Fishes of the Gulf of Mexico, Volume 2: Scorpaeniformes to Tetraodontiformes 9780292794214

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Fishes of the Gulf of Mexico Volume 2

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Fishes of the Gulf of Mexico by John D.McEachran and Janice D.Fechhelm Volume 2: Scorpaeniformes to Tetraodontiformes

University of Texas Press, Austin

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Publication of this book was made possible by the generous financial assistance of the following organizations: Coastal Bend Community Foundation, Harvey Weil Sportsman /Conservationist Award Fund, Corpus Christi, Texas; Harte Research Institute for Gulf of Mexico Studies, Texas A&M University— Corpus Christi; Offshore Operators Committee, New Orleans; Shell Offshore, Inc., New Orleans; Texas Agricultural Experiment Station, Texas A&M University, College Station. Copyright © 2005 by the University of Texas Press All rights reserved Printed in the United States of America First edition, 2005 Requests for permission to reproduce material from this work should be sent to: Permissions University of Texas Press P.O. Box 7819 Austin, TX 78713-7819 www.utexas.edu /utpress/about /bpermission.html  The paper used in this book meets the minimum  requirements of ANSI /NISO Z39.48-1992 (R1997) (Permanence of Paper). ISBN: 0-292-70634-0 (cl. : alk. paper : v. 2) Library of Congress Cataloging-in-Publication Data (from Vol. 1) McEachran, John D. Fishes of the Gulf of Mexico / John D. McEachran and Janice D. Fechhelm. — 1st ed. p. cm. Includes bibliographic references and index. ISBN 0-292-75206-7 (cl. : alk. paper : v. 1) 1. Fishes—Mexico, Gulf of. 2. Fishes— Mexico, Gulf of—Identification. I. Fechhelm, Janice D. II. Title. QL621.56.M383 1998 597.177364 — dc21 98 – 4605

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CONTENTS

Acknowledgments vii Introduction 1 Scope of the Book 1 How to Identify Fishes 2 Biogeography of Fishes of the Gulf of Mexico 4 Appendix: New Records and Emendations to Fishes Included in Volume 1 of Fishes of the Gulf of Mexico 14 Literature Cited 19 Fishes of the Gulf of Mexico Scorpaeniformes 23 Dactylopteridae 24 Scorpaenidae 26 Triglidae 51 Peristediidae 70 Perciformes 85 Centropomidae 97 Moronidae 105 Acropomatidae 111 Howellidae 117 Serranidae 119 Grammatidae 194 Opistognathidae 199 Priacanthidae 211 Apogonidae 216 Epigonidae 235 Malacanthidae 243 Pomatomidae 251 Echeneidae 253 Rachycentridae 263 Coryphaenidae 265 Carangidae 269 Bramidae 304 Caristiidae 311 Emmelichthyidae 313

Lutjanidae 317 Symphysanodontidae 338 Lobotidae 342 Gerreidae 344 Haemulidae 359 Inermiidae 382 Sparidae 385 Polynemidae 405 Sciaenidae 410 Mullidae 447 Pempheridae 453 Bathyclupeidae 455 Chaetodontidae 458 Pomacanthidae 471 Kyphosidae 482 Cirrhitidae 485 Pomacentridae 487 Labridae 504 Scaridae 527 Zoarcidae 545 Chiasmodontidae 550 Percophidae 563 Uranoscopidae 567 Tripterygiidae 572 Dactyloscopidae 578 Labrisomidae 586 Chaenopsidae 610 Blenniidae 624 Gobiesocidae 641 Callionymidae 646 Draconettidae 652 Eleotridae 654 Gobiidae 661 Microdesmidae 730 Ephippidae 735 Luvaridae 738 Acanthuridae 740

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vi CONTENTS

Scombrolabracidae 745 Sphyraenidae 747 Gempylidae 753 Trichiuridae 764 Scombridae 771 Xiphiidae 789 Centrolophidae 796 Nomeidae 801 Ariommatidae 810 Tetragonuridae 815 Stromateidae 817 Pleuronectiformes 820 Bothidae 821 Scophthalmidae 830 Paralichthyidae 832 Poecilopsettidae 861

Achiridae 864 Cynoglossidae 872 Tetraodontiformes 887 Triacanthodidae 889 Balistidae 893 Monacanthidae 901 Ostraciidae 913 Tetraodontidae 919 Diodontidae 930 Molidae 939 Glossary 943 References 951 Index of Scientific Names 987 Index of Common Names 997

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ACKNOWLEDGMENTS

A large number of systematic ichthyologists, many curators of natural history museums, and an editor have made major contributions to this work, and their inputs are gratefully acknowledged. Stuart Poss critically read the scorpaeniform section and offered numerous comments. William Richards read the triglid and peristediid sections, provided unpublished manuscripts, and made many valuable suggestions. G. David Johnson and William SmithVaniz offered several corrections to the key to the families of Perciformes. Tom Orrell read the centropomid and inermiid sections and offered advice. Phil Heemstra reviewed most of the serranid section and provided unpublished manuscripts on moronids, acropomatids, and howellids of the western Central Atlantic. Phil Lobel provided input on the Hyploplectrus (Serranidae) species accounts. Offer Gon reviewed the apogonid and epigonid sections. William Anderson, Jr., read the anthiine serranid, lutjanid, and symphysanodontid sections and provided unpublished manuscripts on anthiine serranids and lutjanids of the western Central Atlantic. William Smith-Vaniz read the carangid section and provided an unpublished manuscript on the carangid fishes of the western Central Atlantic. Labbish Ning Chao provided an unpublished manuscript on the sciaenids of the western Central Atlantic. Randy Mooi critiqued the grammatid and pempherid sections. Bruce B. Collette read the pomatomid, coryphaenid, echeneid, rachycentrid, and scombrid sections; furnished data concerning these taxa; and provided unpublished manuscripts on pomatomids, coryphaenids, rachycentrids, and echeneids of the western Central

Atlantic. Richard E. Matheson read and offered clarifications on the centropomid, gerreid, and sciaenid sections. Mark Leiby and Richard Matheson critiqued and provided data for the sparid section. Mark Westneat critiqued the labrid and scarid sections and provided unpublished manuscripts of the labrids and scarids of the western Central Atlantic. Jeffrey T. Williams read and offered substantial improvements to the very long-winded blennioid sections. Karsten Hartel read and commented on the callionymid and draconettid sections. Edward O. Murdy provided keys to the eleotrids and gobiids of the western Central Atlantic and read the long and tedious gobioid sections. Richard Haedrich read and commented on the stromateoid sections. Thomas A. Munroe read the pleuronectiform sections and provided helpful comments. These colleagues have greatly improved the reliability of the nomenclature and taxonomic treatments in the book. Special thanks are given to Nancy Warrington for her excellent editorial assistance on both volumes of this work. Any remaining problems are strictly the fault of the authors. Various colleagues made available lists of species either that they have recorded from the western Central Atlantic or the Gulf of Mexico or that reside in their natural history collections. William Richards and Mark Leiby provided a list of ray-finned fishes occurring between Cape Hatteras, NC, and the equator. John W. Tunnell, Jr., supplied a checklist of fishes inhabiting the coral reefs of the southwestern Gulf of Mexico that he, Mary E. Vega, and John E. Gourley prepared. Karsten Hartel provided a listing of the species of fishes from

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viii ACKNOWLEDGMENTS

the Gulf of Mexico housed at the Museum of Comparative Zoology (MCZ), Cambridge, MA. Mary Ann Rogers furnished a list of the holdings of fishes of the Gulf of Mexico housed at the Field Museum of Natural History (FMNH), Chicago, IL. Rob Robins supplied a list of the fishes of the Gulf of Mexico housed at the Florida Museum of Natural History (FM), Gainesville, FL. Jeffrey T. Williams made available computer records of the species of fishes of the Gulf of Mexico residing at the National Museum of Natural History (USNM), Washington, D.C. Dan Allen made available a video record of Harriota raleighana from off Louisiana. Glen Parsons, G. W. Lingram, Jr., and R. Havard provided the authors with a prepublication copy of their manuscript describing the record capture of Mitsukurina owstoni (Mitsukurinidae) in the Gulf of Mexico. Tracy Ward, Erik Cordes, and Chuck Fisher, Penn State University, provided specimens of Bellottia apoda (Bythitidae) from the Gulf of Mexico. The following curators are thanked for their hospitality during our visits to their collections and for loaning us specimens: Melanie Stiassny, Scott Schaeffer, and Barbara Brown (AMNH); Bernard Kuhajda, Phillip Harris, and Herbert Boschung (UAIC), Barry Chernoff, Mary Ann Rogers, and Kevin Shlagle (FMNH); Richard E. Matheson and Mark Leiby (FBC); George H. Burgess and Rob Robins (FM); Stuart Poss and Sara LeCroy (GCRL); Karsten Hartel and Ann Everly (MCZ); Randy Mooi (MPM); John Lundberg, Dominique Didier, William G. Saul, Mark Sabaj, and Eugenia Böhlke (ANSP); Kathryn Vaughan and Heather Prestridge (TCWC); Dean Henderson (TNHC); Hank Bart and Mike Taylor (TU); Héctor Espinosa Pérez and Leticia Huidobro-Campos (UNAM); Susan Jewett, Lisa Palmer, David Smith, and Jeffrey T. Williams (USNM). Heather Prestridge also provided valuable assistance in elec-

tronically adapting and labeling many of the illustrations. Natural history collection acronyms are according to Leviton et al. (1985). Students of John D. McEachran whose tenure overlapped this project and who contributed to the effort include the following: Jeff Childs, Timothy Clark, Kathy Dunn, and Hera Konstantinou. Neil Ashilman, an undergraduate student, produced the line drawings of Moronidae and Howellidae that accompany the key to the family of percoid fishes. Several individuals were very instrumental in assisting the authors in obtaining funds for publication of the second volume: Rezneat Darnell, Emeritus Professor, Texas A&M University; Benny Galloway, President of LGL Associates, Bryan, TX; James P. Ray, Manager of Environmental Ecology and Response Shell Global Solutions (US), Inc.; John W. Tunnell, Jr., Associate Director, Harte Institute for Gulf of Mexico Studies, TAMU-Corpus Christi; and Alan Verret, Executive Director, Offshore Operators, New Orleans, LA. Permission to reproduce copyrighted figures was provided by: Species Identification and Data Center of the Fisheries, Food, and Agriculture Organization (FAO) of the United Nations; Academy of Natural Sciences, Philadelphia; and the National Museum of Natural History, Smithsonian Institution. Financial support from the National Science Foundation (NSF), Texas A&M University Sea Grant Office, and Coastal Conservation Association (CCA) enabled the completion of the second and final volume of Fishes of the Gulf of Mexico. Without this support, it would not have been possible to complete the illustrations of the fish species, make numerous visits to natural history institutions housing specimens of fishes from the Gulf of Mexico, and purchase the computers and supplies needed to complete the text in a timely fashion.

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Fishes of the Gulf of Mexico Volume 2

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INTRODUCTION

Scope of the Book This is the second and final volume of the Fishes of the Gulf of Mexico, and it covers 693 species in 303 genera, 84 families, and four orders: Scorpaeniformes, Perciformes, Pleuronectiformes, and Tetraodontiformes. The format follows that of the first volume, except descriptions are somewhat expanded for the families of fishes, pertinent references are provided following the family accounts, and the number of species of the family known from the western Central Atlantic (Cape Hatteras, NC, to the equator) are placed between the total known worldwide and those recorded from the Gulf of Mexico. Also, the species accounts are expanded to include more distributional and ecological data, following advice of peers regarding the first volume. As in the first volume, the classification is according to J. S. Nelson (1994), with the following exceptions: Peristediinae is treated as a family, following advice of William Richards. The percoid genera Howella and Symphysanodon are treated as distinct families, Howellidae and Symphysanodontidae, respectively, because neither shares synapomorphies with other percoid taxa (G. David Johnson, pers. comm., July 1999). Nelson (1994) placed Howella in Acropomatidae and Symphysanodon in Lutjanidae. The family Kyphosidae was interpreted according to G. D. Johnson and R. A. Fritzsche (1989), that is, limited to four genera (Hermosilla, Kyphosus, Neoscorpis, and Sector). Poecilopsettinae was also treated as a family rather than as a subfamily of Pleuronectidae, following Chapleau and Keast (1988) and Chapleau (1993). Unlike the first volume, in which fami-

lies were largely arranged alphabetically within orders, in this volume, families within orders are arranged according to Nelson (1994). Families that are not recognized by Nelson immediately follow the family to which they are thought to be most closely related; for example, Symphysanodontidae follows Lutjanidae. The eastern boundary of the Gulf of Mexico is a line connecting Cape Catoche, near the eastern tip of the Yucatán Peninsula, and Key West, Florida (Fig. A). This boundary, which essentially follows that of R. H. Backus and J. E. Craddock (1977), is the same recognized in the first volume, and it thus excludes the Florida Keys and the northwestern coast of Cuba from the Gulf of Mexico. Florida Bay, on the other hand, is considered to be in the Gulf. As was the case in the first volume, the fish species composition of the Gulf of Mexico was based on extensive review of the literature and unpublished species lists, and on surveys of the major natural history collections of the United States and Mexico. An attempt was made to survey all of the major natural history collections with holdings from the Gulf of Mexico, and the following institutions were included: Academy of Natural Sciences, Philadelphia (ANSP), PA; American Museum of Natural History (AMNH), New York, NY; Field Museum of Natural History (FMNH), Chicago, IL; Florida Board of Conservation Fish Collection (FBC), St. Petersburg, FL; Florida Museum of Natural History (FM), Gainesville, FL; Gulf Coast Research Laboratory (GCRL), Ocean Springs, MS; Museum of Comparative Zoology (MCZ), Cambridge, MA; Milwaukee

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2 INTRODUCTION

FIG A

Public Museum (MPM), Milwaukee, WI; National Museum of Natural History (USNM), Washington, D.C.; Texas Cooperative Wildlife Collection (TCWC), College Station, TX; Texas Natural History Collection (TNHC), Austin, TX; Tulane Museum of Natural History (TU), New Orleans, LA; Universidad Nacional Autónoma de México (UNAM), Mexico City; University of Alabama Ichthyological Collection (UAIC), Tuscaloosa, AL; and University of Michigan Museum of Zoology (UMMZ), Ann Arbor, MI. (The above acronyms follow Leviton et al. [1985]). Over the course of the project, one to four weeks were spent at each of these institutions.

How to Identify Fishes The fishes included in the second volume are more morphologically similar in shape than those covered in the first volume but still display considerable variation in shape and size.

The same methods used in the first volume apply for making measurements and counts, but some of these are further elaborated. Proportional measurements are based on standard length (SL), head length (HL), and occasionally on fork length (FL). Standard length is the distance from the tip of the snout to the end of the hypural plate and is determined by bending the tail against a flat surface and measuring to the seam representing the junction between the hypural plate and the fin rays. The hypural plate forms the surface onto which the caudal fin rays attach proximally. Head length is the straight-line distance between the tip of the snout and the opercular flap (Figs. B and C). Fork length is used in fishes such as carangids that have bony scutes running along the posterior section of the lateral line to the base of the caudal fin. The scutes make it difficult to accurately determine the end of the hypural plate. For this reason, proportional lengths are expressed in percentage of fork length rather

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2 INTRODUCTION

FIG A

Public Museum (MPM), Milwaukee, WI; National Museum of Natural History (USNM), Washington, D.C.; Texas Cooperative Wildlife Collection (TCWC), College Station, TX; Texas Natural History Collection (TNHC), Austin, TX; Tulane Museum of Natural History (TU), New Orleans, LA; Universidad Nacional Autónoma de México (UNAM), Mexico City; University of Alabama Ichthyological Collection (UAIC), Tuscaloosa, AL; and University of Michigan Museum of Zoology (UMMZ), Ann Arbor, MI. (The above acronyms follow Leviton et al. [1985]). Over the course of the project, one to four weeks were spent at each of these institutions.

How to Identify Fishes The fishes included in the second volume are more morphologically similar in shape than those covered in the first volume but still display considerable variation in shape and size.

The same methods used in the first volume apply for making measurements and counts, but some of these are further elaborated. Proportional measurements are based on standard length (SL), head length (HL), and occasionally on fork length (FL). Standard length is the distance from the tip of the snout to the end of the hypural plate and is determined by bending the tail against a flat surface and measuring to the seam representing the junction between the hypural plate and the fin rays. The hypural plate forms the surface onto which the caudal fin rays attach proximally. Head length is the straight-line distance between the tip of the snout and the opercular flap (Figs. B and C). Fork length is used in fishes such as carangids that have bony scutes running along the posterior section of the lateral line to the base of the caudal fin. The scutes make it difficult to accurately determine the end of the hypural plate. For this reason, proportional lengths are expressed in percentage of fork length rather

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INTRODUCTION 3

FIG B

FIG C

than in percentage of standard length. Fork length is measured from the tip of the snout to the midlateral margin of the caudal fin when the fin is expanded dorsoventrally. All measurements are made from point to point. Many of the fishes included in the second volume have the ultimate dorsal fin ray and anal fin divided to the base, thus care must

be taken if this is the case to count the two sections of the ray as one ray. The gill raker counts on the first gill arch in the text include all elements, even the rudiments, unless otherwise stated in the text. Rudiments are generally considered elements that are wider than they are tall. Pectoral fin counts given in the text refer to all elements, whether branched or

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4 INTRODUCTION

unbranched, paired or single. In some literature sources, especially older references, the short unbranched, unpaired elements are given as lowercase Roman numerals. Lateral line scale counts are inclusive of pored, tubed, or grooved scales from above the operculum to the end of the caudal peduncle. In some accounts, lateral line scales posterior to the caudal peduncle are given separately: 35+1+1, as for the scorpaenids. Horizontal scale series are those between the upper free margin of the operculum and the caudal peduncle. The scales between the origin of the dorsal fin and the lateral line are counted diagonally and posteriorly from the base of the first dorsal fin spine (ray) to the lateral line. The scales between the lateral line and the anal fin origin are counted diagonally and anteriorly from the origin of the anal fin to the lateral line.

Biogeography of Fishes of the Gulf of Mexico Species Richness and Composition and Biogeographical Affinities Based on distributional patterns of continental shelf species, the Gulf of Mexico is considered both a warm temperate and a tropical body of water (Briggs 1974). Its northern section, Cape Romano, FL, to Cape Rojo, Veracruz (see Fig. A), is warm temperate and shares faunal elements with the remainder of the Carolinian province (Cape Hatteras, NC, to Cape Canaveral, FL). Its southern sections— Cape Rojo to Cape Catoche, Yucatán, and the west coast of Florida south of Cape Romano—are tropical and share faunal elements with the Caribbean province. Based on the distribution of mesopelagic fishes, such as myctophids, the Gulf of Mexico is considered distinct but most similar to the Atlantic Tropical Region (Backus et al. 1977). J. V. Gartner et al. (1987) and C. Karnella (1987) also noted similarities as well as some differences in species composition of myctophids between the eastern Gulf of Mexico and the tropical Atlantic. According to Backus et al. (1977), the Gulf under-

goes significant winter cooling and, possibly because of this cooling, also shares species with the Atlantic Tropical-Subtropical, the Atlantic Tropical-Semitropical, and the Atlantic Temperate-Semitropical regions. Unlike J. C. Briggs (1974), Backus et al. (1977) considered the Gulf to be undivided and to constitute both a biogeographical region and a province, although no myctophid species are endemic to the Gulf of Mexico. R. H. Gibbs et al. (1983) and T. T. Sutton and T. L. Hopkins (1996) found the stomiid mesopelagic assemblage (Astronesthidae, Chauliodontidae, Idiacanthidae, Malacosteidae, Melasnostomiidae, and Stomiidae) of the Gulf of Mexico to be significantly different from that of the tropical Atlantic in both species richness and composition. According to all of the above interpretations, the Gulf of Mexico shares tropical and warm temperate species with the Atlantic Ocean and is most similar in species composition to the western Central Atlantic region (Cape Hatteras, NC, to the equator). The present two-volume study documents 1,443 fish species in 700 genera, 223 families, and 45 orders in the Gulf of Mexico (767 species in 396 genera, 137 families, and 40 orders in volume 1; 656 species in 296 genera, 83 families, and 4 orders in this volume; and 24 additional species in 8 genera, 3 families, and 1 order and 3 species and 1 genus deletions to accounts included in volume 1). Another 49 species in 11 additional genera and 2 families are included because of the likelihood that they occur but to date have not been documented from the Gulf. Based on several published and unpublished studies (Rosen 1973; Compagno 1984; W. J. Richards and M. Leiby, pers. comm., December 2001; McEachran and de Carvalho 2002), there are about 2,247 species of fishes in 858 genera, 241 families, and 45 orders in the western Central Atlantic. Thus 64.2 percent of the species, 81.6 percent of the genera, 92.5 percent of the families, and all of the orders of fishes recorded from the western Central Atlantic have been documented from the Gulf of Mexico. Families represented in this area but not

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4 INTRODUCTION

unbranched, paired or single. In some literature sources, especially older references, the short unbranched, unpaired elements are given as lowercase Roman numerals. Lateral line scale counts are inclusive of pored, tubed, or grooved scales from above the operculum to the end of the caudal peduncle. In some accounts, lateral line scales posterior to the caudal peduncle are given separately: 35+1+1, as for the scorpaenids. Horizontal scale series are those between the upper free margin of the operculum and the caudal peduncle. The scales between the origin of the dorsal fin and the lateral line are counted diagonally and posteriorly from the base of the first dorsal fin spine (ray) to the lateral line. The scales between the lateral line and the anal fin origin are counted diagonally and anteriorly from the origin of the anal fin to the lateral line.

