et al. 1999; Ivie 2002; Miller 2002), its position within the complex is still in doubt. Crowson (1972) placed the

Fig. 4.12.2. Telegeusis nubifer Martin, male, head and prothorax, lateral (© CSIRO Australia).

Telegeusidae Leng, 1920

family near Phengodidae and hypothesized that the l u m i n o u s larva illustrated by Barber (1908) m i g h t be a larviform female telegeusid. However, the true identity of these glow-worms with red light organs on the head had been established m u c h earlier by Haase (1886,1888), who collected a female in copula with a male of a true phengodid, Phrixothrix heironymi (Hasse) (Wittmer 1976 a; Viviani & Bechara 1993). In a cladogram produced by Branham and Wenzel (2001, 2003) based on adult males only, Telegeusis and Pseudotelegeusis formed a clade w i t h several genera of Phengodidae, Cantharidae and Omethidae. In the molecular cladograms produced by Bocakova et al. (2007), Telegeusis formed a monophyletic g r o u p with Drilonius Kiesenwetter (Omethidae) at or near the base of the Elateroidea (sensu lato) or was located at the base of Elateroidea (excluding Drilonius and the Eucnemidae). In no case was the g r o u p closely related to Phengodidae. Wittmer (1976 b) described Pseudotelegeusis, the two species of which differ f r o m those of the type genus in having strongly serrate antennae and simple, 1-segmented labial palps. Pseudokarumia was compared by its author (Pic 1931) w i t h Drilocephalus Pic and placed in the family Karumiidae (now a subfamily of Dascillidae) by Arnett (1964) and Paulus (1972). After an examination of the type, P. angustata Pic, plus several undescribed species, Ivie (2002) transferred this genus to Telegeusidae. Pseudokarumia species have the short, filiform antennae of Telegeusis combined with the unmodified labial palps resembling those in Pseudotelegeusis.

Acknowledgements CSIRO Australia is acknowledged for s u p p o r t of research a n d for allowing the use of copyrighted images.

Literature Allen, R. T. & Hutton, R. S. (1969): A new species of Telegeusidae (Leng) from Panama (Coleoptera: Cantharoidea). - The Coleopterists Bulletin 23:109-112. Arnett, R. H. (1964): Notes on Karumiidae. - The Coleopterists Bulletin 18: 65-68. Barber, H. S. (1908): The glow-worm Astraptor. - Proceedings of the Entomological Society of Washington 9 (1907): 41-43. - (1913): Observations on the life history of Micromalthus debilis Lec. (Coleoptera). - Proceedings of the Entomological Society of Washintgon 15:31-38. - (1952): Notes on Telegeusis and some relatives. - Pan-Pacific Entomologist 28:163-170. Bocakova, M., Bocak, L., Hunt, T., Teräväinen, Μ. & Vogler, A. P. (2007): Molecular phylogenetics of Elateriformia (Coleoptera): Evolution of bioluminescence and neoteny. - Cladistics 23:477-496. Branham, M. C. & Wenzel, J. W. (2001): The evolution of bioluminescence in cantharoids (Coleoptera, Elateroidea). - Florida Entomologist 84 (4): 565-587.

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(2003): The origin of photic behavior and the evolution of sexual communication in fireflies (Coleoptera: Lampyridae). - Cladistics 19 (1): 1-22. Crowson, R. A. (1955): The Natural Classification of the Families of Coleoptera. 187 pp. Nathaniel Lloyd, London. - (1972): A review of the classification of Cantharoidea (Coleoptera), with the definition of two new families, Cneoglossidae and Omethidae. - Revista de la Universidad de Madrid 21 (82): 35-77. Fleenor, S. B. & Taber, S. W. (2001): A new long-lipped beetle from Texas and a review of the genus Telegeusis Horn (Coleoptera: Telegeusidae). - The Coleopterists Bulletin 55:481-484. Hasse, Ε. (1886): Ein neuer Phengodes. - Entomologische Nachrichten 12:218-219. Hasse, E. (1888): Zur Kentniss vonPhengodes.-Deutsche Entomologische Zeitschrift 32:145-167. Horn, G. H. (1895): Coleoptera of Baja California (Supplement I). - Proceedings of the California Academy of Sciences (2) 5: 242-243. Ivie, M. A. (2002): The transfer of Pseudokarumia Pic from Dascillidae (Karumiinae) to Telegeusidae, with a key to world genera of Telegeusidae. - The Coleopterists Bulletin 56 (4): 582-584. Lawrence, J. F. & Newton, A. F. Jr. (1995): Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names). Pp. 779-1006 in Pakaluk, J. & Slipiriski, S. A. (eds.) Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warsaw. Lawrence, J. F., Hastings, A. M., Dallwitz, Μ. J., Paine, T. A. & Zürcher, Ε. J. (1999): Beetles of the World: A Key and Information System for Families and Subfamilies. CD-ROM, Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne. Leng, C. W. (1920): Catalog of the Coleoptera of America, north of Mexico, χ + 470 pp. John D. Sherman Jr., Mount Vernon, New York. Miller, R. S. (2002): 60. Telegeusidae Leng 1920. Pp. 179-180 in Arnett, R. H. Jr., Thomas, M. C., Skelley, P. Ε & Frank, J. H. (eds.), American Beetles. Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Gainesville, Florida. Olivier, E. (1910): Pars 10. Coleopterorum Catalogus. Pars 10. Rhagophthalmidae, Drilidae. 10 pp. W. Junk, Berlin. Paulus, Η. F. (1972): Die systematische u n d phylogenetische Stellung der Karumiidae, mit einer Beschreibung von Escalerina serraticornis n. sp. aus S.-Persien. - Senckenbergiana 53:37-54. Pic, M. (1931): Coleopteres exotiques en partie nouveaux. - L'Echange 4 7 : 9 2 - 9 6 . Viviani, V. R. & Bechara, E. J. H. (1993): Biophysical and biochemical aspects of phengodid (railroadworm) bioluminescence. - Photochemistry and Photobiology 58 (4): 615-622. Wittmer, W. (1976 a): Arbeiten zu einer Revision der Familie Phengodidae (Coleoptera). -Entomologische Arbeiten aus dem Museum G. Frey 27:415-524. - (1976 b): Eine neue Gattung der Familie Telegeusidae (Col.). - Mitteilungen der Schweizerischen Entomologischen Gesellschaft 49:293-296.

126 Zaragoza Caballero, S. (1975): Una nueva especie de Τelegeusis Horn (Coleoptera: Telegeusidae) de Chamela, Jalisco, Mexico. - Anales Instituto de Biologia NacionalAutönoma de Mexico (Zoologia) 46 (1): 63-68. - (1990): Una especie nueva de Telegeusis Horn, 1895 (Coleoptera: Telegeusidae) de Nuevo Leon, Mexico. - Anales Instituto de Biologia Nacional Autonoma de Mexico (Zoologia) 61 (2): 307-312.

4.13. Phengodidae LeConte, 1861 Cleide Costa and Santiago Zaragoza-Caballero

D i s t r i b u t i o n . The family Phengodidae includes 31 genera and 244 species distributed t h r o u g h o u t the New World f r o m extreme southern Canada to Chile (between 40° Ν and 38° S) including the Nearctic and Neotropical biogeographical regions. [Lawrence 1982; Zaragoza 1984; Costa etal. 1988.] Biology a n d Ecology (Figs. 4.13.3 A-F; 4.13.4 Α-D). Larvae and females are predators, feeding on millipedes and other arthropods occurring in soil and litter. Males are often attracted to lights at night, are short-lived and probably do not feed. Males of Distremocephalus Wittmer bear setose foveae (seemingly of glandular nature) on abdominal ventrites V and VI (Fig. 4.13.5 Α-B). They are formed by two fringes of n u m e r o u s setae that are exposed when the male adopts defensive postures similar to those observed in some Staphylinidae (abdomen raised over back). Male bioluminescence is rare b u t some species may have light organs on the head, prothorax or abdomen (see below). Larvae and females have a set of luminescent organs on the head and t r u n k segments, and are commonly k n o w n as "glow worms" or "railroad worms". The t r u n k organs emit yellow, orange or green light, and some species have red-emitting lanterns on the head. The insect resembles a glowing cigarette when only the red lanterns are shining in the darkness. Females of the mastinocerine genera Mastinomorphus Wittmer and of Phrixothrix Olivier laid a compact mass of eggs d u r i n g a single oviposition in soil fissures or chambers that they dig and encircle the eggs until eclosion. The round eggs are brownish-orange, hard, and have an average diameter of 1 m m . The incubation time observed in the laboratory at 21°C varies f r o m 33 to 51 days. During this period, the females are aggressive and attack intrusive objects. The females die within one week after the eggs hatch. Eggs of the phengodine Zarhipis integripennis (LeConte) are round or oval, white, and become luminous after a m o n t h . Neonatal larvae are active and l u m i n o u s b u t do not feed, the first feeding instar being the second one. Larvae are ambulatory and generally f o u n d on dark soil beneath decaying logs and leaves or litter