Biogeography of Fishes of the Gulf of Mexico Species Richness and Composition and Biogeographical Affinities Based on distributional patterns of continental shelf species, the Gulf of Mexico is considered both a warm temperate and a tropical body of water (Briggs 1974). Its northern section, Cape Romano, FL, to Cape Rojo, Veracruz (see Fig. A), is warm temperate and shares faunal elements with the remainder of the Carolinian province (Cape Hatteras, NC, to Cape Canaveral, FL). Its southern sections— Cape Rojo to Cape Catoche, Yucatán, and the west coast of Florida south of Cape Romano—are tropical and share faunal elements with the Caribbean province. Based on the distribution of mesopelagic fishes, such as myctophids, the Gulf of Mexico is considered distinct but most similar to the Atlantic Tropical Region (Backus et al. 1977). J. V. Gartner et al. (1987) and C. Karnella (1987) also noted similarities as well as some differences in species composition of myctophids between the eastern Gulf of Mexico and the tropical Atlantic. According to Backus et al. (1977), the Gulf under-

goes significant winter cooling and, possibly because of this cooling, also shares species with the Atlantic Tropical-Subtropical, the Atlantic Tropical-Semitropical, and the Atlantic Temperate-Semitropical regions. Unlike J. C. Briggs (1974), Backus et al. (1977) considered the Gulf to be undivided and to constitute both a biogeographical region and a province, although no myctophid species are endemic to the Gulf of Mexico. R. H. Gibbs et al. (1983) and T. T. Sutton and T. L. Hopkins (1996) found the stomiid mesopelagic assemblage (Astronesthidae, Chauliodontidae, Idiacanthidae, Malacosteidae, Melasnostomiidae, and Stomiidae) of the Gulf of Mexico to be significantly different from that of the tropical Atlantic in both species richness and composition. According to all of the above interpretations, the Gulf of Mexico shares tropical and warm temperate species with the Atlantic Ocean and is most similar in species composition to the western Central Atlantic region (Cape Hatteras, NC, to the equator). The present two-volume study documents 1,443 fish species in 700 genera, 223 families, and 45 orders in the Gulf of Mexico (767 species in 396 genera, 137 families, and 40 orders in volume 1; 656 species in 296 genera, 83 families, and 4 orders in this volume; and 24 additional species in 8 genera, 3 families, and 1 order and 3 species and 1 genus deletions to accounts included in volume 1). Another 49 species in 11 additional genera and 2 families are included because of the likelihood that they occur but to date have not been documented from the Gulf. Based on several published and unpublished studies (Rosen 1973; Compagno 1984; W. J. Richards and M. Leiby, pers. comm., December 2001; McEachran and de Carvalho 2002), there are about 2,247 species of fishes in 858 genera, 241 families, and 45 orders in the western Central Atlantic. Thus 64.2 percent of the species, 81.6 percent of the genera, 92.5 percent of the families, and all of the orders of fishes recorded from the western Central Atlantic have been documented from the Gulf of Mexico. Families represented in this area but not

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INTRODUCTION 5

represented in the Gulf are mostly deep-sea and oceanic fishes and, to a lesser extent, temperate fishes rare south of Cape Hatteras or tropical fishes rare north of Central America or west of the Bahamas or Greater Antilles. Deep-sea families present in the western Central Atlantic but not in the Gulf of Mexico include Heterenchelyidae, Derichthyidae, Muraenesocidae, Cyematidae, Saccopharyngidae, Monognathidae, Radiicephalidae, Parabrotulidae, Caulophrynidae, Neoceratiidae, Gigantactinidae, Megalomycteridae, and Oreosomatidae (W. J. Richards and M. Leiby, pers. comm., December 2001). The sill depths of the Straits of Yucatán (2,040 m) and the Straits of Florida (800 m) and, more importantly, the deepest sill depths between the Caribbean Sea and the Atlantic Ocean, which are the Windward (1,540 m), the Jungfern (1,815 m), and the Anegada (1,910) Passages, may exclude some of these families. Temperate fishes known from the western Central Atlantic but not from the Gulf of Mexico include Anotopteridae, Psychrolutidae, Cyclopteridae, Ammodytidae, and Scophthalmidae (W. J. Richards and M. Leiby, pers. comm., December 2001). These families are rare south of Cape Hatteras. Tropical families known from the Caribbean Sea but not from the Gulf of Mexico include Anomalopidae and Synbranchidae. There is only one species of anomalopid in the western Atlantic, and most species of synbranchids are limited to freshwater or brackish water. Most of the species (in families represented in the Gulf) occurring in the western Central Atlantic but not in the Gulf of Mexico are distributed either south of the Gulf of Mexico or in the insular regions to the east (the Bahamas, the Greater Antilles, and the Lesser Antilles). These species may be excluded by a variety of factors, including environmental parameters, habitat availability, and dispersal abilities of the fishes. The Gulf of Mexico has greater seasonal temperature changes than regions to the south or east (Backus et al. 1977), and this greater variation may exclude some stenother-

mal species. The Gulf of Mexico lacks extensive reef habitats, unlike the Caribbean coast of Central and South America, and the Bahamas and the Antilles, and this deficiency may exclude some reef-dependent species. Secretive reef-dwelling or reef-associated species of the western Central Atlantic, such as Grammatidae (with 2 of 11 species in the Gulf), Opistognathidae (with 7 of 17 species), Dactyloscopidae (with 4 of 17 species), Labrisomidae (with 19 of 41 species), Chaenopsidae (with 11 of 40 species), Gobiesocidae (with 2 of 25 species), and Gobiidae (with 58 of 124 species), are relatively less diversely represented and/or widely distributed in the Gulf than the larger and more conspicuous reef dwellers, such as Chaetodontidae (with 6 of 8 species in the Gulf), Labridae (with 17 of 19 species), Scaridae (with 14 of 14 species), Acanthuridae (with 3 of 3 species), and Balistidae (with 6 of 6 species). Pomacentridae (with 14 of 20 species in the Gulf) often occupy small ranges on coral reefs, but they have pelagic eggs and larvae that widely disperse, unlike many of the secretive reef dwellers that either have benthic eggs or are mouth brooders. Many species of Engraulidae (with 8 of 29 species in the Gulf), Ariidae (with 6 of 21 species), Batrachoididae (with 6 of 20 species), and Sciaenidae (with 25 of 59 species) are distributed along the continental coast of the Caribbean, from northern South America to Honduras or Belize, and may be limited by relatively greater seasonal temperature changes in the Gulf than in the areas of the Caribbean. Clupeidae (with 16 of 21 species in the Gulf), Macrouridae (with 32 of 35 species), Holocentridae (with 11 of 11 species), Triglidae (with 14 of 19 species), Serranidae (with 61 of 89 species), Carangidae (with 28 of 32 species), Lutjanidae (with 16 of 18 species), Gerreidae (with 12 of 12 species), Haemulidae (with 18 of 23 species), and Sparidae (with 16 of 19 species) are well represented in the Gulf. These taxa generally are wideranging pelagic or demersal /benthic species, occur on a variety of substrates, or are common on soft bottoms that are widely distributed in the Gulf.

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6 INTRODUCTION

Alternatively, historical factors may be responsible for the absence of species from the continental shelves and slopes of Central and South America in the Gulf of Mexico. Robins (1971) stated that shore fishes of the western central Atlantic are divided into three assemblages: a northern continental assemblage, a central insular assemblage, and a southern continental assemblage. Continental assemblages are typical of continental shelves where water temperatures, salinities, and turbidity levels vary with terrestrial weather patterns and terrigenous substrates predominate. Many species of the continental assemblage are associated with estuaries and areas where rivers meet the coast. Insular assemblages are typical of insular shelves where oceanic water masses buffer the aquatic habitat from terrestrial weather patterns, turbidity is low, and carbonate substrates predominate. Many species of the insular assemblage are associated with coral reefs. The northern continental assemblage extends from Cape Canaveral on the east coast of Florida, around the tip of Florida, and westward throughout the Gulf of Mexico. The southern continental assemblage extends from northern Brazil to Nicaragua. The insular assemblage extends throughout the Bahamas, the Greater and Lesser Antilles, and barrier reefs off Belize and Quintana Roo. Components of the northern and southern continental assemblages are interspersed with the insular assemblage around Cuba, Hispaniola, and Jamaica in areas where rivers discharge into the sea and terrigenous substrates predominate. Correspondingly, components of the insular assemblage are interspersed with the northern and southern continental assemblages along continental shelves where hard substrates predominate and river mouths are lacking. A recent biogeographical study of the fishes of the western Central Atlantic offers support for Robins’ categorization of the region and suggests that faunal differences between the northern and southern continental shelf assemblages are historical (Smith et al. 2002). Smith et al. (2002) mapped the distribution of 987 fish species occurring in the western

Central Atlantic and found areas of highest diversity in waters around southern Florida, the eastern Bahamas, northern Cuba, and to a slightly lesser degree, continental waters of northern South America, Central America, and the northern Gulf of Mexico. They found the highest levels of endemism to be in southern Florida, the coast of South America off Venezuela and Colombia, and the northern Gulf of Mexico. The fishes around southern Florida, eastern Bahamas, and northern Cuba are mostly components of Robins’ insular fish assemblage, and the fishes from northern South America, Central America, and the northern Gulf of Mexico are components of Robins’ southern and northern continental shelf assemblages. Smith et al. (2002) also found that the continental shelf fishes could be placed in four categories: ubiquitous species (found throughout the area; 10 species), species restricted to very small regions of the area (86 species), species that enter the area from the east coast of North America (northern component comprising 49 species), and species that enter the area from the east coast of South America (southern component comprising 67 species). The latter two categories are biogeographically informative. Members of the first category are universally present, thus they do not provide any information. Members of the second category occupy very small ranges that in composite do not reflect obvious vicariant patterns. Only 18 percent of the northern component extends south of Cape Catoche, and only 19 percent of the southern assemblage extends north of Nicaragua. Thus the third and the fourth categories support the presence of two distinct continental faunal assemblages: a warm temperate one that enters the area from the north, and a tropical component that enters the area from the south. These two components represent Robins’ northern and southern continental assemblages, respectively. This pattern suggests that the northern and southern continental assemblages had separate evolutionary patterns, and this hypothesis is supported by tectonic history of the western Central Atlantic. An inter-American sea-

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INTRODUCTION 7

way existed between North and South America for much of the history of the Caribbean Basin (Smith et al. 2002). The seaway opened with the breakup of Pangaea and Gondwanaland in the early Mesozoic and late Mesozoic (see “Geological History of the Gulf of Mexico” in volume 1: 5 – 6). The seaway was open at all depths until the mid Miocene, thus until the mid Miocene it probably served as a barrier to continental shelf and coastal fishes on either side of the seaway. The seaway was partially closed by submarine ridges between the mid and late Miocene and was completely closed by the Panamanian Isthmus in the late Pliocene (2.76 to 2.51 Ma). Thus the deepsea barrier between South America and North America ended between the mid to late Miocene and the late Pliocene, depending on the depth ranges of the fish species on either side. These geological events may explain many of the faunal differences between the two continental shelf assemblages in the western Central Atlantic and why a number of fish species abundant on the Caribbean continental shelf are lacking from the Gulf of Mexico. Considering its size, the Gulf of Mexico has a relatively rich fish fauna (Table 1). The Gulf has about 200 more species and 54 more genera than occur in the eastern Atlantic Ocean between the Arctic and the southern coast of

Morocco, including the Mediterranean (Whitehead et al. 1984 –1986), and within 129 species and 58 genera of the number that occur in the eastern Atlantic from the coast of Morocco to southern Namibia (Quéro et al. 1990). Nearly 10 percent of the world’s known marine fish species occur in the Gulf of Mexico (Nelson 1994).

Endemicity The uniqueness of the Gulf of Mexico stems more from the richness of its fish fauna than from a high percentage of endemic species. Only 66 species of the total 1,443 species (4.6%) can be considered endemic to the Gulf of Mexico (Table 2). Endemicity as used here is liberally defined: species are considered to be endemic if they are largely limited to the Gulf, even if they also occur along the southeastern coast of Florida, the eastern coast of Yucatán, or the Florida Keys. Excluding species along the east coasts of Florida and Yucatán would greatly reduce the percentage of endemic species. Also, Dorosoma petenense is considered endemic despite the fact that only adults are found in brackish to low-salinity waters of the Gulf, with younger life stages limited to freshwater. The majority of the endemic species are limited to one or two of the three subregions

TABLE 1. Comparison of taxonomic richness of fishes in the Gulf of Mexico with that of other regions Number of Species Gulf of Mexico Eastern North Atlantic and Mediterranean 1 Eastern Central Atlantic 2 Eastern South Atlantic and Southwestern Indian Ocean 3 Japanese Archipelago 4 World 5 1 Data

from Whitehead et al. 1984 –1986. 2 Data from Quéro et al. 1990. 3 Data from Smith and Heemstra 1986.

Number of Genera

Number of Families

1,443 1,248 1,572

700 645 757

223 221 236

2,200 3,000 14,652

1,060 1,267 2,939

270 297 389

4 Data 5 Data

from Masuda et al. 1984. from Nelson 1994.

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TABLE 2.

Fish species endemic to the Gulf of Mexico Region

Family Myxinidae Myxinidae Scyliorhinidae Triakidae Rajidae Rajidae Rajidae Rajidae Rajidae Rajidae Ophichthidae Ophichthidae Ophichthidae Ophichthidae Ophichthidae Congridae Clupeidae Clupeidae Clupeidae Clupeidae Clupeidae Melanostomiidae Paralepididae Paralepididae Bythitidae Bythitidae Macrouridae Macrouridae Batrachoididae Batrachoididae Ogcocephalidae Ogcocephalidae Oneirodidae Atherinidae Cyprinodontidae Cyprinodontidae Cyprinodontidae Fundulidae Fundulidae Fundulidae Fundulidae Fundulidae Poeciliidae

Species Eptatretus minor Eptatretus springeri Parmaturus campechiensis Mustelus sinusmexicanus Anacanthobatis folirostris Dipturus olseni Dipturus oregoni Leucoraja lentiginosa Raja ackleyi Raja texana Gordiichthys ergodes Gordiichthys leibyi Ophichthus omorgmus Ophichthus rex Pseudomyrophis nimius Heteroconger luteolus Alosa alabamae Alosa chrysochloris Brevoortia gunteri Brevoortia patronus Dorosoma petenense Eustomias leptobolus Stemonosudis bullisi Uncisudis advena Gunterichthys longipenis Saccogaster rhamphidognatha Coryphaenoides mexicanus Kumba species Opsanus pardus Sanopus reticulatus Ogcocephalus declivirostris Ogcocephalus pantostictus Oneirodes bradburyae Menidia peninsulae Floridichthys carpio Jordanella floridae Jordanella pulchra Adinia xenica Fundulus grandissimus Fundulus jenkinsi Fundulus pulvereus Fundulus seminolis Gambusia affinis

East

NW

South

1 1 0 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 0 1 0 1 1 1 1 0 1 0 1 1 0 0 0 1 1 1 0 1 0 1 0 1 0

1 1 0 1 1 1 1 1 0 1 1 0 0 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 0 1 1 1 1 0 0 0 1 0 1 1 0 1

0 0 1 0 0 0 0 1 1 1 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 0 0 1 0 1 1 0 1 1 0 1 0 1 0 1 0 0

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INTRODUCTION 9

TABLE 2.

(Continued) Region

Family Syngnathidae Triglidae Triglidae Triglidae Sciaenidae Chaenopsidae Blenniidae Blenniidae Blenniidae Blenniidae Blenniidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Gobiidae Microdesmidae Paralichthyidae Achiridae Tetraodontidae

Species

East

NW

South

Syngnathus affinis Prionotus longispinosus Prionotus martis Prionotus paralatus Cynoscion arenarius Emblemariopsis diaphana Chasmodes longimaxilla Chasmodes saburrae Hypleurochilus caudovittatus Hypleurochilus multifilis Lupinoblennius nicholsi Bollmannia communis Bollmannia eigenmanni Chriolepis benthonis Chriolepis vespa Coryphopterus punctipectophorus Ctenogobius claytonii Gobiosoma longipala Varicus marilynae Microdesmus lanceolatus Citharichthys abbotti Gymnachirus texae Sphoeroides parvus

0 1 1 0 1 1 0 1 1 0 0 1 1 0 1 1 0 1 1 0 0 0 0

1 1 1 1 1 0 1 0 0 1 0 1 0 0 0 0 1 1 0 1 0 1 1

1 1 1 1 1 0 1 0 0 1 1 1 0 1 0 1 1 0 0 1 1 1 1

(eastern, northwestern, or southern, described below) and display several distributional patterns. Only 9 endemic species occur throughout the Gulf. Eleven species endemic to the northern or western Gulf have either a sister species or a phenotypically similar species that occurs along the eastern seaboard of the United States (e.g., Leucoraja lentiginosa, Brevoortia gunteri, B. patronis, Alosa alabamae, A. chrysochloris, Prionotus martis, Cynoscion arenarius, Chasmodes longimaxilla, Peprilus burti, Gymnarchirus texae, and Sphoeroides parvus). The east-coast putative sisters of these species are Leucoraja garmani, Brevoortia smithi, B. tyrannus, Alosa sapidissima, A. mediocris, Prionotus carolinus, Cynoscion regalis, Chasmodes bosquianus, Peprilus triacanthus, Gymnachirus melas, and Sphoeroides maculatus,

respectively. The emergence of the Florida Peninsula between the end of the Pliocene and the middle of the Pleistocene (Bowen and Avis 1990) may have been the vicariant event leading to speciation in these cases. Five species that are widely distributed along the eastern coast of the United States, Ascipenser oxyrhynchus (Ascipenseridae), Morone saxatilis (Moronidae), Centropristis striata (Serranidae), Larimus fasciatus (Sciaenidae), Etropus cyclosquamus (Paralichthyidae), have or had isolated populations in the northern Gulf of Mexico, suggesting that the Gulf populations are or were remnants of western extremes of once continuous populations (Smith et al. 2002). Apparently the time interval of 300,000 to 1,000,000 years since the emergence of the Florida Peninsula was insufficient for the Gulf

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10 INTRODUCTION

and east-coast populations of these two species to have formed separate species (Bowen and Avis 1990; Galloway et al. 1991). The Gulf population of Morone saxatilis may no longer exist because of the introduction of specimens of the east-coast population into the Gulf that may have swamped their genome. Two of the northern and western Gulf endemics with sister species in the Atlantic, Chasmodes longimaxilla and Sphoeroides parvus, have another congener (C. saburrae and S. nephelus, respectively) distributed in the eastern Gulf of Mexico and along the Florida Peninsula (Springer 1959; Shipp and Yerger 1969a,b; Williams 1983). In both cases, the peninsular species is likely the sister of the species pair allopatrically distributed along the east coast of the United States and in the northern or western Gulf of Mexico. Florida consisted of one or more islands during the Miocene (Riggs 1984; Galloway et al. 1991; Riggs et al. 1991; Salvador 1991), and the peninsular form may have undergone speciation from the ancestral species of the present-day east-coast and Gulf species during this time. The relationships of the other 55 endemic species are not sufficiently known to comment on their evolutionary patterns. Some of these 55 species are known from very few specimens (e.g., Paramaturus campechiensis, Raja ackleyi, Eustomias leptobolus, Stemonosudis bullisi, Uncisudis advena, and Oreirodes bradburyae), and many or all may ultimately be known from areas outside the Gulf.

The Gulf of Mexico as a BioGeographical Region According to Briggs (1974), a cohesive biogeographic region or biogeographic province is defined by the percentage of endemic species. Areas have to have at least 10 percent endemic species to be considered provinces. Based on endemicity, the Gulf of Mexico cannot be considered a province. However, based on its high diversity of fish species and its unique warm temperate and tropical components of species, the Gulf of Mexico is a unique biogeographical region. Its uniqueness may, in part, be

due to a combination of physical factors. Its mouth straddles the Tropic of Cancer, and it has rather deep sills in the Straits of Yucatán (2,040 m) and in the Straits of Florida (800 m), thus it is accessible to warm temperate and tropical shore fishes and to most deep-sea pelagic and benthic fishes. It is surrounded on three sides by large landmasses, and the proximity of land may be responsible for the significant winter cooling of the water column, compared to the Tropical Atlantic region. Furthermore, the eastern Gulf of Mexico is dominated by the Florida Loop Current, which enters the Gulf through the Yucatán Straits, extends variably westward and occasionally to the mouth of the Mississippi River, and exits through the Florida Straits (McEachran and Fechhelm 1998 and references therein). The current is a regular source of tropical pelagic fauna for the Gulf. Warm-core rings that spin off the Florida Loop Current can transport pelagic organisms into the western Gulf of Mexico.

Distributional Patterns The fishes of the Gulf of Mexico display a number of distributional patterns (see Table 3) but support the conclusions of Briggs (1974) and Backus et al. (1977) that the species assemblages of the Gulf include both warm temperate and tropical components. The Gulf of Mexico shares 812 species with the southeastern United States (Cape Hatteras to Cape Canaveral), and many of these species are in the families Triglidae, Sparidae, Sciaenidae, Blenniidae, and Paralichthyidae. The Gulf shares 1,037 species with the Caribbean coast of Central and South America, and many of these fishes are in the families Ophichthidae, Congridae, Scorpaenidae, Centropomidae, Pomacentridae, Labridae, Scaridae, Labrisomidae, Chaenopsidae, and Bothidae. Gulf of Mexico species of Serranidae, Carangidae, and Lutjanidae appear to be more or less equally distributed along the Atlantic coasts north and south of the Gulf of Mexico. The Gulf shares 563 species with the Bahamas, 848 species

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INTRODUCTION 11

with the Greater Antilles, and 770 species with the Lesser Antilles. The majority of these fishes are wide ranging throughout the western Central Atlantic (Scorpaenidae, Serranidae, Apogonidae, Carangidae, Lutjanidae, Haemulidae, Chaetodontidae, Pomacanthidae, Pomacentridae, Labridae, Scaridae, Balistidae, Monacanthidae, Ostraciidae, Tetraodontidae, and Diodontidae). Other species, representing Tripterygiidae, Chaenopsidae, Labrisomidae, and Gobiidae, are more restricted in their distributions and mostly limited to islands and barrier-reef habitats of the Caribbean Sea. These reef-associated species are generally best represented in the eastern and southern Gulf of Mexico. About half (48.8%) of the 1,443 species occur throughout the Gulf and may, in a broad sense, be considered ubiquitous within their respective depth ranges. A majority of the ubiquitous species are wide-ranging epipelagic fishes (Carcharhinidae, Clupeidae, Belonidae, Exocoetidae, Hemiramphidae, Carangidae, and Scombridae), wide-ranging mesopelagic fishes (Gonostomatidae, Sternoptychidae, Phosichthyidae, and Myctophidae), wide-ranging benthic fishes of the continental shelf (Holocentridae, Serranidae, Lutjanidae, and Sciaenidae), and benthic slope or rise fishes (Ipnopidae, Macrouridae, and Moridae). The ubiquitous species are also widely distributed in other regions of the western Central Atlantic and, in some cases, in other regions of the Atlantic, Pacific, and Indian Oceans. The remaining 51.2 percent of the 1,443 species are restricted to subregions of the Gulf, such as the eastern subregion (Dry Tortugas and Florida Bay to Pensacola, FL, or Mobile Bay, AL), the northwestern subregion (Pensacola, FL, or Mobile, AL, to Cape Rojo, Veracruz), the southern subregion (Cape Rojo to Cape Catoche, Yucatán), or to combinations of these subregions. A total of 14.6 percent (211 species) occur only in the eastern subregion. A large percentage of these species are in the families Ophichthidae, Alepocephalidae, Melanostomiidae, Notosudidae, Paralepididae, Syngnathidae, Opistognathidae, Apogo-

nidae, Chaenopsidae, and Gobiidae. The high diversity of mesopelagic fishes in the eastern subregion may reflect sampling bias or intrusion of the Loop Current into the eastern Gulf of Mexico (Sutton and Hopkins 1996). Thomas Hopkins, of the University of South Florida, has maintained a meso-bathypelagic sampling station on the west coast of Florida for 19 years (Sutton and Hopkins 1996), and this water column by far is the best sampled in the Gulf of Mexico. The high diversity of reefassociated fishes (Opistognathidae, Apogonidae, Chaenopsidae, and Gobiidae) is probably related to the predominance of calcareous substrata along the west coast of Florida and the proximity to the Florida Keys. A number of reef-associated species have been reported only from the Dry Tortugas (Longley and Hildebrand 1941). About 3.6 percent of the Gulf species are limited to the northwestern subsection. A number of these species are either endemic to the Gulf of Mexico or also occur along the southeastern United States. Others such as Harriotta raleighana, Odontapsis noronhai, Aplatophis chauliodus, Labichthys carinatus, Asquamiceps caeruleus, Talismania antillarum, Brotulotaenia brevicauda, Bellotia atra, Kuma sp., Oneirodes bradburyae, Chiasmodon subniger, Pseudoscopelus obtusifrons, and P. scriptus are deep-sea fishes known from relatively few specimens, and they may turn up in other subregions of the Gulf. About 6.4 percent of the species are limited to the southern Gulf of Mexico, and the majority of these species also occur in the Caribbean Sea. Very few species that are restricted to the southern subregion are endemic to the Gulf of Mexico, suggesting that, unlike the Florida Peninsula, the Yucatán Peninsula has not served as a biogeographic barrier (Smith et al. 2002). About 17.5 percent of the species are restricted to the northern Gulf (eastern and northwestern subregions), and the majority of these species are also distributed along the eastern seaboard of the United States. About 3 percent of the species are restricted to the western Gulf (northwestern and southern subregions), and some of these species have disjunct populations along