Cleide Costa and Santiago Zaragoza-Caballero in tropical forest, open fields or marshes. At night, the shining head lantern can frequently pinpoint them, the lateral lanterns remaining dark. Larvae were also observed with all lanterns switched on, making it easy to detect t h e m at great distances. Larvae feed mainly on millipedes, b u t other prey like living termite workers may also be caught. In the laboratory larvae of Mastinomorphus sp. were fed with chicken liver extract. Several phengodid larvae were seen to m o u n t the back of a millipede and cut off its head, consequently paralyzing the prey. After that, the larva first eats the head followed by the body, leaving the chitinized segments empty. Phrixothrix larvae often eject a reddish dark liquid from the anus when disturbed, a behavior not yet understood. Pupae are found buried in soil. In the laboratory the pupal stage of some Mastinocerus Solier and Phrixothrix hirtus Olivier lasted 20 and 22 days, respectively, at 21°C, and that of Zarhipis integripennis lasted 2 0 - 3 5 days. Observations on the adult behavior of Brazilian species f r o m West central Cerrado revealed that they are very sensitive to the climatic conditions, being attracted to white, UV and blue light traps only on windless, warm (19-26°C) and h u m i d nights (85-95% relative humidity). In the laboratory, the longevity of the adult male is very short, in general no longer than one week even when maintained in h u m i d i fied vials and fed sugary water. Bioluminescence f r o m the eggs of Brazilian species is weakly indicated and only detectable f r o m the embryos after day 15, using a photon-counting apparatus (Viviani & Bechara 1997). Eggs of Zarhipis integripennis show a weak glow after 25 days of incubation. Larvae of the genus Brasilocerus Wittmer display eleven pairs of dorsolateral lanterns, emitting green or yellow-green ^ m a x = 5 5 0 - 5 5 7 nm), and 2 large anterior lanterns of yellow-green color (λιη2Χ = 565-570 nm). Those of Mastinocerus spp. exhibit eleven pairs of dorsolateral lanterns emitting yellow light a m a x = 578-580 nm) and the cephalic lantern may be seen frontally, in contrast to other species whose light can be seen dorsally. Larvae of Phrixothrix hirtus have red lights on the head and pairs of lateral lanterns at the edges of the body which emit yellow light. Larvae of Phrixothrix spp. display eleven pairs of dorsolateral lanterns, emitting green or yellow-green light ^ m a x = 5 3 5 - 5 6 8 nm), and two anterior (cephalic and postcephalic) lanterns, appearing visually as a single lantern that emanates reddish light (λπ13Χ = 6 0 0 - 6 3 8 nm). P. hirtus radiates yellow-green light f r o m the lateral lanterns ^ m a x = 568 nm) and the reddest light among the studied phengodids (λιτ13Χ = 636 versus 6 0 0 - 6 2 0 nm) (Viviani & Bechara 1997). In general, male pupae lack the head lantern, b u t bright lateral lanterns are retained. During the pupal stage of Phrixothrix hirtus, females retain the red lanterns b u t males lack them. A diffuse red glow can be observed on the cephalic region only in the first two days of the pupal stage and disappears afterwards. In Mastinomorphus sp. and Phrixothrix hirtus the

Phengodidae LeConte, 1861

color of the lateral rows of lanterns appears to be identical in adult and larval stages, visually in the former species and only spectrophotometrically in the latter species (λ ηΐ2χ = 568 and 563 n m , respectively). Adult females of Phengodes H o f f m a n n s e g g illuminate twelve segments in the yellow-green spectral region, either as spots at the posterolateral corner or transverse shining lines along the posterior margin. The three thoracic segments show a transverse band of bioluminescence, broken in the middle. On the nine abdominal segments there is a transverse band plus a pair of l u m i n o u s spots, with the exception of the abdominal segm e n t IX, where the band was found to be lacking. In Phrixothrix hirtus females the bioluminescence pattern is very similar to that of the larval stage. In female larvae a light organ asymmetrically located on the prothorax between the two lateral lanterns was perceived (Harvey 1952). In general, the color of adult male light emission ranges f r o m green to yellow, b u t some species of Euryopa Gorham and Mastinomorphus emit orange and reddish light. The lanterns of adult males of the Brazilian Mastinocerinae are located dorsolateral^ along the thoracic and abdominal segments, the prothoracic segment having a dorsomedian light organ. In Euryopa laurae Wittmer there is a very small light organ on the head between the eyes, emitting the same orange color as that of the lateral lanterns. The phengodine Pseudophengodes brasiliensis Wittmer was once found radiating a greenish glow. This species has lampyrid-like light organs on the ventral side of the penultimate segment. The function of bioluminescence in the phengodids is not well understood. The continuous glow of the head lanterns when the larva is walking suggests an illumination function, whereas the lateral lanterns may serve a defensive function. A sudden flash m i g h t be used to repel potential predators. The initial bioluminescence intensity quickly decreases, possibly reducing the risk for being targeted by predators. Also, the dorsolateral location of the lanterns in a walking insect suggests that the light is to be perceived by predators above t h e m (Viviani & Bechara 1997). Aposematism associated with distateful properties is also a possible function for the lateral lantern light (Crowson 1981; Sivirisky 1981). In some species the same luciferase isoform t h r o u g h o u t the entire life cycle was identified. This contrasts with lampyrids and elaterids whose bioluminescence colors change in ontogenic succession of different luciferase isoenzymes (Colepicolo-Neto etal. 1986). According to Tiemann (1970) the bioluminescence of adults does not play an important role in sexual attraction. It seems that sexual attraction is mediated by pheromones (Lloyd 1978). Viviani & Bechara (1997) suggested a defensive function to the adult bioluminescence and illumination and self-defense to the bioluminescence of larvae and larviform females. According to Crowson (1981) "the Cantharoidea are noted for the paedomorphic tendencies of their adults,

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which are particularly marked in the l u m i n o u s groups". Among them, the Phengodidae seem to attain the highest level of paedomorphosis, with females retaining many more larval features t h a n neotenic females of other groups of Cantharoidea, including the Rhagophthalmidae. [Buck 1950; Tiem a n n 1967, 1970; Wittmer 1976; Crowson 1981; Costa et al. 1988; LeSage 1991; Viviani & Bechara 1993,1997.] Morphology, Adult Males (Figs. 4.13.1 A-J; 4.13.2 A-K; 4.13.3 D). Total length 3.3-25 m m . Body elongate, parallel. Surface smooth, with or without pubescence. Head (Figs. 4.13.1 F - H ; 4.13.2 A-B; 4.13.3 D) prognathous to slightly declined, subquadrate to slightly transverse. Frontal area variable, projected at apex or vertical. Eyes lateral, usually almost round; strongly emarginate posteriorly only in Νephromma Wittmer (Fig. 4.13.2 Α-B) and slightly emarginate in some Phengodes spp.; p r o t r u d i n g or reduced (between 1.6 and 5.0 times head width); finely facetted, without interfacetal setae. Antennal insertions usually widely separated, rarely approximate (separated by 2.0-0.5 times the length of the first antennomere), sometimes raised to form antennal tubercles (Fig. 4.13.1 H). Frontoclypeal suture present or absent. Labrum free or fused, often partly or completely concealed beneath clypeus. Antennae (Figs. 4.13.1 A-B; 4.13.3 D) usually 12-segmented, b u t occasionally 10-segmented (in Decamastinocerus Wittmer, Penicillophorus Paulus), or 11-segmented (Euryopa, Euryognathus Wittmer, Adendrocera Wittmer, Acladocera Wittmer); antennomere 4 to penultimate antennomere serrate (penicillophorus, Adendrocera, Acladocera) to bipectinate or plumose, with rami beginning on base of antennomere and apical antennomere lanceolate (.Distremocephalus Wittmer, Euryopa, Euryognathus) or filiform (Phengodes, Pseudophengodes Pic, Ptorthodiellus Wittmer). Mouthparts prognathous (Fig. 4.13.1 H). Mandibles (Figs. 4.13.1 E - H ; 4.13.2 E) well developed, falcate, apically acute, with or without additional teeth. Maxilla with galea well developed or reduced, simple or pubescent; lacinia partly reduced (Phrixothrix hirtus) or absent; palps (Figs. 4.13.1 C-E) 3- or 4-segmented; larger in Phengodinae or small and with truncate apex in Mastinocerinae and Penicillophorinae. Labial palps (Fig. 4.13.1 E) with 1-, 2- or 3-segmented, with variable apex. Palps in general variable, usually acuminate or truncate at the apex and lacking an elongate apical segment. Posterior tentorial pits and gular sutures often approximate or fused together. Cervical sclerites well developed. Prothorax (Figs. 4.13.1 H; 4.13.2 A, G) about 0.8-1.65 times as long as wide. P r o n o t u m flattened or slightly biconcave, laterally explanate or not; anterior margin curved or truncate; posterior margin complete or discontinuous; anterior angles straight or truncate; h i n d angles truncate to slightly acute; surface smooth or more strongly punctate,