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12 INTRODUCTION

the eastern seaboard of the United States. About 5.3 percent of the species are restricted to the eastern and southern Gulf of Mexico, and many of these species are associated with reefs and also occur in the Florida Keys, the Bahamas, and the Greater and Lesser Antilles. All of these distributional patterns appear to reflect geological and oceanographic conditions in combination with habitat factors. Benthic species associated with terrigenous substrates occur in the northern and western sections of the Gulf, and benthic fishes associated with calcareous substrates are associated with the calcareous shelves of Florida and Yucatán. Species with warm temperate affinities are generally found in the northern Gulf, and those with tropical affinities are generally found in the southern Gulf. The highest diversity of species (1,259) occurs in the eastern Gulf (Florida Bay to Pensacola or Mobile). The western Gulf (Pensacola or Mobile to Cape Rojo) has slightly fewer species (1,056), and the southern Gulf (Cape Rojo to Cape Catoche) has the lowest number of species (916). These differences may partially reflect sampling or survey intensity, since the northern Gulf has been more intensely sampled than has the southern Gulf. Most of the deepwatercolumn surveys have taken place off northern Florida, and this may account for the larger number of midwater fishes occurring in the eastern Gulf of Mexico. However, it is possible that a number of the midwater fishes captured in the eastern Gulf were collected in the Loop Current, and thus they may not be part of the midwater fauna of the Gulf of Mexico. The Gulf of Mexico shares 512 species with the eastern Central Atlantic, 364 species with the eastern North Atlantic, and 308 species with South Africa. The majority of the amphiAtlantic fishes are neritic coastal species; epipelagic, mesopelagic, or bathypelagic oceanic species; or deep-sea benthic species, including members of Echeneidae, Exocoetidae, Coryphaenidae, and Carangidae (neritic coastal fishes); Scombridae, Xiphiidae, Nomeidae, Ariommatidae, and Molidae (epipelagic fishes); and Stomiiformes, Aulopiformes, Myctophi-

dae, Ceratioidei, Bramidae, Chiasmodontidae, and Gempylidae (meso- and bathypelagic fishes). Benthic continental-shelf species make up about 16 percent of the amphi-Atlantic species and include several species each from the following families: Muraenidae, Congridae, Serranidae, Lutjanidae, and Labridae, and a single species each from the following families: Ophichthidae, Gerreidae, Sparidae, Pomacanthidae, Scaridae, and Bothidae. The majority of these demersal species appear to have achieved their amphi-Atlantic distributions by chance dispersal of eggs and larvae. All of the species have pelagic eggs and larvae that apparently can occasionally survive transatlantic dispersal events. The relatively high percentages of amphi-Atlantic species of Muraenidae, Congridae, and Acanthuridae may be due to their long-lived pelagic larval stages (Leis and Rennis 1983). The high percentage of amphiAtlantic species of Balistidae, Monacanthidae, Tetraodontidae, and Diodontidae may be due to their extended pelagic juvenile stages (Leis and Rennis 1983; Aboussouan and Leis 1984; Leis 1984) and their association with pelagic sargassum (Dooley 1972). Sargassum passively drifts around the North and South Atlantic water gyres, and a diverse fauna of larval and juvenile fishes is associated with the sargassum. The Gulf of Mexico shares 146 species with the eastern Central Pacific and 82 species with the eastern North Pacific. The majority of these are epipelagic, mesopelagic, and bathypelagic oceanic species that are circumtropical. A few are wide-ranging neritic species, such as Coryphaena equisetus, C. hipurus, and Pomatomus saltatrix, and wide-ranging demersal fishes with juvenile stages that are associated with sargassum. The Gulf of Mexico shares 463 (62%) genera and 190 families (85%) with the eastern Central Atlantic, 358 (51%) genera and 172 (77%) families with the eastern North Atlantic, and 421 (60%) genera and 200 (90%) families with South Africa (eastern South Atlantic/southwestern Indian Ocean; Table 3). Most of these genera and families consist of neritic coastal species; epipelagic, meso-

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INTRODUCTION 13

TABLE 3. Number and percentage of species, genera, and families of fishes of the Gulf of Mexico shared with other regions of the Atlantic and eastern Pacific

Gulf of Mexico Florida Keys Bahamas 1 Greater Antilles Lesser Antilles Southeastern United States Central America Bermuda 2 Eastern Central Atlantic 3 Eastern North Atlantic 4 Eastern South Atlantic/ Southwestern Indian Ocean 5 Eastern Central Pacific 6 Eastern North Pacific 7

Number of Species/%

Number of Genera/%

Number of Families/%

1,443/100% 830/57.5% 563/39.0% 848/58.8% 770/53.4% 812/56.3% 1,037/71.9% 460/31.9% 512/35.5% 364/25.2%

700/100% 411/58.7% 308/44.0% 421/60.1% 397/56.7% 486/69.4% 554/79.1% 291/41.6% 463/66.1% 358/51.1%

223/100% 151/67.7% 139/62.3% 188/84.3% 164/73.5% 195/87.4% 190/85.2% 128/57.4% 190/85.2% 172/77.1%

308/21.3% 146/10.1% 82/05.7%

421/60.1% 390/55.7% 242/34.6%

200/89.7% 180/80.7% 152/68.2%

1

Data from Böhlke and Chaplin 1993. Data from Smith-Vaniz et al. 1999. 3 Data from Quéro et al. 1990. 4 Data from Whitehead et al. 1984 –1986.

5

2

6

pelagic, or bathypelagic oceanic species; or deep-sea benthic species. Few clupeid or engraulid genera are amphi-Atlantic, possibly because many of the species in the Gulf of Mexico are coastal pelagics that are associated with estuaries. Among the demersal continentalshelf taxa, a majority to all of the muraenid, ophichthid, congrid, pomocentrid, labrid, and scarid genera are amphi-Atlantic, but the majority of the triglid, serranid, haemulid, sparid, sciaenid, labrisomid, and gobiid genera are not. The Gulf of Mexico shares 390 (56%) genera and 180 (81%) families with the eastern Central Pacific and 242 (36%) genera and 152 (68%) families with the eastern North Pacific (see Table 3). Most of these genera and families are neritic coastal; epipelagic, mesopelagic, or bathypelagic oceanic species; or deep-sea benthic species. More genera of Clupeidae and Engraulidae are shared between the eastern Central Pacific and the Gulf of Mexico

than between the eastern Atlantic and the Gulf of Mexico. Amphi-American distribution patterns are common in several demersal continental-shelf families: Ophichthidae, Syngnathidae, Triglidae, Serranidae, Haemulidae, Sciaenidae, Dactyloscopidae, Chaenopsidae, Labrisomidae, Eleotridae, Gobiidae, and Paralichthyidae. In all cases, the amphi-American genera are endemic to the New World. The families Dactyloscopidae, Chaenopsidae, and Achiridae are also endemic to the New World. The amphi-American genera and families diverged from their ancestral taxa as the Central and South Atlantic Ocean formed and created a barrier between the New World and the source faunas in the eastern Atlantic and IndoPacific. Most of these amphi-American taxa are associated with shore zones and have limited means of dispersal. The ophichthyid genera, with pelagic leptocephalous larvae, may be exceptions, but little is known of their early life history.

Data from Smith and Heemstra 1986. Data from Fischer et al. 1995. 7 Data from Eschmeyer et al. 1983, in part.

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14 INTRODUCTION

Based on this analysis, the Gulf of Mexico is a diverse subregion of the western Central Atlantic but not a biogeographic province. The high diversity of fish taxa is a function of the range of habitat diversity and access to fauna of the western Central Atlantic. Over most of the geological history of the Gulf, the Florida and Yucatan Peninsulas were submerged (Salvador 1991) and did not serve as vicariant mechanisms.

Appendix: New Records and Emendations to Fishes Included in Volume 1 of Fishes of the Gulf of Mexico Between the times of submission of the first and second volumes, a number of discrepancies were noted in the taxa treated in the first volume. These discrepancies are the result of overlooking literature records of fishes reported from the Gulf, overlooking museum specimens from the Gulf, or becoming aware of new literature sources that were not available before the first volume went to press. For these reasons, an appendix is added at the end of the introduction to provide an annotated list of these new records. The species records include distributional information, a designation of where the new record goes in the key to the species of the Gulf of Mexico published in the first volume, and diagnostic characters distinguishing the new record from the species diagnosed by the couplet. Literature records and museum catalog numbers are provided where appropriate.

Rhinochimaeridae New record: Harriotta raleighana (Goode and Bean, 1894) This record is based on a video recording of a live specimen taken off the coast of Louisiana, in Mississippi Canyon (ca. 2730N, 8850W) at 1,920 m. Previously it was known from the western Atlantic between Chesapeake Bay and Nova Scotia, the eastern Atlantic, the southwestern Atlantic, and the Pacific Ocean (Stehmann and Bürkel 1984). This species is

distinguished from the only other species of rhinochimaerid known from the Gulf of Mexico (Rhinochimaera atlantica) by possessing a series of large denticles on dorsal margin of the caudal fin (Bigelow and Schroeder 1953).

Scyliorhinidae New record: Scyliorhinus meadi Springer, 1966 Espinosa Pérez and Huidobro-Campos (1995) reported this species from the southern Gulf of Mexico at three localities: 2232N, 9054W, 413 m; 2230N, 9103W, 431 m; and 2229N, 9104W, 451 m. Previously it was known from North Carolina to the east coast of Florida and from the Bahamas to northern Cuba (Compagno 1984). This species keys to couplet 3a in the scyliorhinid key (McEachran and Fechhelm 1998: 61). It is distinguished from S. retifer by its lack of a color pattern of black lines; instead, it is patterned with seven or eight dusky saddle markings on upper body (Compagno 1984).

Triakidae New record: Mustelus higmani Springer and Lowe, 1963 Heemstra (1997) reported this species from DeSoto Canyon (2911N, 8717W) at 1,281 m. Previously it was known from the Gulf of Venezuela and Trinidad to Santos, Brazil (Heemstra 1997). This species is distinguished from the other three species of Mustelus in the Gulf of Mexico by the following combination of characters: denticles are tridentate between first dorsal fin and pectoral fin origins; orbit diameter is 2.2% to 3.4% of TL; upper labial fold is about equal to lower labial fold; and snout is 6.9% to 10% of TL (Heemstra 1997). New record: Mustelus sinusmexicanus Heemstra, 1997 Heemstra described this species from the Gulf of Mexico, from Panama City to Campeche Bay, between 36 and 229 m. Previously it was confused with M. canis or possibly with

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14 INTRODUCTION

Based on this analysis, the Gulf of Mexico is a diverse subregion of the western Central Atlantic but not a biogeographic province. The high diversity of fish taxa is a function of the range of habitat diversity and access to fauna of the western Central Atlantic. Over most of the geological history of the Gulf, the Florida and Yucatan Peninsulas were submerged (Salvador 1991) and did not serve as vicariant mechanisms.

Appendix: New Records and Emendations to Fishes Included in Volume 1 of Fishes of the Gulf of Mexico Between the times of submission of the first and second volumes, a number of discrepancies were noted in the taxa treated in the first volume. These discrepancies are the result of overlooking literature records of fishes reported from the Gulf, overlooking museum specimens from the Gulf, or becoming aware of new literature sources that were not available before the first volume went to press. For these reasons, an appendix is added at the end of the introduction to provide an annotated list of these new records. The species records include distributional information, a designation of where the new record goes in the key to the species of the Gulf of Mexico published in the first volume, and diagnostic characters distinguishing the new record from the species diagnosed by the couplet. Literature records and museum catalog numbers are provided where appropriate.

Rhinochimaeridae New record: Harriotta raleighana (Goode and Bean, 1894) This record is based on a video recording of a live specimen taken off the coast of Louisiana, in Mississippi Canyon (ca. 2730N, 8850W) at 1,920 m. Previously it was known from the western Atlantic between Chesapeake Bay and Nova Scotia, the eastern Atlantic, the southwestern Atlantic, and the Pacific Ocean (Stehmann and Bürkel 1984). This species is

distinguished from the only other species of rhinochimaerid known from the Gulf of Mexico (Rhinochimaera atlantica) by possessing a series of large denticles on dorsal margin of the caudal fin (Bigelow and Schroeder 1953).

Scyliorhinidae New record: Scyliorhinus meadi Springer, 1966 Espinosa Pérez and Huidobro-Campos (1995) reported this species from the southern Gulf of Mexico at three localities: 2232N, 9054W, 413 m; 2230N, 9103W, 431 m; and 2229N, 9104W, 451 m. Previously it was known from North Carolina to the east coast of Florida and from the Bahamas to northern Cuba (Compagno 1984). This species keys to couplet 3a in the scyliorhinid key (McEachran and Fechhelm 1998: 61). It is distinguished from S. retifer by its lack of a color pattern of black lines; instead, it is patterned with seven or eight dusky saddle markings on upper body (Compagno 1984).

Triakidae New record: Mustelus higmani Springer and Lowe, 1963 Heemstra (1997) reported this species from DeSoto Canyon (2911N, 8717W) at 1,281 m. Previously it was known from the Gulf of Venezuela and Trinidad to Santos, Brazil (Heemstra 1997). This species is distinguished from the other three species of Mustelus in the Gulf of Mexico by the following combination of characters: denticles are tridentate between first dorsal fin and pectoral fin origins; orbit diameter is 2.2% to 3.4% of TL; upper labial fold is about equal to lower labial fold; and snout is 6.9% to 10% of TL (Heemstra 1997). New record: Mustelus sinusmexicanus Heemstra, 1997 Heemstra described this species from the Gulf of Mexico, from Panama City to Campeche Bay, between 36 and 229 m. Previously it was confused with M. canis or possibly with

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M. norrisi. This species is distinguished from the other three Mustelus species in the Gulf of Mexico by the following combination of characters: Denticles are tridentate between first dorsal fin and pectoral fin origins; orbit diameter is 2.2% to 3.4% of TL; upper labial fold is distinctly longer than lower labial fold; and snout is 5.4% to 6.3% of TL (Heemstra 1997).

lantic Oceans, and in the Atlantic only off French Guiana. This family is distinguished from the other families of lamniform sharks by the following combination of characters: Snout is elongated and flattened into a daggerlike blade; precaudal pits are absent; and ventral caudal fin lobe is absent (Compagno 1984).

Narcinidae Squalidae Replacement name: Etmopterus robinsi Schofield and Burgess, 1997 Schofield and Burgess (1997) described this species for specimens that had previously been recognized as E. hillianus (Poey, 1861) from the Gulf of Mexico and off Nicaragua, and for some of the specimens from the Lesser Antilles at 225 to 430 m. Thus, E. hillianus in the first volume is now recognized as E. robinsi.

Oxynotidae New record: Oxynotus caribbaeus Cervigón, 1961 Espinosa Pérez and Huidobro-Campos (1995) listed this species from the southern Gulf of Mexico (2232N, 9054W) at 413 m. Previously it was known from Venezuela (Compagno 1984). This family keys to couplet 1b in the squaliform key (McEachran and Fechhelm 1998: 102). It is distinguished from Squalidae by possessing a very high and compressed body, with very high dorsal fins, rather than a low and cylindrical body and low dorsal fins as in Squalidae. Oxynotus caribbaeus is the only species of Oxynotidae in the western Atlantic (Compagno 1984).

Mitsukurinidae New record: Mitsukurina owstoni Jordan, 1898 Glen Parsons et al. (2002) reported the capture of this species off Mississippi (2844.62N, 8834.13W) at 919 to 1,099 m. This species, the sole member of Mitsukurinidae, was previously known from the Pacific, Indian, and At-

Incorrect record: Benthobatis marcida Bean and Weed, 1909 This species was incorrectly listed from the Gulf of Mexico. It has been recorded from the Straits of Florida but not the Gulf of Mexico (Marcelo de Carvalho, pers. comm., 1998). Replacement name: Narcine bancrofti (Griffith, 1834) McEachran and de Carvalho (2002) elevated this species from synonymy with N. brasiliensis for populations from North Carolina to the Gulf of Mexico, including the Greater and Lesser Antilles, between the shoreline and 37 m. Thus, Narcine bancrofti replaces N. brasiliensis in the first volume.

Rhinopteridae New record: Rhinoptera brasiliensis Castro-Aguirre and Espinosa Pérez (1996) list this species from Tuxpan, Veracruz (205825N, 971715W). Previously it was known from the southern Caribbean off Colombia (McEachran and de Carvalho 2002). This species is distinguished from the other species in the Gulf of Mexico (R. bonasus) by the following combination of characters: Series of tooth plates in each jaw number 8 to 10 (usually 9), and median series of tooth plates is about 5 times as long as it is high (Bigelow and Schroeder 1953; McEachran and de Carvalho 2002).

Congridae New record: Acromycter perturbator (Parr, 1932)

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Three lots of specimens of this species were discovered in the Texas Cooperative Wildlife Collection (TCWC) from the northern Gulf of Mexico off Louisiana and Florida between 896 and 1,170 m. Previously it was known from New Jersey, the Straits of Florida, the Bahamas, and Jamaica (Smith 1989). This species keys to couplet 13b of the congrid key (McEachran and Fechhelm 1998: 292). It is distinguished from Pseudophichthys splendens in that the posterior naris is opposite the upper margin of eye rather than at midlevel of eye (Smith 1989).

Eurypharyngidae New record: Eurypharynx pelecanoides A specimen of this species was discovered in the National Museum of Natural History (USNM) from Louisiana, 2910N, 8757W between 658 and 732 m, R / V Oregon, Station 3660. It is a cosmopolitan species and is distinguished from the other species in the Gulf of Mexico by the following combination of characters: Upper jaw is present; pharynx is highly distensible; gill opening is closer to anus than to tip of snout; and pectoral fin is minute (Böhlke 1966).

Ariidae New record: Arius assimilis (Günther, 1864) This species is reported from the southern Gulf of Mexico from Veracruz, Tabasco, and Campeche (Castro-Aguirre et al. 1999, as Ariopsis assimilis). It was previously known from Yucatán to Honduras along the coast and in estuaries, lagoons, and fresh waters (Taylor and Menezes 1978). It keys to couplet 2a of the ariid key (McEachran and Fechhelm 1998: 360) and is distinguished from A. felis by the following combination of characters: Longitudinal fleshy groove on top of head does not extend anteriorly to level of eyes; gill rakers on outer surface of first gill arch number 16 to 18; and gill rakers on outer surface of second gill arch number 18 to 22 (in A. felis, the longitudinal fleshy groove on top of head extends an-

teriorly to level of eyes; gill rakers on outer surface of first gill arch number 13 to 16; and gill rakers on outer surface of second arch number 13 to 17; Taylor and Menezes 1978). New record: Cathorops aguadulce (Meek, 1904) This species is recorded from Veracruz to Guatemala in estuaries, lagoons, and fresh waters by Castro-Aguirre et al. (1999) and replaces (in part) Cathorops sp. in McEachran and Fechhelm (1998: 363). It was previously known from Venezuela to Panama. It is distinguished from the other species of the genus in the Gulf of Mexico by the following combination of characters: Axillary pore is much smaller than nasal opening; internal surface of pectoral and pelvic fins is gray to grayish brown; first gill arch has 14 to 17 gill rakers; second gill arch has 14 to 19 gill rakers; and serrations on inner aspect of pectoral spine are straight and well developed (Castro-Aguirre et al. 1999). New record: Cathorops melanopus (Günther, 1864) This species is reported from Tamaulipas to Honduras in estuaries, lagoons, and fresh waters by Castro-Aguirre et al. (1999) and replaces (in part) Cathorops sp. in McEachran and Fechhelm (1998: 363). It is distinguished from the other species of the genus in the Gulf of Mexico by the following combination of characters: Axillary pore is as large as nasal opening, and internal surface of pectoral and pelvic fins is darkly pigmented (Castro-Aguirre et al. 1999). New record: Cathorops spixi (Agassiz, 1829) This species is reported from Tabasco, Campeche, and Yucatán in estuaries, lagoons, and fresh waters by Castro-Aguirre et al. (1999) and replaces (in part) Cathorops sp. in McEachran and Fechhelm (1998: 363). This species was previously known from the northern coast of South America from Colombia to Rio de Janeiro. It is distinguished from the other

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species of Cathorops by the following combination of characters: Axillary pore is much smaller than nasal opening; internal surface of pectoral and pelvic fins is gray to grayish brown; first gill arch has 18 to 22 gill rakers; second gill arch has 17 to 21 gill rakers; serrations on inner aspect of pectoral fin spine are weak and oblique (Castro-Aguirre et al. 1999).

Alepocephalidae New record: Alepocephalus australie Barnhard, 1923 Two lots of specimens of this species were discovered in the Texas Cooperative Wildlife Collection (TCWC) that were collected off Texas and Florida at depths from 752 to 936 m. Previously this species was known from the western Atlantic between 10 and 39 and the eastern Atlantic from South Africa to southeastern Ireland (Markle and Quéro 1984). It keys to couplet 10b in the alepocephalid key (McEachran and Fechhelm 1998: 382) and is distinguished from A. productus by the following combination of characters: Transverse scale rows from dorsal fin origin to first lateral line scale number fewer than 30, and posterior margin of eye is slightly nearer to tip of snout than to posterior margin of operculum (Markle and Quéro 1984).

Melanostomiidae New record: Eustomias achirus Parin and Pokhilskaya, 1974 Clarke (1998) recorded this species from four localities in the northern Gulf of Mexico: 2700N, 8600W; 2914N, 8740W; 2900N, 8802W; and 2833N, 8848W. It also occurs between 20C and 40C in the Atlantic and in the Coral Sea in the Pacific. It keys to couplet 33a in McEachran and Fechhelm (1998: 479) and is distinguished from E. lipochirus by the following combination of characters: Chin barbel is short, with three branches off stem that extend beyond tip of barbel and bear internal ovoid bodies; ovoid bodies are about the same diameter as stem

and are spaced about three ovoid body diameters apart; branches are lightly pigmented or unpigmented between bodies (Clarke 1998). Replacement name: Eustomias monoclonus Regan and Trewavas, 1930 Clarke (1999) reclassified E. dendriticus Regan and Trewavas, 1930, as E. monoclonus Regan and Trewavas, 1930, in the western Atlantic, including the Gulf of Mexico.

Aulopodidae Replacement name: Aulopus filamentosus (Bloch, 1792) According to B. A. Thompson, S. W. Ross, and K. Sulak (pers. comm., April 1998; in Smith-Vaniz et al. 1999), Aulopus filamentosus is the senior synonym of A. nanae.