128

Cleide Costa and Santiago Zaragoza-Caballero

Fig. 4.13.1. A, Mastinowitttnerus mexicanus Zaragoza, segments 4—12 of antenna (after Zaragoza 1984); Β, Paraptorthodius queretaroensis Zaragoza, antenna (after Zaragoza 1999); C, Phengodes (Phengodes) vazquezae Zaragoza, right maxillary palp (after Zaragoza 1978); D, Phengodes (Phengodella) leonilae Zaragoza & Wittmer, maxillary palp (after Zaragoza & Wittmer 1986); E, Mastinowittmerus mexicanus, labium, mandible apex and maxillary palp (after Zaragoza 1984); F, Cenophengus guerrerensis Zaragoza, head, ventral view (after Zaragoza & Wittmer 1986); G, Phengodes (Phengodella) atezcanus Zaragoza, head, ventral view (after Zaragoza & Wittmer 1986); H, Phengodes (Phengodes) bipennifera Gorham, dorsal view of head and pronotum (after Zaragoza & Wittmer 1986); I, Phrixothrix hirtus Olivier, hind wing (after Costa etal. 1999); J, Cenophengus guerrerensis, hind wing, (after Zaragoza 1991), line= 1 m m .

Fig. 4.13.2. A, Β, Nephromma barberi Wittmer, dorsal view of head and pronotum, lateral view of eyes (after Wittmer 1976); C, Mastinowittmerus mexicanus, first tarsomeres (after Zaragoza 1984); D, Ptorthodius mandibularis Gorham, claws; E, Oximastinocerus unicolor (Pic), mandibles; F, Howdenia sp., metepisternum; G. Phrixothrix sp., pronotum; H, Pseudophengodes sp., abdomen; I, Phengodes (Phengodes) vazquezae, abdomen; J, K, Phrixothrix hirtus, aedeagus (after Costa et al. 1999), Fig. A-H, line = 1 mm; Fig. J - K , line = 0.1 mm.

130

Cleide Costa and Santiago Zaragoza-Caballero

Fig. 4.13.3. A-F, Phrixothrix hirtus. A, larva, dorsal view; B-C, male pupa, dorsal and lateral view; D, habitus, adult male; E, larva, 9th and 10th ventral abdominal segments; F, female adult, apex of abdomen, ventral (arrow pointing the ooporus) (after Costa et al. 1999), Fig. Α-D, line = 5 mm; Fig. E-F, line = 1 mm.