Ophidiidae Replacement name: Barathrites parri Nybelin, 1957 This species replaces Barathrites sp. in McEachran and Fechhelm (1998: 705). It is distinguished from the other species of the genus by the following combination of characters: Branchiostegal rays number seven, and basibranchial tooth plate is single. The other species, B. iris, has six branchiostegal rays and no basibranchial tooth plate (Nielsen et al. 1999). New record: Bassozetus normalis Gill, 1884 J. G. Nielsen and N. G. Merritt (2000) record this species from throughout the Gulf of Mexico at depths from 1,775 to 3,843 m. It was previously known from the tropical Atlantic Ocean between 1,760 and 5,062 m (Nielsen and Merrett 2000). This species is distinguished from the other two congeners in the Gulf of Mexico by the following combination of characters: Long gill rakers on first arch number 13 to 20; scales in oblique row between dorsal fin origin and anus number 25 to 35; and body depth is 8.4% to 11% of SL. In B. compressus, long gill rakers on first arch number 11 to 14; scales in oblique row be-

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tween dorsal fin origin and anus number 15 to 20; and body depth is 8% to 14% of SL. In B. robustus, long gill rakers on first arch number 13 to 16; scales in oblique row between dorsal fin origin and anus number 30 to 40; and body depth is 10% to 19% of SL (Nielsen and Merritt 2000). Incorrect record: Ophidion holbrooki (Putnam, 1874) J. G. Nielsen et al. (1999) consider Ophidion beani a junior synonym of O. holbrooki. New record: Ophidion josephi Girard, 1859 J. G. Nielson et al. (1999) consider this species valid but provide no characters to distinguish it from the other species of the genus. It occurs from Georgia to the northern Gulf of Mexico. New record: Ophidion selenops Robins and Böhlke, 1959 J. G. Nielsen et al. (1999) list this species from South Carolina to the southeastern Gulf of Mexico. It is distinguished from the other species of Ophidion in the Gulf of Mexico by the following combination of characters: Tip of snout has prominent spine, and body is silvery.

Bythitidae New record: Bellottia apoda Giglioli, 1883 Several specimens of this species were captured in the northern Gulf of Mexico at 2747N, 9130.5W in 540 to 580 m and at 2744.7N, 9113.3W in 540 m. It was previously known from the tropical Atlantic and the Mediterranean Sea. This species keys to 3b of the bythitid key (McEachran and Fechhelm 1998: 742) and is distinguished from Cataetyx laticeps by lacking pelvic fins and possessing several sharp-pointed spines on the preoperculum (Cohen and Nielsen 1978).

Macrouridae New record: Caelorinchus ventilux Marshall and Iwamoto, 1973

A single specimen of this species was discovered in the Texas Cooperative Wildlife Collection (TCWC 4490.02) from Campeche Bay (210730N, 861712W, depth unknown). Previously it was known from the West Indies. This species is distinguished from the other species of Caelorinchus from the Gulf of Mexico by the following combination of characters: Ventral side of snout is scaled except for naked broad band anterior to premaxilla, and black band is present between dorsal fins. The other species either lack scales on ventral side of snout (C. occa) or have fully scaled ventral snouts, and lack black bands between dorsal fins (Marshall and Iwamoto 1973).

Moridae Replacement name: Laemonema goodebeanorum Meléndez and Markle, 1997 This species replaces Laemonema sp. in volume 1 (McEachran and Fechhelm 1998: 806). It occurs from Canada to Brazil, including the Gulf of Mexico, between 180 and 792 m.

Batrachoididae New record: Batrachoides surinamensis (Bloch and Schneider, 1801) Castro-Aguirre et al. (1999) report this species from Campeche to Brazil in coastal waters and estuaries. It is distinguished from the other batrachoidids in the Gulf of Mexico in possessing embedded scales; the other species are naked. New record: Opsanus dichrostomus Collette, 2001 This species occurs in the southern Gulf of Mexico at Tampico, Veracruz, and along the west coast of the Yucatán Peninsula. It is also known from Cuba, the east coast of the Yucatán Peninsula, and Belize (Collette 2001). It is distinguished from the other batrachoidid species in the Gulf of Mexico by the following combination of characters: Second dorsal fin rays number 23 to 25; pectoral fin rays number 17 or 19; interorbital distance is less than orbit diameter (Collette 2001).

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Mugilidae Incorrect record: Mugil gaimardianus Desmarest, 1831 This species is included in volume 1 but is a junior synonym of M. curema Valenciennes, 1836, according to Harrison (2002). New record: Joturus pichardi Poey, 1860 This species occurs in the Gulf of Mexico, from the mouth of the Rio Grande to Yucatán, along the Caribbean coast of Central America and Colombia, and in the Bahamas and the Antilles. Adults inhabit freshwater streams but spawn in brackish water or the ocean (Harrison 2002). This species is distinguished from the other species of mugilids in the Gulf of Mexico in possessing a subterminal mouth (Harrison 2002).

Syngnathidae New record: Cosmocampus brachycephalus (Poey, 1868) This species is recorded from the eastern Gulf of Mexico at the Dry Tortugas. It ranges from the southern coast of Florida and the Bahamas to Venezuela, including the Greater and Lesser Antilles (Dawson 1982). This species keys to couplet 16b in the syngnathid key (McEachran and Fechhelm 1998: 1049) and is distinguished from C. albirostris in having a snout length 26.3% to 34.5% of head length and a snout depth 45.5% to 71.4% of snout length rather than a snout length 35.7% to 45.5% of head length and a snout depth 25% to 38.5% of snout length as in C. albirostris.

Literature Cited Aboussouan, A., and J. M. Leis. 1984. Balistoidei: development. In: OSF: 450 – 459. Backus, R. H., and J. E. Craddock. 1977. Data report for Atlantic pelagic zoogeography. Tech. Rep. Woods Hole Oceanographic Institution. WHOI-77-4. 8 pp., 3 appendixes. Backus, R. H., J. E. Craddock, R. L. Haedrich,

and B. H. Robison. 1977. Atlantic zoogeography. In: FWNA 7: 266 –287. Bigelow, H. B., and W. C. Schroeder. 1953. Sawfishes, guitarfishes, skates and rays, and chimaeroids. In: FWNA 2: 1–588. Böhlke, J. E. 1966. Order Lyomeri. In: FWNA 5: 603 – 625. Böhlke, J. E., and C. C. G. Chaplin. 1993. Fishes of the Bahamas and adjacent tropical waters. 2d ed. Austin: University of Texas Press. Bowen, B. W., and J. C. Avis. 1990. Genetic structure of Atlantic and Gulf of Mexico populations of sea bass, menhaden, and sturgeon: influence of zoogeographic factors and life-history patterns. Mar. Biol. 107: 371–381. Briggs, J. C. 1974. Marine zoogeography. New York: McGraw-Hill. 475 pp. Castro-Aguirre, J. L., and H. S. Espinosa Pérez. 1996. Listados faunísticos de México. Vol. 7, Catálogo sistemático de las rayas y especies afines de México (Chondrichthyes: Elasmobranchii: Rajiformes: Batoideomorpha). Mexico City: UNAM. 75 pp. Castro-Aguirre, J. L., H. S. Espinosa Pérez, and J. J. Schmitter-Soto. 1999. Ictiofauna estuarino-lagunar y vicaria de México. Mexico City: LIMUSA Grupo Noriega Editores. 711 pp. Chapleau, F. 1993. Pleuronectiform relationships: a cladistic reassessment. Bull. Mar. Sci. 52: 516 –540. Chapleau, F., and A. Keast. 1988. A phylogenetic reassessment of the monophyletic status of the family Soleidae, with comments on the suborder Soleoidei (Pisces: Pleuronectiformes). Can. J. Zool. 66: 2797–2810. Clarke, T. A. 1998. Pelagic fishes of the genus Eustomias (Melanostomiidae) presently associated with Eustomias achirus Parin and Pokhilskaya, with the description of five new species. Copeia: 676 – 686. Clarke, T. A. 1999. Pelagic fishes of the genus Eustomias (Melanostomiidae) similar to Eustomias dendriticus Regan and Trewavas, with the description of seven new species. Copeia: 1001–1013.

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Mugilidae Incorrect record: Mugil gaimardianus Desmarest, 1831 This species is included in volume 1 but is a junior synonym of M. curema Valenciennes, 1836, according to Harrison (2002). New record: Joturus pichardi Poey, 1860 This species occurs in the Gulf of Mexico, from the mouth of the Rio Grande to Yucatán, along the Caribbean coast of Central America and Colombia, and in the Bahamas and the Antilles. Adults inhabit freshwater streams but spawn in brackish water or the ocean (Harrison 2002). This species is distinguished from the other species of mugilids in the Gulf of Mexico in possessing a subterminal mouth (Harrison 2002).

Syngnathidae New record: Cosmocampus brachycephalus (Poey, 1868) This species is recorded from the eastern Gulf of Mexico at the Dry Tortugas. It ranges from the southern coast of Florida and the Bahamas to Venezuela, including the Greater and Lesser Antilles (Dawson 1982). This species keys to couplet 16b in the syngnathid key (McEachran and Fechhelm 1998: 1049) and is distinguished from C. albirostris in having a snout length 26.3% to 34.5% of head length and a snout depth 45.5% to 71.4% of snout length rather than a snout length 35.7% to 45.5% of head length and a snout depth 25% to 38.5% of snout length as in C. albirostris.

Literature Cited Aboussouan, A., and J. M. Leis. 1984. Balistoidei: development. In: OSF: 450 – 459. Backus, R. H., and J. E. Craddock. 1977. Data report for Atlantic pelagic zoogeography. Tech. Rep. Woods Hole Oceanographic Institution. WHOI-77-4. 8 pp., 3 appendixes. Backus, R. H., J. E. Craddock, R. L. Haedrich,

and B. H. Robison. 1977. Atlantic zoogeography. In: FWNA 7: 266 –287. Bigelow, H. B., and W. C. Schroeder. 1953. Sawfishes, guitarfishes, skates and rays, and chimaeroids. In: FWNA 2: 1–588. Böhlke, J. E. 1966. Order Lyomeri. In: FWNA 5: 603 – 625. Böhlke, J. E., and C. C. G. Chaplin. 1993. Fishes of the Bahamas and adjacent tropical waters. 2d ed. Austin: University of Texas Press. Bowen, B. W., and J. C. Avis. 1990. Genetic structure of Atlantic and Gulf of Mexico populations of sea bass, menhaden, and sturgeon: influence of zoogeographic factors and life-history patterns. Mar. Biol. 107: 371–381. Briggs, J. C. 1974. Marine zoogeography. New York: McGraw-Hill. 475 pp. Castro-Aguirre, J. L., and H. S. Espinosa Pérez. 1996. Listados faunísticos de México. Vol. 7, Catálogo sistemático de las rayas y especies afines de México (Chondrichthyes: Elasmobranchii: Rajiformes: Batoideomorpha). Mexico City: UNAM. 75 pp. Castro-Aguirre, J. L., H. S. Espinosa Pérez, and J. J. Schmitter-Soto. 1999. Ictiofauna estuarino-lagunar y vicaria de México. Mexico City: LIMUSA Grupo Noriega Editores. 711 pp. Chapleau, F. 1993. Pleuronectiform relationships: a cladistic reassessment. Bull. Mar. Sci. 52: 516 –540. Chapleau, F., and A. Keast. 1988. A phylogenetic reassessment of the monophyletic status of the family Soleidae, with comments on the suborder Soleoidei (Pisces: Pleuronectiformes). Can. J. Zool. 66: 2797–2810. Clarke, T. A. 1998. Pelagic fishes of the genus Eustomias (Melanostomiidae) presently associated with Eustomias achirus Parin and Pokhilskaya, with the description of five new species. Copeia: 676 – 686. Clarke, T. A. 1999. Pelagic fishes of the genus Eustomias (Melanostomiidae) similar to Eustomias dendriticus Regan and Trewavas, with the description of seven new species. Copeia: 1001–1013.

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Cohen, D. M., and J. G. Nielsen. 1978. Guide to the identification of the genera of the fish order Ophidiiformes, with a tentative classification to the order. NOAA Tech. Rep. NMFS Circ. 417: 1–72. Collette, B. B. 1978. Batrachoididae. In: FAO species identification sheets for fishery purposes, Western Central Atlantic (Fishing area 31), ed. W. Fischer. Rome: FAO. Collette, B. B. 2001. Opsanus dichrostomus, a new toadfish (Teleostei: Batrachoididae) from the western Caribbean Sea and southern Gulf of Mexico. Occas. Pap. Mus. Univ. Mich. 781: 1–16. Compagno, L. J. V. 1984. Sharks of the world. In: FAO Species Catalogue, Vol. 4, Sharks of the world. An annotated and illustrated catalogue of sharks species known to date. FAO Fisheries Synopsis 125 (4, parts 1 and 2): 1– 655. Dawson, C. E. 1982. Family Syngnathidae. In: FWNA 8: 1–172. Dooley, J. K. 1972. Fishes associated with the pelagic sargassum complex, with a discussion of the sargassum community. Contrib. Mar. Sci. 16: 1–32. Eschmeyer, W. N., E. S. Herald, and H. Hammann. 1983. A field guide to the Pacific coast fishes of North America. Peterson Field Guide Series. Boston: Houghton Mifflin. 336 pp. Espinosa Pérez, H. S., and L. HuidobroCampos. 1995. Nuevos registros de tiburones en México. Rev. Soc. Mex. Nat. 46: 113 –120. Fischer, W., F. Krupp, W. Schneider, C. Sommer, K. E. Carpenter, and V. H. Niem, eds. 1995. Guía FAO para la identificación de especies para los fines de la pesca, Pacífico Centro-oriental. Rome: FAO. 3 vols., 1813 pp. Galloway, W. E., D. G. Bebout, W. L. Fisher, J. B. Dunlap, Jr., R. Cabrera-Castro, J. E. Lugo-Rivera, and T. M. Scott. 1991. Cenozoic. In: The geology of North America, Vol. J, The Gulf of Mexico Basin, ed. A. Salvador, 245 –324. Boulder: Geological Society of America.

Gartner, J. V., Jr., T. L. Hopkins, R. C. Baird, and D. M. Milliken. 1987. The lanternfishes (Pisces: Myctophidae) of the eastern Gulf of Mexico. Fish. Bull. 85: 81–98. Gibbs, R. H., Jr., T. A. Clarke, and J. R. Gomon. 1983. Taxonomy and distribution of the stomioid fish genus Eustomias (Melanostomiidae). Part 1, Subgenus Nominostomias. Smithsonian Contrib. Zool. 380: 1–139. Gomon, J. R., and R. H. Gibbs, Jr. 1985. Taxonomy and distribution of the stomioid fish genus Eustomias (Melanostomiidae). Part 2, Biradiostomias, new subgenus. Smithsonian Contrib. Zool. 409: 1–58. Harrison, I. J. 2002. Mugilidae. In: FAO-WCA: 1071–1085. Heemstra, P. C. 1997. A review of the smoothhound sharks (genus Mustelus, family Triakidae) of the western Atlantic Ocean, with descriptions of two new species. Bull. Mar. Sci. 60: 894 –928. Karnella, C. 1987. Biology of midwater fishes of Bermuda Ocean Acre: family Myctophidae, lanternfishes. Smithsonian Contrib. Zool. 452: 51–168. Leis, J. M. 1984. Tetraodontoidei: development. In: OSF: 459 – 463. Leis, J. M., and D. S. Rennis. 1983. The larvae of Indo-Pacific coral reef fishes. Sydney, Australia: New South Wales University Press. 269 pp. Leviton, A. E., R. H. Gibbs, Jr., E. Heal, and C. E. Dawson. 1985. Standards in herpetology and ichthyology. Part 1, Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia: 802 – 832. Longley, W. H., and S. F. Hildebrand. 1941. Systematic catalogue of the fishes of Tortugas, Florida, with observations on color, habits, and local distribution. Carnegie Inst. Wash. Publ. 535: 1–331. Markle, D. F., and J.-C. Quéro. 1984. Alepocephalidae. In: FN-EAM 1: 228 –253. Marshall, N. B., and T. Iwamoto. 1973. Genus Coelorhynchus. In: FWNA 6: 538 –563. Masuda, H., K. Amaoka, C. Araga, T. Uyeno,

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and T. Yoshino. 1984. The fishes of the Japanese Archipelago. Tokyo: Tokai University Press. 437 pp., 370 pls. McEachran, J. D., and M. R. de Carvalho. 2002. Batoid fishes. In: FAO-WCA: 508 – 589. McEachran, J. D., and J. D. Fechhelm. 1998. Fishes of the Gulf of Mexico, vol. 1, Myxiniformes to Gasterosteiformes. Austin: University of Texas Press. 1120 pp. Meléndez, C. R., and D. F. Markle. 1997. Phylogeny and zoogeography of Laemonema and Guttigadus (Pisces; Gadiformes; Moridae). Bull. Mar. Sci. 61: 593 – 670. Nelson, J. S. 1994. Fishes of the world. 3rd ed. New York: John Wiley and Sons. 600 pp. Nielsen, J. G., and N. G. Merritt. 2000. Revision of the cosmopolitan deep-sea genus Bassozetus (Pisces: Ophidiidae) with two new species. Galathea Rep. 18: 1–57. Nielsen, J. G., D. M. Cohen, D. F. Markle, and C. R. Robins. 1999. Ophidiiform fishes of the world (Order Ophidiiformes). FAO species catalogue. Vol. 18: 1–178. Parsons, G. R., G. W. Lingram, Jr., and R. Havard. 2002. First record of the goblin shark Mitsukurina owstoni, Jordan (family Mitsukurinidae) in the Gulf of Mexico. Southeast. Nat. 1: 189 –192. Quéro, J.-C., J.-C. Hureau, C. Karrer, A. Post, and L. Saldanha. 1990. Check-list of the fishes of the eastern tropical Atlantic. Vols. 1–3. Lisbon: UNESCO. Riggs, S. R. 1984. Paleoceanographic model of neogene phosphorite deposition, U.S. Atlantic continental margin. Science 223: 123 –131. Riggs, S. R., S. P. Ellison, Jr., W. L. Fisher, W. E. Galloway, M. L. W. Jackson, and R. A. Morton. 1991. Mineral resources and geopressured-geothermal energy. In: The geology of North America, Vol. J, The Gulf of Mexico Basin, ed. A. Salvador, 495 –528. Boulder: Geological Society of America. Robins, C. R. 1971. Distributional patterns of fishes from coastal and shelf waters of the tropical western Atlantic. Symposium on investigations and resources of the Carib-

bean Sea and adjacent regions. Papers on Fisheries Resources. Rome: FAO. 90 –255. Rosen, D. E. 1973. Suborder Cyprinodontoidei. In: FWNA 6: 229 –262. Salvador, A. 1991. Origin and development of the Gulf of Mexico basin. In: The geology of North America, Vol. J, The Gulf of Mexico Basin, ed. A. Salvador, 389 – 444. Boulder: Geological Society of America. Schofield, P. J., and G. H. Burgess. 1997. Etmopterus robinsi (Elasmobranchii, Etmopteridae), a new species of deepwater lantern shark from the Caribbean Sea and western Atlantic, with a redescription of Etmopterus hillianus. Bull. Mar. Sci. 60: 1060 – 1073. Shipp, R. L., and R. W. Yerger. 1969a. A new puffer fish, Sphoeroides parvus, from the western Gulf of Mexico, with a key to species of Sphoeroides from the Atlantic and Gulf coasts of the United States. Proc. Biol. Soc. Wash. 82: 477– 488. Shipp, R. L., and R. W. Yerger. 1969b. Status, characters, and distribution of the northern and southern puffers of the genus Sphoeroides. Copeia: 425 – 433. Smith, D. G. 1989. Family Congridae. In: FWNA 9(1): 460 –567. Smith, M. L., K. E. Carpenter, and R. W. Waller. 2002. An introduction to oceanography, geology, biogeography, and fisheries of the tropical and subtropical Atlantic. In: FAOWCA: 1–23. Smith-Vaniz, W. F., B. B. Collette, and B. E. Luckhurst. 1999. Fishes of Bermuda: History, zoogeography, annotated checklist, and identification keys. Lawrence (KS): Allen Press. 424 pp. Springer, V. G. 1959. Blenniid fishes of the genus Chasmodes. Texas J. Sci. 9: 321– 334. Stehmann, M., and D. L. Bürkel. 1984. Rhinochimaeridae. In: FN-EAM 1: 216 –218. Suttkus, R. D. 1963. Family Lepisosteidae. In: FWNA 3: 68 – 88. Sutton, T. T., and T. L. Hopkins. 1996. Species composition, abundance, and vertical distribution of the stomiid (Pisces: Stomii-

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22 INTRODUCTION

formes) fish assemblage of the Gulf of Mexico. Bull. Mar. Sci. 59: 530 –542. Taylor, W. R., and N. A. Menezes. 1978. Ariidae. In: FAO species identification sheets for fishery purposes, Western Central Atlantic (Fishing area 31), ed. W. Fischer. Rome: FAO. Whitehead, P. J. P., M.-L. Bauchot, J.-C. Hu-

reau, J. Nielsen, and E. Tortonese, eds. 1984 –1986. Fishes of the North-eastern Atlantic and the Mediterranean. Vols. 1–3. Paris: UNESCO. Williams, J. T. 1983. Taxonomy and ecology of the genus Chasmodes (Pisces: Blenniidae) with a discussion of its zoogeography. Bull. Fla. St. Mus. Biol. Sci. 29: 66 –100.

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23

SCORPAENIFORMES Scorpaeniformes form an unresolved polytomy with the Perciformes, Pleuronectiformes, and Tetraodontiformes. The order comprises 26 families, and representatives of 4 of these families (Dactylopteridae, Scorpaenidae, Triglidae, and Peristediidae) occur in the Gulf of Mexico. Johnson and Patterson (1993) place Dactylopteridae in their own order, Dactylopteriformes, because of a lack of derived characters shared between dactylopterids and the remaining families; however, most modern authors retain dactylopterids in Scorpaeniformes. Nelson (1994) placed Peristediinae in Triglidae, but the two taxa are treated as separate families, following William Richards (pers. comm., January 2002).

Key to the Families of the Gulf of Mexico (Adapted from Smith and Heemstra 1986c) 1a. Head encased in heavy bony armor . . . . . . . . . . . . . . . . . 2 1b. Head not encased in bony armor [body moderately compressed to robust; head large, with numerous spines and ridges; pectoral fin large and fanlike; dorsal fin single] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenidae p. 26 2a. Lower two or three pectoral fin rays free of membranes and enlarged; preopercular spine of moderate length . . . . 3 2b. Lower two or three pectoral fin rays attached to membranes and not enlarged; preopercular spine very long, extending beyond pelvic fin base [body elongate and squarish in cross section; head bones united to form bony helmet] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dactylopteridae p. 24 3a. Barbels on lip and chin; pectoral fin with two lowermost rays free of membranes and thickened; body covered with bony plates [body slender to moderately robust and covered with rows of spinous scutes; head bony, with many ridges and spines]. . . . . . . . . . . . . . . . . . . . . . . Peristediidae p. 70 3b. Barbels absent on lip and chin; pectoral fin with three lowermost rays free of membranes and thickened; body covered with scales [body robust and elongate; head rugose and bony; snout broad and flat]. . . . . . . . . . Triglidae p. 51

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24

DACTYLOPTERIDAE

Flying gurnards

The flying gurnards are elongate and squarish in cross section, with a large, bony head and greatly enlarged pectoral fins. Bones of head are united to form bony helmet, preopercular and posttemporal spines are greatly elongated, and opercular spine is absent in adults and juveniles of most species. Interorbital region is wide and concave. Mouth is slightly subterminal, small, and nearly horizontal. Jaw teeth are granular. Vomer and palatine lack teeth. Gill rakers are minute. Gill membranes are broadly joined to isthmus. Pectoral fin has a nearly horizontal base and 28 to 37 rays and is divided at base into anterior and posterior sections. Anterior section is narrow and has short rays, and posterior section is broad and has very long rays. First dorsal fin has seven or eight spines, with first one or two generally free and distinctly anterior to remainder of fin. Second dorsal fin has one spine and 8 rays. Pelvic fin is thoracic and consists of one spine and 4 rays. Anal fin originates below or slightly posterior to origin of second dorsal fin and has 6 or 7 rays. Caudal fin has concave posterior margin. Body is covered with scutelike scales, and two enlarged, modified scales are located on base of caudal fin. Lateral line is present or absent. Vertebrae number 22, with first 2 elongated. Gas bladder is divided into right and left halves, joined near midlength, and possesses drumming muscles. Flying gurnards occur in the tropical to warm temperate Atlantic and Indo-West Pacific Oceans. They are benthic in shallow to moderate depths and most commonly found on sandy bottoms. The enlarged pectoral fins are possibly used for defensive displays, courtship displays, and feeding, but they are not used for flying or gliding through the air, as they are in exocoetids. These fishes are seldom observed off the bottom, and their bodies are too heavy and fins too delicate for gliding. Pelvic fins are used to propel the body over the bottom. Eggs are pelagic and slightly less than 1 mm in diameter. There are about seven species in two genera, and one species occurs in the western Central Atlantic, including the Gulf of Mexico. references: Jordan and Evermann 1898; Breder 1963; Washington, Eschmeyer, et al. 1984; Washington, Moser, et al. 1984; Eschmeyer 1986b, 1994, 1997; Nelson 1994.

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DACTYLOPTERIDAE 25

Dactylopterus volitans Flying gurnard

(Linnaeus, 1758)

Dactylopterus volitans has a blunt head, extremely long preopercular spines, and fanlike pectoral fins. Interorbital region is strongly concave. Preopercular spine extends to or beyond base of pectoral fin. Measurements are expressed as percent of SL: head length 25%– 28%, snout length 6%–9%, orbit diameter 7%– 8%, upper jaw length 7%– 8%, pectoral fin length 49%–77%, pelvic fin length 25%– 26%, body depth 16%–17%. Pectoral fin extends beyond anal fin base and has 34 rays, with lower 6 rays shorter and forming separate lobe from remainder of fin. First dorsal fin has six spines, with anterior two on nape connected basally by membranes and separated from more posterior four spines. Second dorsal fin has one spine and 8 rays. Anal fin originates posterior to origin of second dorsal fin and has 6 rays, with last ray single rather than double. Color varies with background but often is orange brown to dusky, with marbling of blue spots on back; lower body is pinkish; and

pectoral fin is patterned with blue or lavender spots and blue lines. This species occurs in the tropical to warm temperate Atlantic Ocean. In the western Atlantic it occurs to a depth of about 80 m, from Massachusetts and Bermuda to Argentina, including the Gulf of Mexico, the Bahamas, and the Caribbean Sea. It is found throughout the northern Gulf of Mexico and off Veracruz in the southern Gulf. Food consists of small shrimps and crabs. Maximum known size is about 460 mm SL.

references: Longley and Hildebrand 1941; Bigelow and Schroeder 1953; Breder 1963; Böhlke and Chaplin 1968; Hoese and Moore 1977, 1998; Uyeno and Sato 1983e; Eschmeyer 1986b; Robins and Ray 1986; Roux 1986; Cervigón 1991; Boschung 1992; Randall 1996; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; SmithVaniz 2003a.

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26

SCORPAENIDAE

Scorpionfishes

Scorpionfishes are moderately compressed to robust, with a large head bearing numerous spines and ridges, large fanlike pectoral fins, and a single dorsal fin. Snout is short to moderately long and moderately oblique to nearly vertical. Premaxilla is protractile. Eye is moderate to large and laterally located. Bony ridge (suborbital stay), formed by infraorbital bone and bearing spines, is generally present below and behind eye, extending from just behind mouth to preoperculum. Preoperculum has 3 to 5 spines, with uppermost usually the largest and often bearing a smaller supplementary spine at its base. Operculum has 1 or 2 spines. Other spines on head are variably developed. Slit behind fourth gill arch is usually small or absent in western Atlantic species. Gill opening is wide, and gill membranes are free of each other and of isthmus. Pseudobranch is usually well developed. Pectoral fin is rounded to wedge shaped and has 11 to 24 rays, with inner 2 or 3 rays deeply incised or free of membranes. Dorsal fin originates posterior to eye and is usually strongly notched between anterior section of 7 to 18 spines and posterior section of 4 to 14 rays. Pelvic fin is thoracic and has 1 spine and 2 to 5 rays. Anal fin has 2 to 4 spines and 5 to 14 rays. Last ray of dorsal fin and anal fin is split at base. Caudal fin is rounded to slightly forked. Scales are usually present (always present in western Atlantic species) and, when present, are usually ctenoid, although some species have cycloid scales. Vertebrae number 24 to 31. Venom gland is often present at base of dorsal, pelvic, and anal spines. Physoclistous gas bladder is usually present and has extrinsic or intrinsic musculature. Scorpionfishes occur worldwide in tropical to temperate seas. Most species are rather sedentary benthic fishes that are associated with filamentous algae, rocks, sand, or coral structures. Species range from the intertidal zone to 2,200 m. However, one species is mesopelagic. Food consists mostly of crustaceans and ray-finned fishes that are ambushed when they approach these predominately cryptic predators. Venom glands associated with the fin spines can cause painful wounds. Many species practice internal fertilization, and some retain the fertilized eggs in the oviducts. Other species are oviparous and have small pelagic eggs. Eggs of oviparous species are often laid in gelatinous masses. There are about 388 species in 56 genera, 30 or 31 species in 11 genera occur in the western Central Atlantic, and 20 or 21 species in 9 genera occur in the Gulf of Mexico. references: Fritzsche and Fuiman 1982; Eschmeyer 1986a,c, 1994; Hureau and Litvinenko 1986a; Nelson 1994; Smith 1997; Poss and Eschmeyer 2002.

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SCORPAENIDAE 27

Key to the Species of the Gulf of Mexico (Adapted from Eschmeyer 1969b) 1a. Dorsal fin spines 13 or more; palatine teeth absent . . . . . 2 1b. Dorsal fin spines 12 or fewer; palatine teeth present. . . . . 3 2a. Pectoral fin rays 18 to 20 (usually 19); one or more small spines below row of suborbital spines (Fig. 1) Scorpaenodes caribbaeus p. 42

FIG 1

2b. Pectoral fin rays 16 to 18 (usually 17); suborbital ridge with single row of spines . . . . . . . . . . . . . . . . . Scorpaenodes tredecimspinosus p. 48 3a. Lateral line consists of continuous channel, roofed by thin scales; supraocular and postocular spines absent (except in juveniles); bones of head relatively weakly ossified and cavernous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3b. Lateral line consists of tubed scales; supraocular and postocular spines present; bones of head strongly ossified. . . . 5 4a. Pectoral fin rays 18 to 20; anal fin rays usually 6; orbit diameter about half interorbital width . . . . . . . . . . . . . . . . . . . . . . . . . Ectreposebastes imus p. 30 4b. Pectoral fin rays 21 to 24; anal fin rays usually 5; orbit diameter about equal to interorbital width . . . . . . . . . . . . . . . . . . . . . . . . . . Setarches guentheri p. 49 5a. Second preopercular spine longest; dorsal fin rays 11 or more . . . . . . . . . . . . . . . . . Helicolenus dactylopterus p. 31 5b. First preopercular spine longest; dorsal fin rays 10 or fewer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6a. Lateral line incomplete, only anterior four to six tubed scales present . . . . . . . . . . . . Phenacoscorpius nebris p. 34 6b. Lateral line complete, extending to base of caudal fin. . . . 7 7a. Scales ctenoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7b. Scales cycloid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

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28 SCORPAENIFORMES

8a. Pectoral fin not wedge shaped; longest fin rays near upper edge (fin slightly bilobed in large specimens); pectoral fin rays 21 to 24 (rarely 20) . . . Trachyscorpia cristulata p. 50 8b. Pectoral fin wedge shaped; pectoral fin rays usually 19 or fewer (rarely 20). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9a. Pectoral fin rays unbranched . . . . . . . . . . . . . . . . . . . . . 10 9b. Some or all pectoral fin rays branched (except in juveniles) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10a. Third dorsal spine distinctly elongate in specimens longer than 120 to 130 mm SL; first preorbital spine usually directed anteriorly; sides of body usually spotted . . . . . . . . . . . . . . . . . . . . . . . . . Pontinus longispinis p. 35 10b. Third dorsal spine not distinctly elongate in specimens longer than 120 to 130 mm SL; first preorbital spine directed posteriorly; sides of body with several dusky smudges but without spots . . . . . . Pontinus rathbuni p. 36 11a. Dorsal fin rays usually 10 (rarely 9), last ray double; scales small, vertical scale rows more than 55; orbit 58.8% to 91.1% of snout length. . . . Neomerinthe hemingwayi p. 33 11b. Dorsal fin rays 9 (rarely 8 or 10), last ray double; scales relatively large, vertical scale rows fewer than 50; orbit 70.9% to 142% of snout length . . . Neomerinthe beanorum p. 32 12a. Preorbital bone with two spinous points over maxilla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 12b. Preorbital bone with three or four spinous points over maxilla, except two spinous points in Scorpaena plumieri and possibly in S. dispar less than 40 mm SL . . . . . . . . . 20 13a. Supplemental preopercular spine absent; occipital pit absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13b. Supplemental preopercular spine present; occipital pit present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14a. Eye with inverted heart-shaped tabs extending downward in front of cornea and from ocular tabs at upper boundary of cornea. . . . . . . . . . . . . . . . . . . . Scorpaena inermis p. 45 14b. Eyes without inverted heart-shaped tabs . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaena calcarata p. 41 15a. Axil of pectoral fin and side of body between pectoral fin and pelvic fin with brown spots on pale background; vertical scale rows more than 50. . . Scorpaena brasiliensis p. 40 15b. Axil of pectoral fin without brown spots on pale background; vertical scale rows fewer than 50. . . . . . . . . . . . 16 16a. Snout 46% to 67% of eye length; appressed pectoral fin extending to insertion of anal fin (shorter in juveniles) . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaena agassizii p. 37 16b. Snout 67% to 111% of eye length; appressed pectoral fin extending to third anal fin spine . . . . . . . . . . . . . . . . . . . 17 17a. Spinous part of dorsal fin with dark blotch between dorsal spines three to five and six to seven . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaena bergii p. 39

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SCORPAENIDAE 29

17b. Spinous part of dorsal fin without dark blotch . . . . . . . . 18 18a. Axil of pectoral fin with white specks (less than 0.5 mm in diameter) surrounded by brown or on tan background; supraocular tentacle about 2 times orbit diameter; tympanic spine reduced or absent . . . . . . . . . . . . . . . . . . . . . . . Scorpaena grandicornis p. 44 18b. Axil of pectoral fin without white specks; supraocular tentacle usually less than orbit diameter; tympanic spine present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 19a. Pectoral fin rays 16 to 18; suborbital ridge with one or (rarely) two spines. . . . . . . . . . . . . Scorpaena elachys p. 43 19b. Pectoral fin rays 19 or 20; suborbital ridge with three or (rarely) two spines . . . . . . . . . . Scorpaena albifimbria p. 38 20a. Axil of pectoral fin with large white spots or irregular blotches on black background on specimens greater than 30 mm SL; caudal peduncle paler than anterior portion of body; pectoral fin rays 19 to 21 . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaena plumieri p. 46 20b. Axil of pectoral fin unmarked; body pale except for scattered dark blotches; pectoral fin rays usually 17 or 18 . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaena dispar p. 42

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30 SCORPAENIFORMES

Ectreposebastes imus

Garman, 1899

Ectreposebastes imus is black, with the anterior spine on the preorbital bone greatly reduced, remaining preorbital spines strongly developed, and lateral line in a continuous trough. Head and snout are relatively large, and eye is of moderate size. Teeth are small; arranged in narrow rows; and present in jaws, palatine, and vomer. Preorbital bone has 3 spines over maxilla, with first greatly reduced and others strongly developed. Suborbital ridge lacks spinous points. Nasal, preocular, parietal, upper and lower posttemporal, cleithral, supracleithral, and pterotic spines are present. Preoperculum has 5 spines, with third the largest, fifth the smallest, and all about equally spaced. Gill rakers on first arch number 6 or 7 on upper limb and 9 to 11 on lower limb. Small slit is present behind fourth gill arch. Measurements are expressed as percent of SL: snout length 13%–15%, interorbital width 10%–15%, body depth 27%–52%. Orbit diameter is about half of snout length. Pectoral fin has 18 to 20 rays, with upper 1 or 2 rays unbranched,

middle rays branched, and lower rays unbranched. Dorsal fin has 12 spines and 9 or 10 rays. Anal fin has 3 spines and 6 (rarely 5 or 7) rays. Scales are small, cycloid, and present on head and body. Lateral line is covered by scales. Vertebrae number 24, and pyloric caecae number four or five. Gas bladder is absent or rudimentary. Adults are maroon and black, and juveniles are black. Mouth cavity is black with patches of bright orange and red in life. This species occurs between 274 and 732 m in the tropical to warm temperate Atlantic and the eastern and Central Pacific. Adult specimens are occasionally captured in midwater nets. In the western Atlantic it occurs in the northern Gulf of Mexico and off Honduras, Colombia, and Puerto Rico. Food consists of amphipods and midwater shrimps (Sergestes sp.). Maximum known size is 171 mm SL. references: Eschmeyer and Collette 1966; Eschmeyer 1969b; Uyeno and Sato 1983b; Poss and Eschmeyer 2002.

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SCORPAENIDAE 31

Helicolenus dactylopterus Blackbelly rosefish

(Delaroche, 1809)

Helicolenus dactylopterus is pale red, without cirri or tabs on head. Head is of moderate size, snout is short, and eye is large. Small teeth occur in jaws, palatine, and vomer. Preorbital bone has rounded lobes over maxilla. Suborbital ridge either lacks spines or has a single spine. Nasal, preocular, supraocular, postocular, and pterotic spines are poorly developed. Right and left tympanic spines are widely separated. Parietal spine is longer than nuchal spine. Preoperculum has 5 spines, with second the longest, first lacking supplemental spine and about equal to third, and fourth and fifth of moderate length. Gill rakers on first arch number 7 or 8 on upper limb and 16 to 18 on lower limb. Slit is lacking behind fourth gill arch. Measurements are expressed as percent of SL: head length 39%– 43%, snout length 8%– 10%, orbit diameter 13%–14%, interorbital width 4%–5%, jaw length 21%–22%, body depth 31%–33%. Pectoral fin rays number 19 (rarely 17 or 18). Dorsal fin has 12 spines and 11 to 13 rays. Anal fin has 3 spines and 5 rays (rarely 2 spines and 4 or 6 rays). Caudal fin is

very slightly emarginate. Body is covered with ctenoid scales. Vertebrae number 25, and gas bladder is absent. Dark markings are mainly concentrated on dorsal half of body. Young specimens have dark area at base of dorsal fin. Mouth and opercular cavity are black. Preserved specimens are straw colored with dark markings. This species occurs in temperate to tropical waters of the Atlantic Ocean. In the western Atlantic it occurs between 110 and 735 m, from Nova Scotia to Guyana, including the northern Gulf of Mexico, the Yucatán Channel, and the Bahamas. Food consists of benthic and pelagic crustaceans and ray-finned fishes, including myctophids. Maximum known size is 380 mm SL. references: Longley and Hildebrand 1941 (as Helicolenus maderensis); Bigelow and Schroeder 1953; Eschmeyer 1969b; Robins and Ray 1986; Boschung 1992; Cervigón 1993a; Poss and Eschmeyer 2002.

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Neomerinthe beanorum Spotwing scorpionfish

(Evermann and Marsh, 1900)

Neomerinthe beanorum is reddish brown, with spines on suborbital ridge and an elongated third dorsal spine. Head is of moderate size, snout is short, and eye is large. Teeth are small; granular; arranged in narrow bands; and present in jaws, palatine, and vomer. Preorbital bone has two spinous points over maxilla, with first directed ventrally or slightly forward. Suborbital ridge has three or four spinous points. Small nasal, preocular, supraocular, postocular, pterotic, parietal, nuchal, and cleithral spines are present. Preoperculum has 4 or 5 spines, with first the longest and bearing supplemental spine, second very small to absent, third pointing downward, fourth pointing forward, and fifth small to absent. Measurements are expressed as percent of SL: head length 44%–50%, snout length 12%–15%, orbit diameter 10%–14%, interorbital width 4%– 5%, jaw length 20%–24%, body depth 29%– 33%. Gill rakers on first arch number 5 or 6 on upper limb and 9 to 11 on lower limb. Slit be-

hind fourth gill arch is present. Pectoral fin has 17 rays (rarely 16 or 18), with rays 3 or 4 through 7 to 9 branched in specimens greater than 50 mm SL. Dorsal fin has 12 spines, with third greatly elongated, and 8 or 9 rays. Anal fin has 3 spines and 5 rays. Vertebrae number 24, and gas bladder is present. Dorsal half of body is mottled red and brown, pectoral fin has elongate dusky spots, midlength of spinous section of dorsal fin is darkly pigmented, and rayed section of dorsal fin and upper half of caudal fin are spotted. This species occurs between 91 and 375 m in the western Atlantic from the Bahamas; the Greater and Lesser Antilles; and the coasts of Central America, Colombia, and Venezuela. There are no verified records of this species in the Gulf of Mexico, but it may occur there. Maximum known size is 155 mm SL. references: Eschmeyer 1969b; Poss and Eschmeyer 2002.

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SCORPAENIDAE 33

Neomerinthe hemingwayi Spinycheek scorpionfish

Fowler, 1935

Neomerinthe hemingwayi is reddish brown, with a moderately long snout and spines on suborbital ridge. Head is of moderate size and somewhat pointed, and eye is less than snout length. Teeth are small; pointed; and arranged in narrow bands in jaws, palatine, and vomer. Preorbital bone has two spinous points over maxilla, with first directed downward and second directed posteriorly. Suborbital ridge has three or four spinous points. Small nasal, preocular, supraocular, postocular, pterotic, parietal, and nuchal spines are present. Preoperculum has 3 to 5 spines, with first the longest and bearing supplemental spine, second absent in all but very small specimens, third and fourth moderate to small, and fifth small or absent. Measurements are expressed as percent of SL: head length 42%– 48%, snout length 12%– 14%, orbit diameter 8%–11%, interorbital width 5%– 6%, jaw length 20%–22%, body depth 28%–33%. Gill rakers on first arch number 6 or 7 on upper limb and 9 to 12 on lower limb. Slit is present behind fourth arch.

Pectoral fin has 17 rays (rarely 16), with rays 3 through 7 or 8 branched (rarely rays 2 through 9 branched) in specimens greater than 50 mm SL. Dorsal fin has 12 spines and 10 rays (rarely 9). Anal fin has 3 spines and 5 rays. Scales are ctenoid. Vertebrae number 24, and gas bladder is present. Back and side are mottled red and brown; dorsal, caudal, and anal fins are spotted; and posterior part of lateral line bears three dark spots. This species occurs in the western North Atlantic from New Jersey to southern Florida and the Gulf of Mexico. In the Gulf of Mexico it occurs from Texas to Tampa Bay, FL, Veracruz, and Campeche between 31 and 230 m. Maximum known size is 315 mm SL.

references: Longley and Hildebrand 1941 (as S. tortugae); Springer and Bullis 1956 (as Pontinus castor or P. pollux); Bullis and Thompson 1965 (as P. pollux); Eschmeyer 1969b; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Poss and Eschmeyer 2002.

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34 SCORPAENIFORMES

Phenacoscorpius nebris Short-tube scorpionfish

Eschmeyer, 1965

Phenacoscorpius nebris is pinkish red, with strong spines on head. Teeth are small and arranged in narrow bands in jaws, palatine, and vomer. Preorbital bone has two spinous points over maxilla, with first rounded and second broadly pointed. Suborbital ridge has five to eight spinous points. Nasal, preocular, supraocular, postocular, frontal, anterior and posterior parietal, postorbital, pterotic, lower posttemporal, supracleithral, cleithral, suborbital, and preopercular spines are present. Preoperculum has 5 spines, with first the longest and bearing supplemental spine, and third, fourth, and fifth well developed. Measurements are expressed as percent of SL: head length 46%– 48%, snout length 12%–14%, orbit diameter 13%–14%, interorbital width 4%– 6%, jaw length 22%–23%, body depth 35%– 40%. Gill rakers on first arch number 5 or 6 on upper limb and 14 or 15 on lower limb. Slit behind fourth gill arch is very small. Pectoral fin

rays number 16 or 17, with upper 1 or 2 unbranched, adjacent 3 to 6 branched, and remaining unbranched. Dorsal fin has 12 spines and nine rays. Anal fin has 3 spines and five rays. Scales on body are ctenoid. Lateral line consists of several tubed scales behind head. Vertebrae number 25, and gas bladder is present. Saddle-shaped patches of dark pigment occur at dorsal fin origin, below center of spinous section of dorsal fin, between dorsal insertion and anal fin, and at base of caudal fin. Dark patch is present between fifth and ninth dorsal fin spines. This species occurs in the western North Atlantic from the north-central Gulf of Mexico and Venezuela between 348 and 476 m. Maximum known size is 84 mm SL.

references: Eschmeyer 1965a, 1969b; Robins and Ray 1986; Poss and Eschmeyer 2002.