Phengodidae LeConte, 1861

with or without setae. Prosternum in front of coxae usually very short, almost reduced to slender transverse bar (longer in Cenophengus LeConte), triangularly produced behind. Functional spiracles produced, located laterad of procoxae. Prosternal process reduced. Hypomera simple; width and internal margin variable. Procoxae cone-shaped, projecting and narrowly separated; procoxal cavities posteriorly open. Trochantin almost triangular. Protrochanter strongly obliquely articulated to profemurbase. Scutellar shield well developed; posterior edge wide, truncate, lobulate or emarginate; anterior edge carinate and deeply emarginate. Elytra (Fig. 4.13.3 D) slightly to extremely shortened, exposing abdominal apex or all tergites; distal portion lobulate or acuminate; epipleura wide at base, reduced posteriorly; surface irregularly punctured and pubescent. Mesoventrite short; anterior edge deeply emarginate. Mesocoxae projecting, conical, moderately separated. Mesocoxal cavities closed by mesanepisternum and mesepimeron. Mesotrochantins setiferous. Mesotrochanter oval and obliquely articulated to mesofemur base. Mesothoracic spiracles between mesanepisternum and mesepimeron. Metaventrite wide and slightly convex; enlarged posteriorly; posterior margin triangularly projecting; discrimen moderately long or short. Metanepisternum (Fig. 4.13.2 F) enlarged, triangular, posteriorly reduced. Metepimeron curved and reduced. Metacoxae well developed, transverse, narrowly separated; metatrochanters oval and obliquely articulated to metafemur base. Metendosternite with stalk well-developed, very narrow and elongate, with a conspicuous nearly central indentation, broadening out forward into a small, lobe-like lamina and then narrowing abruptly to apex; tendons indistinct (Phrixothrix hirtus). Hind wing (Fig. 4.13.11-J) not folded under the elytra, usually 2.2 as long as wide; radial cell closed (Phengodes, Nephromma, Distremocephalus) or open (Penicillophorus, Adrendocera, Howdenia Wittmer, Spanglierella Wittmer and some species of Cenophengus); r4 well marked, complete or not; wedge cell absent; medial field containing a varied number of free veins: five (MP3, MP 4 , CUA2, AA3, AA4) in some Phengodes species; four in some Mastinocerus and Eurymastinocerus; three in some species of Ptorthodiellus, Phrixothrix, Pseudophengodes, Cenophengus, and one in some Cenophengus spp. Legs elongate; tarsi 5-5-5, simple, often densely setose; tarsomeres 3 and 4 with a ventral, densely setose lamella (Phrixothrix); tarsomere 1 sometimes with a row of aligned spines forming ventral comb (Mastinowittmerus Zaragoza, Phrixothrix) (Fig. 4.13.2 C) or with combs on tarsomeres 1 and 2 in Distremocephalus, Brasilocerus, Nephromma. Claws simple or dentate, without setae (Fig. 4.13.2 D). Tibial spurs usually absent, present in some species of Phengodes, Phrixothrix and Eurymastinocerus; a crown of apical spines in some Phengodes and Mastinowittmerus species. Abdomen with eight completely free ventrites (sternites II-IX); sternites V and VI (Fig. 4.13.5 A-B)

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occasionally with setose foveae (Distremocephalus spp.), formed by two fringes of numerous short and long setae. In Penicillophorus species setose patches are on the pleural region of the first two abdominal segments and in Howdenia spp on the metanepisternum. The penultimate ventrite in Pseudophengodes sp. (Fig. 4.13.2 H) is pointed, while that in Zarhipis species and some species of Phengodes is more or less emarginate (Fig. 4.13.21). Functional spiracles posteriorly in pleural membrane. Luminous organs rare in males but if present may be located: frontally on head between eyes; dorsolateral^ along the thoracic and abdominal segments, dorsomedially on the pronotum and on penultimate or last ventrite. Aedeagus (Fig. 4.13.2 J-K); trilobate; phallobase small, parameres enveloping penis or not; flagellum conspicuous and freely extensible (not encapsulated inside the penis), at rest, encircled or not around penis; when distended about as long as penis length or in Euryopa several times as long. [Wittmer 1976, 1988; Lawrence 1982; Zaragoza 1989; Wittmer 1993; 1996; Costa etal. 1999; Lawrence etal. 1999 b.] Morphology, Adult Females (Figs. 4.13.3 F; 4.13.4 Α-D). Females are much larger than the males and are larviform, retaining many larval features. Description based mainly on the mastionocerine species Phrixothrix hirtus and P. hieronymi (Haase). Length: 34 mm; width of pronotum: 5 mm. Larviform, orthosomatic, less depressed than mature larva, integument and pilosity yellowish, very similar to mature larva, except for the following characters: stronger punctures on head and a type of flattened scale-like ornamentations more evident on the integument; ventral area of mandibles with dentiform microtrichia; dorsal region of maxillae with longer setae; epipharynx with denser microtrichia; hypopharynx formed by a simple, membranous lobe, clothed with short hairs; hypopharyngeal sclerome weakly sclerotized, apparently not associated with mandibles, clothed with microtrichia forming oval patches and an apical fringe of ramose setae; annular spiracles present on mesothorax and abdominal segments I—VIII; sternite IX with a large transverse oopore (Fig. 4.13.3 F); anal opening bearing two transverse dark sclerotized lamellae. Internal reproductive system (Fig. 4.13.4 D) consisting of a pair of ovaries connected with a pair of asymmetrical oviducts which join to form a median oviduct opening into vagina; bilobed spermatheca continuing as a small weakly sclerotized pouch that opens into vagina near common oviduct. Ovaries lying between abdominal segment IV to VIII (Fig. 4.13.4 C); right oviduct longer than left one. Other than oopore, special structures associated with oviposition absent. Females of the genus Phrixothrix are easily distinguished from larvae by the presence of an oopore, which is absent in larvae, and by the annular spiracles, biforous in larvae. According to Buck (1950) in Phengodes the cream-colored individuals are adult females while