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SCORPAENIDAE 35

Pontinus longispinis Goode and Bean, 1896 Longspine scorpionfish

Pontinus longispinis is light orange and marked by dusky spots on head and body, with a large spiny head and the third dorsal fin spine greatly elongated. Head is relatively slender, snout is moderately long, and eye is large. Occiput is relatively flat. Teeth are small and pointed and arranged in bands in jaws, palatine, and vomer. Preorbital bone has two spinous points over maxilla, with first pointing anteriorly and second pointing posteriorly. Suborbital ridge has three or four spinous points. Nasal, preocular, supraocular, postocular, tympanic, parietal, nuchal, pterotic, sphenotic, lower posttemporal, supracleithral, and cleithral spines are present. Preoperculum has 4 or 5 spines, with first the longest and bearing supplemental spine, second small or absent, third pointing ventrally or posteriorly, fourth present, and fifth acute. Measurements are expressed as percent of SL: head length 40%– 45%, snout length 10%– 13%, orbit diameter 11%–15%, interorbital width 3%–5%, jaw length 19%–21%, body depth 29%–34%. Gill rakers on first arch number 6 or 7 on upper limb and 8 to 10 on lower limb. Slit behind fourth arch is present. Pectoral fin has 17 unbranched rays (rarely 16 or 18). Dorsal fin has 12 spines, with third

greatly elongated in specimens larger than about 150 mm SL, and 9 rays (rarely 8 or 10). Anal fin has 3 spines and 5 rays. Supraocular tentacle is small or absent. Scales are ctenoid and occur on body, cheek, postorbital region, and occiput. Vertebrae number 24, and gas bladder is present. Upper section of head and body and rayed section of dorsal fin and caudal fin are covered with dusky spots. This species occurs between 77 and 440 m in the western Atlantic from South Carolina to Brazil, including the Gulf of Mexico. In the Gulf of Mexico it occurs from the Dry Tortugas and the Florida Panhandle to southern Texas and from Veracruz to Yucatán. It is generally captured over soft bottoms. Food consists of polychaetes, squids, euphausids, penaeid shrimps, porcellanid crabs, and rayfinned fishes. Shrimps and crabs are the most numerous food items. Maximum known size is 250 mm SL. references: Longley and Hildebrand 1941; Eschmeyer 1969b; Hoese and Moore 1977, 1998; Uyeno and Sato 1983b; Robins and Ray 1986; Cervigón 1991; Boschung 1992; Schaldach et al. 1997; Poss and Eschmeyer 2002.

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36 SCORPAENIFORMES

Pontinus rathbuni Goode and Bean, 1896 Highfin scorpionfish

Pontinus rathbuni is orangish red, with a spiny head and a short snout. Occiput is relatively flat. Teeth are small and arranged in narrow bands in jaws, palatine, and vomer. Preorbital bone has two spinous points over maxilla, with first pointing ventroposteriorly and second curved posteriorly. Suborbital ridge usually has four spinous points. Nasal, preocular, supraocular, postocular, tympanic, parietal, nuchal, pterotic, sphenotic, lower posttemporal, supracleithral, and cleithral spines are present. Preoperculum has 4 or 5 spines, with first the longest and bearing supplemental spine, second moderate to small and closer to first than to third, third directed posteriorly, fourth directed ventrally to ventroposteriorly, and fifth small or absent. Measurements are expressed as percent of SL: head length 42%– 48%, snout length 11%–14%, orbit length 11%–15%, interorbital width 3%–5%, jaw length 19%–23%, body depth 30%–38%. Gill rakers on first arch number six or seven on upper limb and eight or nine on lower limb. Slit behind fourth gill arch is present. Pectoral fin

has 17 unbranched rays (rarely 16 or 18). Dorsal fin has 12 spines, with third slightly elongated, and 9 rays. Anal fin has 3 spines and 5 rays. Supraorbital tentacle is thick, shorter than orbit, usually expanded distally, and occasionally split. Scales are ctenoid and occur on body, cheek, postorbital region, and occiput. Vertebrae number 24, and gas bladder is present. Upper section of body below dorsal fin is patterned with four saddles of dusky pigment. Dusky smudges may also occur on lateral line scales, axil of pectoral fin, head, supraorbital tentacle, pectoral fin, anal fin, and caudal fin. Smudges may be orange or red in life. This species occurs between 73 and 366 m in the western Atlantic from Virginia to Suriname, including the northern Gulf of Mexico. It is more common in the northeastern than in the northwestern Gulf of Mexico. Maximum known size is 197 mm SL. references: Eschmeyer 1969b; Uyeno and Sato 1983b; Robins and Ray 1986; Boschung 1992; Poss and Eschmeyer 2002.

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SCORPAENIDAE 37

Scorpaena agassizii Goode and Bean, 1896 Longfin scorpionfish

Scorpaena agassizii is uniformly reddish brown, with an occipital pit, relatively long pectoral fins, and cycloid scales. Snout is short, and eye is large. Small teeth are arranged in narrow bands in jaws, palatine, and vomer. Preorbital bone has two spinous points over maxilla, with second long and curved posteriorly in large specimens. Suborbital ridge has three or (rarely) four spines. Well-developed nasal, preocular, supraocular, postocular, frontal, anterior and posterior parietal, upper and lower posttemporal, supracleithral, postorbital, and opercular spines are present. Cleithral spine is small or absent. Preoperculum has 3 spines, with first bearing supplemental spine and reaching half the distance from base to opercular margin, and second smaller than third. Gill rakers on first arch number 4 to 6 on upper limb and 8 to 11 on lower limb. Slit behind fourth arch is absent. Measurements are expressed as percent of SL: head length 43%– 49%, snout length 8%–10%, orbit diameter 14%–20%, interorbital width 7%–11%, upper jaw length 20%–25%, body depth 33%–

39%. Pectoral fin has 18 to 21 rays, with uppermost unbranched, next 6 to 9 branched, and remainder unbranched. Dorsal fin has 12 spines (rarely 13) and 9 rays. Anal fin has 3 spines and 5 rays. Scales are cycloid, and lateral line scales are pored and number 23 to 25. Vertebrae number 24, and gas bladder is absent. Body pigment is concentrated into poorly defined spots behind head. This species occurs between 46 and 300 m in the western Atlantic from North Carolina to Brazil, including the northern and southern Gulf of Mexico. In the Gulf of Mexico it is most common off Florida, Alabama, and Mississippi, and it moves offshore during the spring and onshore during the fall. It is associated with soft to partially hard bottoms. Maximum known size is 155 mm TL. references: Longley and Hildebrand 1941; Eschmeyer 1965b, 1969b; Uyeno and Sato 1983b; Robins and Ray 1986; Cervigón 1991; Boschung 1992; Smith 1997; Poss and Eschmeyer 2002.

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38 SCORPAENIFORMES

Scorpaena albifimbria Evermann and Marsh, 1900 Coral scorpionfish

Scorpaena albifimbria is mottled brown and red, with a shallow occipital pit and 3 spines on suborbital ridge. Head is relatively large, snout of moderate length, and eye large. Teeth are small; arranged in narrow rows; and present in jaws, palatine, and vomer. Preorbital bone has two spinous points over maxilla. Suborbital ridge has spinous points below middle of eye, below posterior margin of eye, and at posterior end of ridge. Nasal, preocular, supraocular, postocular, frontal, anterior and posterior parietal, sphenotic, upper and lower posttemporal, cleithral, supracleithral, and pterotic spines are present. Preoperculum has 5 spines; first bears supplemental spine and extends twothirds of distance to opercular margin, second is closer to first and smaller than third, and fourth and fifth are of moderate size. Measurements are expressed as percent of SL: head length 47%–52%, snout length 10%–13%, orbit diameter 15%–17%, interorbital width 6%–7%, upper jaw length 24%–25%, body depth 39%– 44%. Gill rakers on first arch number 4 or 5 on upper limb and 7 to 10

on lower limb. Slit behind fourth gill arch is absent. Pectoral fin rays number 19 to 21, with uppermost unbranched, following 4 to 6 branched, and remaining 12 to 15 unbranched. Dorsal fin has 12 spines and 8 or 9 rays. Anal fin has 3 spines and 5 rays. Scales are cycloid, and lateral line scales are pored and number 23. Vertebrae number 24, and gas bladder is absent. Saddle of dark pigment is located above base of pectoral fin, distal section of pelvic fin is dark, and pectoral fin is dusky. This species occurs between the shoreline and 36 m in the western North Atlantic from southern Florida and Bermuda to the southern Caribbean Sea, including the southern Gulf of Mexico, the Bahamas, and the Antilles. It has been reported only once in the Gulf of Mexico, in southern Campeche Bay. Maximum known size is about 51 mm SL. references: Eschmeyer 1965b, 1969b; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Poss and Eschmeyer 2002.

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SCORPAENIDAE 39

Scorpaena bergii Evermann and Marsh, 1900 Goosehead scorpionfish

Scorpaena bergii is mottled dark brown, with a well-developed occipital pit and a single spine at end of suborbital stay. Head and snout are relatively short, and eye is of moderate size. Teeth are small and arranged in narrow bands in jaws, palatine, and vomer. Preorbital bone has two spinous points over maxilla. Nasal, preocular, supraocular, postocular, single or double sphenotic, anterior and posterior parietal, upper and lower posttemporal, supracleithral, pterotic, and cleithral spines are present. Preoperculum has 3 spines, with first bearing supplemental spine and extending from base to half the distance to opercular margin, and second longer than third and closer to first. Measurements are expressed as percent of SL: head length 39%– 46%, snout length 8%– 13%, orbit diameter 12%–16%, interorbital width 5%– 6%, upper jaw length 19%–24%, body depth 35%– 40%. Gill rakers on first arch number 3 or 4 on upper limb and 7 to 10 on lower limb. Slit behind fourth gill arch is lacking. Pectoral fin has 16 or 17 rays, with

some branched. Dorsal fin has 12 (rarely 11) spines and 9 rays. Anal fin has 3 spines and 5 rays. Scales are cycloid, and lateral line scales are pored and number 23. Vertebrae number 24, and gas bladder is absent. Dorsal fin has dark spot between third, fourth, or fifth to seventh or eighth fin spines; posterior section of dorsal fin and caudal fin bear three dark bands; pectoral fin is irregularly pigmented; and anal fin has three transverse dark bands. This species occurs between the shoreline and 75 m in the western Atlantic from New York to Brazil, including the eastern Gulf of Mexico and the Caribbean Sea. In the Gulf of Mexico it has been captured only off the Dry Tortugas. It is associated with turtle grass, coral reefs, rocky outcroppings, and sandy bottoms. Maximum known size is about 63 mm SL. references: Longley and Hildebrand 1941; Eschmeyer 1965b, 1969b; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997; Poss and Eschmeyer 2002.

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40 SCORPAENIFORMES

Scorpaena brasiliensis Barbfish

Cuvier, 1829

Scorpaena brasiliensis is mottled red, orange, or blue above and pale below, with a welldeveloped occipital pit and small to no upper posttemporal spine. Head and snout are moderately short, and eye is of moderate size. Teeth are small and arranged in narrow bands in jaws, palatine, and vomer. Preorbital bone has two spinous points over maxilla. Suborbital ridge has three spinous points, with first two blunt. Nasal, preocular, supraocular, postocular, anterior and posterior parietal, lower posttemporal, one or two sphenotic, supracleithral, and pterotic spines are present. Cleithral spine is small in juveniles and often absent in adults. Preoperculum has 5 spines, with first bearing small supplemental spine and extending to middle of operculum, second equal or subequal to third, and fourth and fifth moderate to small. Measurements are expressed as percent of SL: head length 39%– 47%, snout length 9%–11%, orbit diameter 10%–14%, interorbital width 4%–7%, jaw length 20%– 24%, body depth 32%– 40%. Gill rakers on first arch number 4 or 5 on upper limb and 8 to 10 on lower limb. Slit behind fourth gill arch is lacking. Pectoral fin has 18 to 20 rays, with uppermost unbranched, next 7 to 10 branched, and remainder unbranched in specimens greater than 50 mm SL. Dorsal fin has

12 spines and 8 (rarely 9) rays. Anal fin has 3 spines and 5 rays. Supraorbital tentacle is well developed but shorter than orbit diameter. Scales are cycloid, and lateral line scales are pored and number 23 or 24. Vertebrae number 24, and gas bladder is absent. Large dark spot is present on upper flank behind operculum, several spots occur on pale axil of pectoral fin, and two dark bands are located on tail. This species occurs between the shoreline and 163 m in the western Atlantic from Virginia to southern Brazil, including the Gulf of Mexico and the Greater and Lesser Antilles. It is common in the northern Gulf of Mexico and also occurs off Veracruz, Campeche, and Yucatán. Formerly it occurred in Bermuda, but it has not been observed or collected there since 1927 (Smith-Vaniz et al. 1999). This species is generally associated with soft bottoms but is occasionally found on coral reefs. Food consists of crabs, shrimps, and ray-finned fishes. Maximum known size is 250 mm SL.

references: Longley and Hildebrand 1941; Eschmeyer 1965b, 1969b; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Uyeno and Sato 1983b; Robins and Ray 1986; Cervigón 1991; Boschung 1992; Schaldach et al. 1997; Smith 1997; Poss and Eschmeyer 2002.

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SCORPAENIDAE 41

Scorpaena calcarata Goode and Bean, 1882 Smoothhead scorpionfish

Scorpaena calcarata is plain reddish, with a very shallow occipital pit and no supplemental preopercular spine. Head and snout are moderately short, and eye is of moderate size. Teeth are small and arranged in narrow bands in jaws, palatine, and vomer. Preorbital bone has two free spinous points over maxilla. Suborbital ridge has two or three spinous points. Nasal, preocular, supraocular, postocular, frontal, anterior and posterior parietal, lower posttemporal, supracleithral, and pterotic spines are present. Sphenotic bone has several closely located spinules. Preoperculum has 5 spines, with first extending half the distance to opercular flap, second smaller than third, fourth flat, and fifth flat and often curved ventrally. Gill rakers on first arch number 4 or 5 on upper limb and 7 to 11 on lower limb. Slit behind fourth gill arch is absent. Measurements are expressed as percent of SL: head length 40%– 46%, snout length 9%–12%, orbit length 12%–16%, interorbital width 5%– 6%, upper jaw length 19%–23%, body depth 32%–38%. Pectoral fin has 19 to 21 rays, with uppermost unbranched, following 5 to 7 branched, and remaining 11 to 15 unbranched.

Dorsal fin has 12 spines and 9 (rarely 7 or 8) rays. Anal fin has 3 spines and 5 rays. Scales are cycloid, and lateral line scales are pored and number 22 to 24+0 –1. Vertebrae number 24, and gas bladder is absent. Upper body has three slightly darker patches along dorsal fin base, upper part of pectoral fin is slightly pigmented, and ventral body is pale. This species occurs from near shore to 91 m in the western Atlantic from North Carolina to Brazil, including the Gulf of Mexico and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico, and in the northern Gulf it is most common between 20 and 60 m. Food consists of mysid shrimps, mantas shrimps, penaeid shrimps, alphaeid shrimps, and portunid crabs. Maximum known size is 125 mm SL.

references: Longley and Hildebrand 1941; Eschmeyer 1965b, 1969b; Hoese and Moore 1977, 1998; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Cervigón 1991; Boschung 1992; Schaldach et al. 1997; Smith 1997; Poss and Eschmeyer 2002.

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42 SCORPAENIFORMES

Scorpaena dispar Longley and Hildebrand, 1940 Hunchback scorpionfish

Scorpaena dispar is reddish, with three or four spinous points on preorbital bone and a supplemental preopercular spine. Head is of moderate size, snout is moderately long, and eye is of moderate size. Teeth are small and arranged in narrow bands in jaws, palatine, and vomer. Suborbital ridge has three spinous points (rarely two or four). Nasal, preocular, supraocular, postocular, frontal, one or two sphenotic, anterior and posterior parietal, upper and lower posttemporal, cleithral, supracleithral, and pterotic spines are present. Preoperculum has 5 spines: the first bears a supplemental spine and extends less than half the distance across operculum; the second, third, and fourth are of moderate size; and the fifth is minute. Gill rakers on first arch number 5 on upper limb and 10 to 12 on lower limb. Slit behind fourth gill arch is absent. Measurements are expressed as percent of SL: head length 43%– 48%, snout length 11%–15%, orbit diameter 11%–14%, interorbital width 5%– 6%, upper jaw length 20%–24%, body depth 32%–38%. Pectoral fin has 17 to 19 rays, with uppermost unbranched, next 6 to 8 branched,

and following 8 to 12 unbranched in specimens greater than 50 mm SL. Dorsal fin has 12 spines and 9 (rarely 8) rays. Anal fin has 3 spines and 5 rays. Scales are cycloid, and lateral line scales are pored and number 22 or 23. Supraorbital tentacle is long and slender. Vertebrae number 24, and gas bladder is absent. Upper body has two dark spots behind head, and pectoral, dorsal, and caudal fins bear small dark spots. Dark spots also occur on medial surface of pectoral fin. This species occurs between 36 and 125 m in the western Atlantic from Florida to Brazil, including the Gulf of Mexico and the Caribbean Sea. It is common throughout the Gulf of Mexico but is rare to absent in the Antilles. Maximum known size is about 181 mm SL.

references: Longley and Hildebrand 1941; Eschmeyer 1965b, 1969b; Hoese and Moore 1977, 1998; Castro-Aguirre and Márquez-Espinoza 1981; Uyeno and Sato 1983b; Robins and Ray 1986; Cervigón 1991; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Poss and Eschmeyer 2002.

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SCORPAENIDAE 43

Scorpaena elachys Dwarf scorpionfish

Eschmeyer, 1965

Scorpaena elachys is reddish, with a tympanic spine and one or more spines on suborbital ridge. Head is relatively large, snout is of moderate length, and eye is of moderate size. Teeth are small and arranged in narrow bands in jaws, palatine, and vomer. Preorbital bone has two spinous points over maxilla. Suborbital ridge occasionally has two small spinous points posteriorly. Nasal, preocular, supraocular, postocular, one or two sphenotic, frontal, anterior and posterior parietal, pterotic, upper and lower posttemporal, and cleithral spines are present. Preoperculum has 5 spines, with first bearing small supplemental spine and extending less than half the width of operculum, second smaller than third, fourth broader and shorter than fifth, and fifth pointing anteroventrally. Gill rakers on first arch number 4 or 5 on upper limb and 10 or 11 on lower limb. Slit behind fourth gill arch is absent. Measurements are expressed as percent of SL: head length 47%–51%, snout length 11%–13%, orbit diameter 14%–16%, interorbital width

about 5%, upper jaw length 24%–27%, body depth 38%– 41%. Pectoral fin has 17 (rarely 16) rays, with uppermost unbranched, second or third to fifth or sixth branched, and remainder unbranched in specimens greater than 50 mm SL. Dorsal fin has 12 spines and 8 or 9 rays. Anal fin has 3 spines and 5 rays. Scales are cycloid, and lateral line scales are pored and number 23. Supraorbital tentacle is relatively short. Vertebrae number 24, and gas bladder is absent. Upper body has several narrow light and dark lines below dorsal fin, and inner surface of operculum is blackish. This species occurs between 55 and 90 m in the western Atlantic from Florida to the Antilles, including the eastern Gulf of Mexico off southern Florida. Maximum known size is about 50 mm SL. references: Eschmeyer 1965b, 1969b; Robins and Ray 1986; Smith 1997; Poss and Eschmeyer 2002.

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44 SCORPAENIFORMES

Scorpaena grandicornis Cuvier, 1829 Plumed scorpionfish

Scorpaena grandicornis is dark brown on body and fins, with long supraocular tentacles and a well-developed occipital pit. Head and snout are relatively short, and eye is of moderate size. Teeth are small and arranged in narrow bands in jaws, palatine, and vomer. Preorbital bone has two spinous points over maxilla. Suborbital ridge has two or three spinous points. Nasal, preocular, supraocular, postocular, anterior and posterior parietal, one or two sphenotic, upper and lower posttemporal, cleithral, supracleithral, and pterotic spines are present. Preoperculum has 3 spines; the first bears a supplemental spine and extends less than half the width of operculum, and the second is about as long or slightly longer than the third. Gill rakers on first arch number 4 to 7 on upper limb and 7 to 10 on lower limb. Slit behind fourth gill arch is absent. Measurements are expressed as percent of SL: head length 40%– 43%, snout length 8%–10%, orbital diameter 10%–13%, interorbital width 6%– 8%, upper jaw length 18%–22%, body depth 38%– 41%. Pectoral fin has 18 or 19 rays, with uppermost unbranched, next 7 to 9 branched, and re-

mainder unbranched in specimens greater than 50 mm SL. Dorsal fin has 12 spines and 9 rays. Anal fin has 3 spines and 5 rays. Scales are cycloid, and lateral line scales are pored and number 23. Vertebrae number 24, and gas bladder is absent. Head and anterior part of body have white spots, and caudal fin has three dark bands. Axil of pectoral fin has patch of white spots on tan or dark background. This species occurs in shallow water in the North Atlantic from Florida and Bermuda to southern Brazil, including the eastern Gulf of Mexico and the Caribbean Sea. In the Gulf of Mexico it is known from only the Dry Tortugas. Formerly it occurred in Bermuda, but it has not been observed or collected there in nearly 100 years (Smith-Vaniz et al. 1999). It is common in sea-grass beds, channels, and bays. Maximum known size is about 140 mm SL. references: Longley and Hildebrand 1941; Eschmeyer 1965b, 1969b; Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986; Smith 1997; Castro-Aguirre et al. 1999; Poss and Eschmeyer 2002.

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SCORPAENIDAE 45

Scorpaena inermis Cuvier, 1829 Mushroom scorpionfish

Scorpaena inermis is mottled red, with fleshy, mushroom-shaped growths on upper surface of pupil of eyes and no occipital pit. Head is relatively large, snout is of moderate length, and eye is relatively large. Teeth are small and arranged in bands in jaws, palatine, and vomer. Preorbital bone has two spinous points over maxilla. Suborbital ridge has two (rarely one) spinous points. Nasal spine is blunt. Preocular, supraocular, posttemporal, supracleithral, and pterotic spines are poorly developed. Preoperculum has 3 spines, with first lacking a supplemental spine and extending less than half the width of operculum, and second generally smaller than third. Gill rakers on first arch number three to five on upper limb and six to eight on lower limb. Slit behind fourth gill arch is absent. Measurements are expressed as percent of SL: head length 46%–50%, snout length 10%–12%, orbit diameter 14%–18%, interorbital width 3%– 4%, upper jaw length 24%–26%, body depth 31%–37%. Pectoral fin has 19 to 21 rays, with uppermost unbranched, next 5 or 6 branched, and remaining 13 or 14 unbranched in specimens greater

than 50 mm SL. Dorsal fin has 12 spines and 7 or 8 rays. Anal fin has 3 spines and 5 rays. Scales are cycloid, and lateral line scales are pored and number 22 to 24. Vertebrae number 24, and gas bladder is absent. Dark pigment is generally concentrated below middle of spinous part of dorsal fin, below rayed section of dorsal fin, and on dorsal part of caudal peduncle. Brown spots often occur behind head below lateral line. Pelvic fin is dusky or has dark tip, pectoral fin is dusky along distal margin, and caudal fin has two diffuse transverse bands. This species occurs between the shoreline and 73 m in the western Atlantic from Georgia to Venezuela, including the eastern Gulf of Mexico, the Bahamas, and the Antilles. It is generally associated with clear water over sandy bottoms or coral reefs. Maximum known size is 77 mm SL. references: Longley and Hildebrand 1941; Eschmeyer 1965b, 1969b; Uyeno and Sato 1983b; Robins and Ray 1986; Cervigón 1991; Boschung 1992; Smith 1997; Poss and Eschmeyer 2002.

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46 SCORPAENIFORMES

Scorpaena plumieri Bloch, 1789 Spotted scorpionfish

Scorpaena plumieri is olive brown, with three spinous points on suborbital bone and a welldeveloped occipital pit. Head is of moderate size, snout is moderately long, and eye is relatively small. Teeth are small and arranged in narrow bands in jaws, palatine, and vomer. Second preorbital spinous point is closer to anterior spine than to posterior spine, and specimens less than 50 mm SL may have only two spinous points. Nasal, preocular, supraocular, postocular, anterior and posterior parietal, upper and lower posttemporal, pterotic, cleithral, and supracleithral spines are present. Preoperculum has 5 spines, with first bearing a supplemental spine and extending less than half the width of the operculum, second longer than third, and fourth and fifth variable in development. Gill rakers on first arch number 4 to 6 on upper limb and 8 to 12 on lower limb. Slit behind fourth gill arch is absent. Measurements are expressed as percent of SL: head length 43%– 47%, snout length 11%–14%, orbit diameter 8%–11%, interorbital width 8%–10%, upper jaw length 21%–25%, body depth 35%– 42%. Pectoral fin has 18 to 21 rays, with uppermost unbranched, next 7 to 10 branched, and remaining 9 to 12 unbranched in specimens greater than 50 mm SL. Dorsal fin has

12 spines and 9 rays. Anal fin has 3 spines and 5 rays. Scales are cycloid, and lateral line scales are pored and number 22 to 24. Dark pigment is concentrated between bases of rayed section of dorsal fin and anal fin, and caudal peduncle is pale. Fins are banded or blotched with dark pigment on pale background, caudal fin has three distinct bars, and axil of pectoral fin is black with white spots. This species occurs in the western Atlantic and eastern Pacific over rocky areas from near shore to 55 m. In the western Atlantic it occurs from Massachusetts and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It occurs throughout the Gulf of Mexico and is most abundant in shallow water, especially on reefs, jetties, and oil platforms. Maximum known size is 430 mm SL.

references: Longley and Hildebrand 1941; Eschmeyer 1965b, 1969b; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Castro-Aguirre and Márquez-Espinoza 1981; Uyeno and Sato 1983b; Robins and Ray 1986; Cervigón 1991; Boschung 1992; Smith 1997; Castro-Aguirre et al. 1999; Poss and Eschmeyer 2002.

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SCORPAENIDAE 47

Scorpaenodes caribbaeus Reef scorpionfish

Meek and Hildebrand, 1928

Scorpaenodes caribbaeus is mottled olive brown and yellow, with small spines just below suborbital spines and spinous procurrent caudal fin rays. Head is of moderate size, snout is relatively short, and eye is relatively large. Occipital pit is absent. Teeth are small and arranged in narrow bands in jaws and vomer. Preorbital bone has three lobes, with second and third over maxilla. Suborbital ridge has two to five spinous points and one to six small additional spines below main series. Nasal, preocular, supraocular, postocular, tympanic, parietal, nuchal, sphenotic, pterotic, upper and lower posttemporal, cleithral, and supracleithral spines are present. Spines are also present near end of interorbital ridge and on frontal bone (two coronal spines). Preoperculum has 3 (rarely 4) spines, with first bearing a supplemental spine and extending less than half the width of the operculum and second closer to first than to third. Gill rakers on first arch number five to eight on upper limb and eight or nine on lower limb. Small slit behind fourth gill arch is present. Measurements are expressed as percent of SL: head length 42%– 47%, snout length 10%–12%, orbit diameter 12%–15%, interorbital width 5%–7%, upper jaw length

22%–24%, body depth 35%– 40%. Pectoral fin has 18 to 20 rays, with uppermost and occasionally next unbranched, next 5 to 7 branched, and remaining unbranched in specimens longer than 45 mm SL. Dorsal fin has 13 spines and 9 (rarely 8) rays. Anal fin has 3 spines and 4 or 5 rays. Scales are ctenoid, occur on head, and number 40 to 45 in vertical rows. Vertebrae number 24, and gas bladder is present. Pectoral, dorsal, and caudal fins are spotted, and occasionally dark spots occur on cheek and operculum. This species occurs from the shoreline to 28 m in the western Atlantic from Florida and Bermuda to Venezuela, including the northwestern and southern Gulf of Mexico, the Bahamas, and the Antilles. It has been recorded only once from the northwestern Gulf, at the West Flower Garden Bank. This species is associated with hard substrates, including coral reefs. Maximum known size is 85 mm SL. references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Eschmeyer 1969a,b; Bright and Cashman 1974; Castro-Aguirre and MárquezEspinoza 1981; Robins and Ray 1986; Cervigón 1991; Smith 1997; Poss and Eschmeyer 2002.

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48 SCORPAENIFORMES

Scorpaenodes tredecimspinosus Deepreef scorpionfish

(Metzelaar, 1919)

Scorpaenodes tredecimspinosus is brown spotted, with a single row of spines on suborbital ridge and spinous procurrent caudal fin rays. Head and snout are relatively short, and eye is relatively large. Occipital pit is absent. Teeth are small and arranged in narrow bands in jaws and vomer. Preorbital bone has three lobes, with second and third over maxilla. Suborbital ridge has one to three spinous points. Nasal, preocular, supraocular, postocular, tympanic, parietal, nuchal, sphenotic, pterotic, upper and lower posttemporal, cleithral, and supracleithral spines are present. Spines near end of interorbital ridge and coronal spines on temporal bone are often absent. Preoperculum has 3 (rarely 4) spines, with the first bearing a supplemental spine and extending less than half the width of the operculum, and the second closer to first than to third. Gill rakers on first arch number four to six on upper limb and seven to nine on lower limb. Small slit behind fourth gill arch is present. Measurements are expressed as percent of SL: head length 40%– 44%, snout length 9%–11%, orbit diameter

12%–15%, interorbital width 4%–7%, upper jaw length 24%–27%, body depth 34%– 40%. Pectoral fin has 16 to 18 rays, with uppermost and occasionally next unbranched, following 4 to 6 branched, and remainder unbranched. Dorsal fin has 13 spines and 9 (rarely 8) rays. Anal fin has 3 spines and 4 or 5 rays. Scales are ctenoid, occur on head and body, and number 40 to 45 in vertical rows. Vertebrae number 23, and gas bladder is present. Color is olive brown or yellow, with dusky pigment on head and body. Fins are transparent except for black spot at end of spinous part of dorsal fin. This species occurs between 7 and 82 m in the western Atlantic from North Carolina and the Bahamas to Venezuela, including the eastern Gulf of Mexico and the Antilles. It is associated with rocky or coral reefs. Maximum known size is about 45 mm SL. references: Longley and Hildebrand 1941 (as Scorpaenodes floridae); Böhlke and Chaplin 1968; Eschmeyer 1969a,b; Robins and Ray 1986; Cervigón 1991; Smith 1997; Poss and Eschmeyer 2002.

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SCORPAENIDAE 49

Setarches guentheri Johnson, 1862

Setarches guentheri is orangish red, with head bones cavernous and weakly ossified, and lateral line consisting of a continuous trough covered by thin, membranous scales. Head is of moderate length, snout is relatively long, and eye is of moderate size. Teeth are small and arranged in narrow bands in jaws, palatine, and vomer. Preorbital bone has moderate-sized spines. Suborbital ridge lacks spinous points. Nasal, preocular, parietal, upper and lower posttemporal, pterotic, cleithral, and supracleithral spines are present. Preoperculum has 4 or 5 spines of similar size. Gill rakers number 2 to 5 on upper limb and 8 to 11 on lower limb. Measurements are expressed as percent of SL: snout length 10%–14%, interorbital width 7%–9%, body depth 28%– 40%. Pectoral fin has 20 to 25 rays, with uppermost 1 or 2 unbranched, next 12 to 16 branched, and remainder unbranched. Dorsal fin has 12 spines and 9 or 10 rays (for western Atlan-

tic specimens). Anal fin has 3 spines and 4 to 6 rays. Scales are cycloid and small. Vertebrae number 24, and gas bladder is well developed. Head and body are covered with reddish black spots. Juveniles are usually reddish gray, with black specks. This species occurs in the Atlantic, Indian, and western Pacific Oceans. In the western Atlantic it occurs generally between 220 and 760 m, from New Jersey to northern Brazil, including the Gulf of Mexico and the Caribbean Sea. It has been reported from as shallow as 18 m off Alabama in the Gulf of Mexico. Food consists of caridean shrimps and amphipods. Apparently this species feeds on pelagic crustaceans near the bottom. Maximum known size is 250 mm SL. references: Eschmeyer and Collette 1966; Eschmeyer 1969b; Uyeno and Sato 1983b; Cervigón 1991; Boschung 1992; Poss and Eschmeyer 2002.

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50 SCORPAENIFORMES

Trachyscorpia cristulata (Goode and Bean, 1896) Atlantic thornyhead

Trachyscorpia cristulata is dusky on back and sides, with well-developed spines on head and no occipital pit. Head is relatively large, snout is short, and eye is moderately large. Teeth are small and arranged in narrow bands in jaws, palatine, and vomer. Preorbital bone has 2 spines over maxilla, with first directed posterolaterally and second directed posteriorly. Suborbital ridge has many sharp-pointed spines. Nasal, preocular, supraocular, postocular, tympanic, parietal, nuchal, pterotic, sphenotic, lower posttemporal, cleithral, and supracleithral spines are present. Preoperculum has 4 or 5 spines, with first longest and bearing a supplemental spine, second slightly closer to first than to third, fourth present, and fifth very small to absent. Slit behind fourth gill arch is absent. Measurements are expressed as percent of SL: head length 46%– 49%, snout length 10%–11%, orbit diameter 11%–16%, interorbital width 4%–7%, upper jaw length 22%–24%, body depth 32%–36%. Pectoral

fin is wedge shaped and has 21 to 24 rays, with uppermost unbranched, next 10 to 12 branched, and remainder unbranched. Upper pectoral fin rays are longest. Dorsal fin has 12 spines and 9 (rarely 8) rays. Anal fin has 3 spines and 5 rays. Scales are ctenoid and occur on body, cheek, postorbital region, and occiput. Lateral line is covered with tubed scales. Vertebrae number 25, and gas bladder is absent. Body is darkest below third or fourth to tenth dorsal spines and below rayed section of dorsal fin. Fins are often dusky. This species occurs in the Atlantic Ocean. The western Atlantic form is considered a separate subspecies, Trachyscorpia cristulata cristulata, and occurs between 130 and 1,100 m, from Massachusetts to the Straits of Florida and in the northern Gulf of Mexico. Maximum known size is about 400 to 500 mm SL. references: Eschmeyer 1969b; Robins and Ray 1986; Poss and Eschmeyer 2002.

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51

TRIGLIDAE

Searobins

Searobins are robust and elongate, with a large, rugose, and bony head and 3 lowermost rays of pectoral fins thickened and free of membranes. Snout is broad and flat, moderately to strongly oblique, and of moderate length. Lachrymal bone is moderately projected as flattened rostrum that often bears short rostral projections or rostral exsertions on each side. Mouth is short and terminal to slightly inferior or subterminal. Barbels are absent on lip, chin, and mandible. Villiform teeth are present in jaws, palatine, and vomer. Tongue is present. Premaxilla is protractile, and maxilla slips under preorbital bone. Eye is moderate to rather small in size and laterally located. Bony ridge (bony stay), formed by infraorbital bones, is generally present below eye and extends from just behind mouth to preoperculum. Preoperculum and operculum have well-developed spines. Gill membrane is free of isthmus or narrowly joined to isthmus anteriorly. Gills are present on four arches, with large slit behind fourth arch. Pectoral fin is fanlike, with 11 to 16 attached rays in addition to swollen free rays. First dorsal fin has 7 to 11 spines, and second dorsal fin has 10 to 19 rays. Pelvic fin is thoracic and has 1 spine and 5 rays. Anal fin is located below second dorsal fin and has 0 or 1 spine and 11 to 18 rays. Caudal peduncle is moderately slender, and caudal fin is narrow and truncate. Body is covered with ctenoid scales. Physoclistous gas bladder and pyloric caecae are present. Gas bladder is bilobed. Searobins occur worldwide in tropical to temperate seas. All species are benthic on continental and insular shelves and slopes, mostly on sandy to muddy bottoms. The enlarged and free pectoral fin rays are innervated with sensory nerves and apparently used to locate prey. The gas bladder generally has intrinsic and extrinsic muscles that produce drumming sounds when contracted. These sounds may be used in reproduction to attract mates. All species are oviparous and produce planktonic eggs ranging from 1.0 to 1.7 mm in diameter. There are about 125 species in eight genera, 19 species in two genera occur in the western Central Atlantic, and 14 or 15 species in two genera occur in the Gulf of Mexico. references: Washington, Eschmeyer, et al. 1984; Washington, Moser, et al. 1984; Heemstra 1986a; Hureau and Litvinenko 1986b; Eschmeyer 1994; Nelson 1994; Richards 1996; Smith 1997.

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52 SCORPAENIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Richards and Miller 2002a) 1a. Dorsal fin spines normally 11 (rarely 10 or 12); dorsal fin rays 11; first 1 or 2 dorsal fin spines often elongated and filamentous in males (except B. brachychir); opercular membrane scaleless (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . 2

FIG 2

1b. Dorsal fin spines normally 10 (rarely 9 or 11), posterior dorsal fin spines short and possibly difficult to see; dorsal fin rays 12 or 13; first 1 or 2 dorsal fin spines not elongated and filamentous; opercular membrane partially scaled above opercular spine. . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2a. Cleithral spine distinctly extending beyond tip of opercular spine; chest scaled; rostral projection extending well beyond snout . . . . . . . . . . . . . . . . . Bellator militaris p. 57 2b. Cleithral spine not extending beyond tip of opercular spine; chest naked; rostral projection barely extending beyond snout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. First free pectoral fin ray shorter than remainder of fin; eye with tabs or tentacles on upper surface; nasal spine present; maxilla not extending to anterior margin of eye; pectoral fin with alternating patches of light and dark pigment, appearing as brown patches on uppermost pectoral fin rays (often resemble bands). . . . . . . . . . . . Bellator egretta p. 56 3b. First free pectoral fin ray distinctly longer than remainder of fin; eye without tabs or tentacles; nasal spine absent; maxilla extending beyond anterior margin of eye; pectoral fin dusky or with elongate black spot on upper portion and white posterior margin . . . . . . . . Bellator brachychir p. 55 4a. Mouth terminal; lower jaw with small bony knob at symphysis; pectoral fin shorter than head length and not extending to anal fin origin; body distinctly silvery; lower preopercular spine extending only to distal end of operculum; body cavity extending distally to anal fin origin . . . . . . . . . . . . . . . . . . . . . . . . . . . Prionotus stearnsi p. 68

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TRIGLIDAE 53

4b. Mouth subterminal; lower jaw lacks bony knob; pectoral fin as long as or longer than head length and extending beyond anal fin origin; body coloration variable but not silvery; preopercular spine extending beyond operculum; body cavity terminates at anal fin origin . . . . . . . . . . . . . 5 5a. Nasal and supraocular cirri present . . . . . . . . . . . . . . . . . 6 5b. Nasal and supraocular cirri absent . . . . . . . . . . . . . . . . . . 7 6a. Pectoral fin rounded, upper 2 rays not extending to caudal fin base as filaments . . . . . . . . . . . Prionotus ophryas p. 62 6b. Pectoral fin not rounded, upper 2 rays produced and extending to caudal fin base as filaments . . . . . . . . . . . . . . . . . . . . . . . . . . Prionotus murielae p. 61 7a. First dorsal fin with two non-ocellated spots, one between first and second spines and other between fourth and fifth spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7b. First dorsal fin without two non-ocellated spots, either with a single ocellated spot or with no distinct spot . . . . . 9 8a. Throat (gular region) naked; joined pectoral fin rays 12 to 14 (usually 13); gill rakers on lower limb (including angle) 10 to 13 (usually 11); branchiostegal rays white . . . . . . . . . . . . . . . . . . . . . . . . . . . Prionotus scitulus p. 67 8b. Throat (gular region) entirely scaled; joined pectoral fin rays 14 or 15 (usually 14); gill rakers on lower limb (including angle) 8 to 11 (usually 9); branchiostegal rays dusky or black . . . . . . . . . . . . . . . . . . . . . . . . Prionotus martis p. 60 9a. First dorsal fin with ocellated spot between fourth and fifth spine . . . . . . . . . . . . . . . . . . Prionotus longispinosus p. 59 9b. First dorsal fin without ocellated spot . . . . . . . . . . . . . . 10 10a. Pectoral fin emarginate, with uppermost joined rays about same length as lowermost joined rays . . . . . . . . . . . . . . . . . . . . . . . . . . Prionotus paralatus p. 63 10b. Pectoral fin rounded or with lowermost rays much longer than uppermost joined rays . . . . . . . . . . . . . . . . . . . . . . 11 11a. Pectoral fin with lowermost joined rays much longer than uppermost rays. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11b. Pectoral fin rounded, lowermost rays not longer than uppermost attached rays. . . . . . . . . . . . . . . . . . . . . . . . . 13 12a. Posterior margin of pectoral fin concave; pectoral fin with black bands; nasal spine present. . . . Prionotus alatus p. 58 12b. Posterior margin of pectoral fin straight or obliquely truncate; pectoral fin black with blue ventral margin; nasal spine absent . . . . . . . . . . . . . . . . . . . Prionotus rubio p. 66 13a. Dark band or lines extending across pectoral fin; two lateral stripes or series of spots extending partially or totally from head to caudal peduncle . . . . Prionotus tribulus p. 69 13b. No dark bands or lines extending across pectoral fin; no lateral stripes on body . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14a. Uppermost free pectoral fin ray not reaching distal end of pelvic fin; pectoral fin covered with bright blue or dark

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54 SCORPAENIFORMES

ocellated spots; ventral pectoral fin margin dark or brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prionotus roseus p. 65 14b. Uppermost free pectoral fin ray extending to distal end of pelvic fin; pectoral fin without ocellated spots; ventral pectoral fin margin blue . . . . . . . . . Prionotus punctatus p. 64

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TRIGLIDAE 55

Bellator brachychir (Regan, 1914)

Bellator brachychir has poorly developed rostral projections, a long opercular spine, and a relatively long upper pectoral free ray. Head is relatively large and moderately broad, and snout is of moderate length. Rostral projection is broad and triangular shaped and bears several anteriorly directed spines. Preopercular spine lacks supplemental spine. Opercular spine extends beyond cleithral spine. Gill rakers on first arch number 1 to 3 on upper limb and 12 to 17 on lower limb. Measurements are expressed as percent of SL: head length 37%– 45%, snout length 12%–15%, upper jaw length 14%–20%, orbit diameter 12%–15%, interorbital width 3%– 6%, length of joined pectoral fin 23%–36%, length of first free pectoral fin ray 31%– 41%, body depth 26%– 32%. Joined pectoral fin rays number 12. First dorsal fin has 10 or 11 spines, with first 3 or 4 strongly serrated, and second dorsal fin has 11 rays, with first serrated at base. Anal fin

rays number 10 or 11. Body is covered with small ctenoid scales. Nape is scaled, and breast and usually anterior section of belly are naked. Color is brown on upper half of body and white on lower half. Spinous dorsal fin is brown, with black spot between fourth and fifth spines. Pectoral fin is dusky and has large black spot between third and fifth fin rays. This species occurs between 27 and 366 m in the western Atlantic from North Carolina to Uruguay, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it has been captured off the western coast of Florida and in the Bay of Campeche. Maximum known size is 160 mm SL. references: Ginsburg 1950; Cervigón 1966, 1991; Lewis and Yerger 1976; Robins and Ray 1986; Miller and Richards 1991a; Smith 1997; Richards and Miller 2002a.

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56 SCORPAENIFORMES

Bellator egretta (Goode and Bean, 1896) Streamer searobin

Bellator egretta has poorly developed rostral projections, a relatively long opercular spine, and papillae on orbits. Head is relatively long and broad, and snout is long. Rostral projection is short and has a rounded, granular margin. Preopercular spine lacks supplemental spine. Opercular spine extends beyond cleithral spine. Gill rakers on first arch number 1 to 3 on upper limb and 11 to 17 on lower limb. Measurements are expressed as percent of SL: head length 37%– 43%, snout length 17%– 20%, upper jaw length 12%–15%, orbit diameter 9%–11%, interorbital width 4%– 6%, joined pectoral fin length 26%–33%, first free pectoral fin ray length 21%–27%, body depth 20%–28%. Joined pectoral fin rays number 12. First dorsal fin has 10 or 11 spines, and second dorsal fin has 11 rays. First dorsal fin terminates as long filament in some specimens. Anal fin rays number 11. Body is covered with small ctenoid scales. Nape and breast are

naked, and belly is naked anteriorly but often scaled posteriorly. Color in life is reddish or yellowish brown on upper half of body and light on lower half, with sides banded. First dorsal fin has black spot or smudge between fourth and fifth dorsal spines. Pectoral fin is dusky, especially medial aspect, with upper 3 rays banded with dark and tan. This species occurs between 40 and 230 m in the western Atlantic from North Carolina to Barbados, including the Gulf of Mexico. In the Gulf of Mexico it occurs from the west coast of Florida to Alabama. Maximum known size is 150 mm SL.

references: Longley and Hildebrand 1941; Ginsburg 1950; Bullis and Thompson 1965; Lewis and Yerger 1976; Darcy and Gutherz 1984; Robins and Ray 1986; Cervigón 1991; Boschung 1992; Miller and Richards 1991a; Smith 1997; Richards and Miller 2002a.

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TRIGLIDAE 57

Bellator militaris Horned searobin

(Goode and Bean, 1896)

Bellator militaris has well-developed diverging rostral projections, a short opercular spine, and a broad interorbital space. Head is relatively long and moderately broad, and snout is relatively long. Rostral projection is triangular shaped and bears small spines along margin. Preopercular spine has supplemental spine. Opercular spine falls short of tip of cleithral spine. Gill rakers on first arch number 0 to 3 on upper limb and 10 to 15 on lower limb. Measurements are expressed as percent of SL: head length 38%–50%, snout length 15%–23%, upper jaw length 10%–17%, orbit diameter 9%–14%, interorbital width 8%–13%, pectoral fin length 29%–59%, first free pectoral fin ray length 23%–31%, body depth 28%– 41%. Joined pectoral fin rays number 12. First dorsal fin has 10 or 11 spines, with first 3 serrated, and second dorsal fin has 11 rays, with first serrated at base. First 2 dorsal fin spines terminate as long filaments in mature males. Anal fin rays number 9 or 10. Body is covered with small ctenoid scales. Nape is naked but may have rows of small papillae. Breast and posterior half of belly are scaled, and anterior half of belly is naked. Color in life is brown on upper half of body and white to slightly dusky

on lower half, with two or three yellow stripes along upper sides. First dorsal fin is light, with yellow spots and yellow pigment on filaments of first two dorsal spines, and black spot between fourth and fifth spines. Uppermost pectoral fin rays are marked with black and white bands. This species occurs between 20 and 216 m in the western Atlantic from North Carolina to Colombia, including the Gulf of Mexico. In the Gulf of Mexico it is widespread along the outer continental shelf. Food consists of copepods, mysid shrimps, penaeid shrimps, and ray-finned fishes, with juveniles consuming a higher proportion of copepods and mysids, and adults consuming a higher proportion of penaeid shrimps and ray-finned fishes. Maximum known size is 160 mm SL. references: Longley and Hildebrand 1941; Ginsburg 1950; Hildebrand 1954; Miller 1965; Chittenden and McEachran 1976; Lewis and Yerger 1976; Chittenden and Moore 1977; Hoese and Moore 1977, 1998; Ross 1977, 1983; Robins and Ray 1986; Miller and Richards 1991a; Boschung 1992; Huidobro-Campos and Schmitter-Soto 1993; Schaldach et al. 1997; Smith 1997; Richards and Miller 2002a.