132 the brown pigmented forms are larvae. [Haase 1888; Costa etal. 1999; Lawrence etal. 1999 b.] Morphology, Mature Larva. Length 15-65 mm. Orthosomatic, elongate and depressed (Fig. 4.13.3 A). Integument moderately sclerotized, shiny, covered dorsally and ventrally by varying setae. Body color varies from cream, orange, yellow or red, to brown or black. Head and segment X usually more sclerotized. Head prognathous, one third to half the width of pronotal base, depressed, retractable. Epicranial suture absent. Endocarina absent. One stemma on each side of head below antenna. Labrum usually fused with clypeus and frons, separate of frons by a weak suture in Phrixothrix hirtus. Epipharynx with microtrichia and sensory pores. Antennae 3-segmented; basal segment telescopic, with dorsal and ventral sensory pores; second segment longer bearing an apical sensorium; distal segment reduced. Mandibles falciform, dorsally channeled from base to near tip, with or without a retinaculum. Maxilla with distinct stipes and cardo; galea 2-segmented, or palp-like; lacinia lobate; maxillary

Cleide Costa and Santiago Zaragoza-Caballero

palp 3- or 4-segmented. Labium with prementum narrowed at base; postmentum elongate; ligula membranous, well developed. Labial palp 2-segmented. Hypopharynx forming a simple membranous lobe densely covered by microtrichia. Gular sutures present or absent. Pronotum transverse, narrowed apically; meso- and metanotum similar in length. Mesothorax with a pair of ventrolateral anterior biforous spiracles. Legs increasing in size from pro- to metathoracic, 5-segmented including a claw-like pretarsus; coxae elongate, wider at base, bearing simple setae of varied size; trochanter subtriangular, femora and tibiotarsus elongate; trochanters, femora and tibiotarsus bearing simple and stout setae of varied size; pretarsus bearing two stout, simple setae. Abdominal segments I—IX transverse; segments I-VIII laterally bearing paired biforous spiracles, smaller than those on thorax; segment IX smaller; segment X (Fig. 4.13.3 E) reduced and probably acting as a proleg. Anal slit transverse. Urogomphi absent. Paragraph LeSage (1991), probably based only on the mandible of Phengodes laticollis LeConte, described and illustrated by Peterson (1951), stated that the

Fig. 4.13.4. Α-D, Phrixothrix hirtus. A-C, female adult, dorsal, lateral and ventral view; D, reproductive system, ventral (after Costa etal. 1999), line = 5 mm.

Phengodidae LeConte, 1861

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robust divergent projections, with truncate apex; a pair of rounded ventral spiracles located at apex of small tuberculiform projection. Mesonotum half as long as metanotum. Segments I-VII transverse; segment VIII narrower; segments I-VIII bearing paired lateral rounded spiracles located at apex on small tubular tubercle. Tergites IX-X, sternite IX and aedeagus partially extroverted. [Costa et al. 1999.] Morphology, Female Pupa. Larviform, similar to larva and adult female, but integument lighter and less pilose; mandible not sulcate, broad, triangular shaped, shorter than in larva and in the female, not falciform, large dorsal seta, not spatulate, retinaculum indicated as an indentation in the mesal margin; spiracles annular; antenna proportionatelly smaller than in larva and neotenic female; oopore indicated as a weak transversal carina on the sternite IX. The larviform female pupa of Phrixothrix hirtus can be easily distinguished from the larva and larviform female by the lighter coloration of the integument, and by the mandible triangular-shaped and not sulcate. [Costa etal. 1999.]