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58 SCORPAENIFORMES

Prionotus alatus Spiny searobin

Goode and Bean, 1883

Prionotus alatus has nasal spines and greatly elongated pectoral fins. Snout is relatively long, concave in profile, and depressed anteriorly. Rostral projection is well developed and strongly rounded, with crenate margin. Mouth is subterminal and rather short. Supplemental preopercular spine is moderately developed but decreases in size with growth. Rostral and infraorbital spines are persistent throughout life. Villiform teeth are arranged in bands in jaws, vomer, and palatine. Gill rakers on first arch number 1 to 4 on upper limb and 9 to 21 on lower limb. Measurements are expressed as percent of SL: head length 34%–37%, snout length 16%–18%, orbital diameter 7%– 8%, interorbital width 3%– 4%, longest upper pectoral fin ray length 38%– 44%, longest lower pectoral fin ray length 56%– 63%, first free pectoral fin ray length 21%–23%, preopercular spine length 12%–16%. Joined pectoral fin rays number 12 or 13. Lower rays of joined pectoral fin are greatly elongated, and distal margin is oblique and concave. First dorsal fin has 10 (rarely 8 or 9) spines, and second dorsal fin has 12 (rarely 11) rays. Anal fin has 11 fin rays. Body is covered with small ctenoid scales. Nape is scaled, but belly is naked anterior to line connecting innermost pelvic rays.

Color is brown on dorsal and lateral aspects of body and light tan to white ventrally. Dark blotch is located on first dorsal fin. Pectoral fin is light, with medial and distal bands across all but outermost fin rays. This species occurs between 55 and 457 m in the western Atlantic from Virginia to the eastern Gulf of Mexico, from the west coast of Florida to Mobile Bay, and on Campeche Bank. In the eastern Gulf of Mexico it is common on carbonate sediments, including broken shell and bryozoan fragments. It apparently hybridizes with P. paralatus between Pensacola and Mobile Bay. Food consists of polychaetes, ostracods, stomatopods, penaeid shrimps, brachyuran crabs, and ray-finned fishes such as Bregmaceros atlanticus. Spawning occurs between March and June in the eastern Gulf of Mexico. Maximum known size is about 200 mm SL.

references: Longley and Hildebrand 1941; Ginsburg 1950; Teague 1951; Miller and Kent 1971; Lewis and Yerger 1976; Hoese and Moore 1977, 1998; Ross 1977, 1983; Robins and Ray 1986; McClure and McEachran 1992; Boschung 1992; Miller and Richards 1991b; Smith 1997; Richards and Miller 2002a.

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TRIGLIDAE 59

Prionotus longispinosus Bigeye searobin

Teague, 1951

Prionotus longispinosus has a large head and moderately long and acutely rounded pectoral fins. Snout is moderately long and depressed at tip. Rostral projection is well developed and broadly rounded, with a crenate margin. Mouth is subterminal and rather large. Nasal spine is absent. Rostral and anterior infraorbital spines are well developed, and posterior infraorbital and supplemental preopercular spines are moderately developed. Villiform teeth are arranged in bands in jaws, palatine, and vomer. Gill rakers on first arch number 0 or 1 on upper limb and 8 to 11 on lower limb. Measurements are expressed as percent of SL: head length 33%–36%, head depth 19%– 23%, snout length 14%–15%, orbit diameter 7%– 8%, interorbital width 4%–5%, upper jaw length 13%–15%, preopercular spine length 14%–16%, pectoral fin length 44%– 48%, first free pectoral fin ray length 23%– 27%, body depth 20%–23%. Joined pectoral fin rays number 12 or 13. First dorsal fin has 9 or 10 spines, and second dorsal fin has 12 or 13 rays. Anal fin rays number 10 to 12. Body is covered with small ctenoid scales. Nape, belly, and chest are fully scaled. Lateral scale rows number 88 to 106. Color is brownish gray dorsally, grading to white ventrally. Dorsal fins are light gray, with dark ocellated spot on first dorsal fin. Pectoral fin is nearly black,

with white, yellow, or light blue spots. Anal fin is light gray, with dark median band. Caudal fin is light gray, with dusky margin. This species occurs from near shore to 219 m in the western Atlantic from the northern and western Gulf of Mexico. It is most abundant between 9 and 91 m. In the Gulf of Mexico it is most common west of Apalachicola Bay on muddy bottoms. Young are common inshore and move into deeper water with growth. Spawning occurs from January through August or September but is most intense from January through April or May. Food consists of polychaetes, amphipods, mysid shrimps, penaeid shrimps, crabs, and rayfinned fishes. Penaeid shrimps and crabs appear to be the dominant prey, especially for large individuals. Small individuals consume a high proportion of crab megalops. Maturity is reached in the first year of life at 120 to 130 mm TL. Maximum known size is 350 mm SL.

Ginsburg 1950 (as P. rubio); Teague 1951; Hoese and Moore 1977 (as P. rubio), 1998; Ross 1983 (as P. rubio); Robins and Ray 1986; Miller and Richards 1991b; Boschung 1992; Hoff 1992; Russell et al. 1992; Schaldach et al. 1997 (as P. rubio); Smith 1997; Castro-Aguirre et al. 1999; Richards and Miller 2002a.

REFERENCES:

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60 SCORPAENIFORMES

Prionotus martis Ginsburg, 1950 Barred searobin

Prionotus martis has a relatively small, conical head and moderately short, truncate pectoral fins. Snout is moderately long and depressed at tip. Rostral projection is poorly developed, with rounded, crenate margin. Mouth is subterminal and relatively short. Rostral and infraorbital spines are moderately developed in small specimens but disappear with growth. Nasal and supplemental preopercular spines are poorly developed. Villiform teeth are arranged in bands in jaws, vomer, and palatine. Gill rakers on first arch number 1 on upper limb and 8 to 11 on lower limb. Measurements are expressed as percent of SL: head length 29%–32%, head depth 17%–20%, snout length 14%–15%, upper jaw length 9%– 10%, orbit diameter 5%– 6%, interorbital width 3%– 4%, preopercular spine length 8%– 10%, pectoral fin length 36%– 43%, first free pectoral fin ray length 18%–21%, body depth 17%–22%. Joined pectoral fin rays number 14 or 15. First dorsal fin has 10 spines, and second dorsal fin has 12 or 13 rays. Anal fin rays number 11 to 13. Body is covered with small ctenoid scales. Nape is scaled, and chest is fully scaled in specimens longer than 25 mm SL. Body is mottled with brownish gray dorsally

and laterally, and lower part of body is light tan. Three or four dark, broad diagonal bands or saddles often occur on back and caudal peduncle. Branchiostegal rays are dusky. Dorsal fins have dark, narrow horizontal bands. First dorsal fin has dark spot with pale margin on spines and often has narrow dark spot near margin between first and second dorsal fin. Pectoral fin is dusky or dark, with light mottling to nearly black. This species occurs between 11 and 46 m in the western Atlantic from the Gulf of Mexico. It occurs throughout the Gulf of Mexico within its depth range. Food consists of polychaetes, cumaceans, mysids, amphipods, various shrimps, and amphioxi. Spawning takes place from early spring to summer in the eastern Gulf of Mexico. Maximum known size is about 170 mm SL.

references: Ginsburg 1950; Lewis and Yerger 1976; Hoese and Moore 1977, 1998 (as P. carolinus); Ross 1977, 1983; Robins and Ray 1986; Boschung 1992; Huidobro-Campos and SchmitterSoto 1993 (as P. carolinus); Smith 1997; VegaCendejas et al. 1997 (as P. carolinus); CastroAguirre et al. 1999; Richards and Miller 2002a.

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TRIGLIDAE 61

Prionotus murielae

Mowbray, 1928

Prionotus murielae has nasal and supraocular cirri and filamentous upper pectoral fin rays. Head bears long, tapering cirrus behind anterior naris, and large, fimbriated cirrus on upper section of eye. Snout is of moderate length, depressed anteriorly, and concave in profile. Rostral projection is broadly rounded, with crenate margin. Supplemental preopercular spine is absent. Villiform teeth are arranged in rows in jaws, palatine, and vomer. First gill arch has three or four rudimentary gill rakers on upper limb and seven gill rakers on lower limb. Measurements are expressed as percent of SL: head length about 40%, orbit diameter about 11%, interorbital width about 6%, upper jaw length about 17%, pectoral fin length about 54%. Joined pectoral fin rays number 14, and uppermost two rays extend as fila-

ments to caudal fin base. First dorsal fin has 10 spines, and second dorsal fin has 12 fin rays. Anal fin rays number 11. Body is covered with small ctenoid scales. Nape and belly are scaled, and chest is scaled posterior to axil of pectoral fin. Lateral scale rows number 97. Pectoral fin has three transverse bands of pigment. This species occurs in the western Atlantic from Cay Sal Bank in the Bahamas. It is known from only a single specimen and thus has not been captured from the Gulf of Mexico but may occur there. Maximum known size is 200 mm SL.

references: Mowbray 1928, in Borodin 1928; Miller and Richards 1991b; Richards and Miller 2002a.

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62 SCORPAENIFORMES

Prionotus ophryas Jordan and Swain, 1885 Bandtail searobin

Prionotus ophryas has nasal and supraocular cirri and long, fan-shaped pectoral fins. Head bears long, tapering cirrus behind anterior naris, and large, fimbriated cirrus on upper section of eye, followed by several short, simple cirri. Snout is of moderate length, depressed anteriorly, and concave in profile. Rostral projection is broadly rounded, with crenate margin. Nasal, rostral, infraorbital, and supplemental preopercular spines are absent in smallest specimens. Villiform teeth are arranged in rows in jaws, palatine, and vomer. First gill arch has several rudimentary gill rakers on upper limb and six or seven gill rakers and five to eight rudiments on lower limb. Measurements are expressed as percent of SL: head length 32%–35%, head depth 18%– 21%, snout length 16%–17%, orbit diameter 6%– 8%, interorbital width 4%–5%, upper jaw length 13%–15%, preopercular spine length 13%–14%, pectoral fin length 60%– 70%, first free pectoral fin length 22%–25%, body depth 20%–22%. Joined pectoral fin rays number 14. First dorsal fin has 9 or 10 spines, and second dorsal fin has 11 to 13 rays. Anal fin rays number 10 or 11. Posterior margin of pectoral fin is concave between fourth and ninth rays. Body is covered with small ctenoid scales. Nape and belly are scaled, and chest is scaled posterior to axil of pectoral fin. Lateral

scale rows number 93 to 105. Color is grayish brown on dorsal surface and sides, and lower sides and ventral surface are light except for elongate extensions of dorsal coloration and dusky blotches. First dorsal fin has proximal and distal series of spots forming irregular band, and second dorsal fin has dark blotch on anterior part of fin and faint spots on remainder. Pectoral fin is dark, with darker spots between upper rays. Caudal fin has three broad, dark transverse bands. This species occurs between 7 and 110 m in the western Atlantic from Cape Hatteras to Venezuela, including the entire Gulf of Mexico. It is most abundant between 18 and 64 m. This species is associated with soft bottoms. Food consists of mysids, shrimps, and crabs. Spawning takes place between January and June. Maturity is reached at 100 to 129 mm TL at age one. Maximum known size is about 200 mm SL.

references: Longley and Hildebrand 1941; Ginsburg 1950; Teague 1951; Hoese and Moore 1977, 1998; Ogren and Brusher 1977; Ross 1983; Robins and Ray 1986; Cervigón 1991, 1993b; Boschung 1992; Hoff 1992; Russell et al. 1992; HuidobroCampos and Schmitter-Soto 1993; Schaldach et al. 1997; Smith 1997; Richards and Miller 2002a.

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TRIGLIDAE 63

Prionotus paralatus Mexican searobin

Ginsburg, 1950

Prionotus paralatus has a long preopercular spine and rather short, emarginate pectoral fins. Snout is moderately long, depressed anteriorly, and concave in profile. Mouth is subterminal and relatively small. Rostral projection is broadly rounded and has crenate margin. Rostral, infraorbital, and supplemental preopercular spines are well developed, and nasal spine is absent. Villiform teeth are arranged in bands in jaws, vomer, and palatine. First gill arch has 1 to 3 tubercles on upper limb and 11 to 16 tubercles and gill rakers on lower limb. Measurements are expressed as percent of SL: head length 35%– 41%, head depth 19%–23%, snout length 16%–18%, orbit diameter 7%–9%, interorbital width 3%– 4%, upper jaw length 11%–13%, preopercular spine length 20%–27%, longest joined upper pectoral fin ray length 32%– 40%, longest joined lower pectoral fin ray 38%– 44%, uppermost free pectoral fin ray length 24%– 29%, body depth 17%–20%. Joined pectoral fin rays number 12 or 13. First dorsal fin has 9 or 10 spines, and second dorsal fin has 11 to 13 rays. Anal fin rays number 10 or 11. Body is covered with small ctenoid scales except on chest anterior to outermost pelvic fin rays.

Color is light grayish brown on dorsal and upper lateral surfaces, and light tan to whitish on lower lateral and ventral surfaces. Pectoral fin is uniformly black or has black and white crossbands on upper part of fin. First dorsal fin has dark oval-shaped spot between fourth and fifth spines. This species occurs between 9 and 274 m in the western Atlantic in the Gulf of Mexico from Panama City, FL, to the Gulf of Campeche. It is most abundant in the northern Gulf of Mexico between 60 and 120 m. Maturity is reached at 100 to 120 mm TL at age one. Specimens between Gulfport, MS, and Panama City, FL, are apparently hybrids of P. paralatus and P. alatus. Food consists of polychaetes, bivalve mollusks, shrimps, and crabs. Maximum known size is about 180 mm SL.

references: Ginsburg 1950; Miller and Kent 1971; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Hoff 1992; McClure and McEachran 1992; Russell et al. 1992; Huidobro-Campos and Schmitter-Soto 1993; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; Richards and Miller 2002a.

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64 SCORPAENIFORMES

Prionotus punctatus

(Bloch, 1793)

Prionotus punctatus has a large, deep head and moderately long and rounded pectoral fins. Snout is relatively long, straight in profile, and depressed anteriorly. Rostral projection is broadly rounded, with crenate margin. Nasal spine is absent, but supplemental preopercular, rostral, and infraorbital spines are persistent through life. Mouth is long and subterminal. Villiform teeth are arranged in narrow rows in jaws, palatine, and vomer. Gill rakers on first arch number 8 to 11. Measurements are expressed as percent of SL: head length 35%– 40%, snout length 17%–18%, orbit diameter 6%–11%, interorbital width 4%– 6%, upper jaw length 15%–18%, body depth 22%–25%. Joined pectoral fin rays number 13. First dorsal fin has 10 spines, and second dorsal fin has 11 to 13 rays. Anal fin rays number 11. Body is covered with small ctenoid scales. Nape and chest are fully scaled. Lateral scale rows range from 97 to 110. Color is brown to grayish brown, with a few to many dark spots irregularly distributed on upper half

of body and white to yellowish white or light brown on lower half of body. First dorsal fin of small specimens has a dark spot between fourth and fifth spines, but the spot is indistinct in large specimens. Pectoral fin is dark to nearly black, with upper section occasionally marked with darker spots or irregular crossbands. This species occurs between 7 and 115 m on soft, sandy, or mud bottoms in the western Atlantic from Cuba and the southern Gulf of Mexico to Argentina. In the Gulf of Mexico it is known from Tuxpan, Veracruz; Laguna de Términos, Campeche; and Celestún, Yucatán. Food consists of shrimps, decapod crustaceans, and gobiid fishes. Maximum known size is about 200 mm SL. references: Ginsburg 1950; Cervigón 1991, 1993b; Russell et al. 1992; Huidobro-Campos and Schmitter-Soto 1993; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999; Richards and Miller 2002a.

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TRIGLIDAE 65

Prionotus roseus Jordan and Evermann, 1887 Bluespotted searobin

Prionotus roseus has a moderately large head and very long pectoral fins covered with blue spots. Snout is moderately long, depressed anteriorly, and slightly concave in profile. Rostral projection is poorly developed and broadly rounded, with crenate margin. Mouth is subterminal and relatively small. Nasal, rostral, infraorbital, and supplemental preopercular spines are absent except in very small specimens. Villiform teeth are arranged in bands in jaws, palatine, and vomer. Gill rakers on first arch number 7 to 10, with none on upper limb and 1 in corner. Measurements are expressed as percent of SL: head length 28%–33%, head depth 16%–21%, snout length 13%–15%, orbit diameter 5%– 6%, interorbital width 3%– 4%, upper jaw length 10%–11%, preopercular spine length 6%–10%, pectoral fin length 58%–74%, first free pectoral ray length 19%– 22%, body depth 14%–19%. Joined pectoral fin rays number 12 to 14. First dorsal fin has 10 spines, and second dorsal fin has 12 rays. Anal fin rays number 10 to 12. Body is covered with small ctenoid scales, including belly, chest, and throat. Lateral scale rows number 89 to 104. Color is light brown to grayish brown on dorsal surface and upper sides, and whitish to tan on lower sides and ventral surface. Dorsal

fin is dark, with black spot between fourth and fifth spines. Pectoral fin is dark, with bright blue oval spots symmetrically arranged along rays. Anal fin is unpigmented or peppered with black spots distally. This species occurs from 9 to 183 m on soft bottoms in the western Atlantic from North Carolina to Brazil, including the entire Gulf of Mexico and the Greater and Lesser Antilles. It is most abundant between 27 and 91 m. Food consists of polychaetes, mysids, gammaridean amphipods, shrimps, crabs, and rayfinned fishes, with shrimps composing the majority of the organisms. Gonadal development begins in the late fall, and spawning takes place from April to September. Maturity is reached at about 90 mm SL, and maximum known size is about 200 mm SL.

references: Longley and Hildebrand 1941 (as P. microlepis); Ginsburg 1950; Lewis and Yerger 1976; Hoese and Moore 1977, 1998; Ross 1983; Uyeno and Sato 1983c; Robins and Ray 1986; Cervigón 1991, 1993a; Boschung 1992; Russell et al. 1992; Huidobro-Campos and Schmitter-Soto 1993; Schaldach et al. 1997; Smith 1997; CastroAguirre et al. 1999; Richards and Miller 2002a.

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66 SCORPAENIFORMES

Prionotus rubio Jordan, 1886 Blackwing searobin

Prionotus rubio has very long, obliquely truncate pectoral fins. Snout is relatively long, slightly concave in profile, and depressed anteriorly. Rostral projection is relatively broad and rounded, with crenate margin. Mouth is subterminal and relatively long. Nasal spine and supplemental preopercular spine are absent. Rostral spine and first infraorbital spine are present at all sizes. Villiform teeth are arranged in bands in jaws, vomer, and palatine. Gill rakers (including tubercles) number 3 or 4 on upper limb and 10 to 14 on lower limb. Measurements are expressed as percent of SL: head length 31%–36%, head depth 17%– 21%, snout length 14%–16%, orbit diameter 6%–7%, interorbital width 4%–5%, upper jaw length 13%–15%, preopercular spine length 7%–9%, pectoral fin length 54%– 63%, first free pectoral fin ray length 20%–26%, body depth 16%–21%. Pectoral fin has 12 or 13 joined rays, with ninth longest. First dorsal fin has 10 spines, and second dorsal fin has 12 or 13 rays. Anal fin rays number 11 or 12. Body is covered with small ctenoid scales. Nape is scaled, and chest is completely scaled to isthmus. Color is light brown to grayish brown on

upper body and side, and light tan to white ventrally. First dorsal fin has black spot that fades with age. Second dorsal fin is lightly pigmented. Pectoral fin is dark, with indistinct light or dark spots on upper section and blue on lower margin. Anal fin is unpigmented. This species occurs between the shoreline and 212 m in the western North Atlantic from North Carolina to Cuba, including the entire Gulf of Mexico. It is most common between 9 and 55 m. Food consists mostly of mysids, stomatopods, shrimps, crabs, ray-finned fishes, and to a lesser extent polychaetes. Maturity is reached between 80 and 120 mm TL at age one. Maximum known size is about 230 mm SL.

references: Ginsburg 1950 (as P. pectoralis and P. salmonicolor); Teague 1951 (as P. vanderbilti); Hoese and Moore 1977 (as P. salmonicolor), 1998; Ross 1977, 1983 (as P. salmonicolor); Robins and Ray 1986; Cervigón 1991; Miller and Richards 1991b; Boschung 1992; Russell et al. 1992; Huidobro-Campos and Schmitter-Soto 1993; Schaldach et al. 1997 (as P. salmonicolor); Smith 1997; Castro-Aguirre et al. 1999; Richards and Miller 2002a.

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TRIGLIDAE 67

Prionotus scitulus Leopard searobin

Jordan and Gilbert, 1882

Prionotus scitulus has a conical head and moderately long and strongly rounded pectoral fins. Snout is relatively long, slightly concave in profile, and depressed anteriorly. Rostral projection is broadly rounded, with crenate margin. Mouth is subterminal and short. Nasal and supplemental preopercular spines are slightly developed in small specimens but absent in large juveniles and adult specimens. Rostral and infraorbital spines are poorly developed in small specimens and are absent in large juveniles and adults. Villiform teeth are arranged in narrow rows in jaws, palatine, and vomer. Gill rakers on first arch number 9 to 13. Measurements are expressed as percent of SL: head length 27%–30%, head depth 13%–16%, snout length 13%–15%, orbit diameter 4%– 5%, interorbital width 2%–3%, upper jaw length 9%–11%, preopercular spine length 7%–9%, pectoral fin length 28%–37%, first free pectoral fin ray length 15%–19%, body depth 13%–15%. Joined pectoral fin rays number 12 to 14. First dorsal fin has 10 spines, and second dorsal fin has 12 to 14 rays. Anal fin rays number 11 to 13. Body is covered with small ctenoid scales. Nape is scaled, but chest is naked anterior to base of lower free pectoral

fin rays. Lateral scale rows number 103 to 130. Color is brown dorsally, with reddish brown spots, and white to light tan ventrally. First dorsal fin has two dark spots, one between first and second spines, and second between fourth and fifth spines. Pectoral fin is dark, with brightly colored spots. This species occurs from shallow bays to 91 m in the western North Atlantic from Virginia to Venezuela, including the entire Gulf of Mexico. It is most common from the shoreline to 46 m and is more abundant in the eastern than in the northwestern Gulf of Mexico. Food consists mostly of polychaetes, shrimps, crabs, and amphioxus, with amphipods and mysids the most abundant items. Spawning takes place between September and April or May. Maturity is reached at about 110 mm SL, and maximum known size is about 250 mm SL.

references: Ginsburg 1950; Teague 1951; Hoese and Moore 1977, 1998; Ross 1977, 1978, 1983; Robins and Ray 1986; Boschung 1992; Hoff 1992; Russell et al. 1992; Huidobro-Campos and Schmitter-Soto 1993; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999; Richards and Miller 2002a.

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68 SCORPAENIFORMES

Prionotus stearnsi Jordan and Swain, 1885 Shortwing searobin

Prionotus stearnsi has a rather large and compressed head; a cluster of small papillae on eyeball; and small, rounded pectoral fins. Snout is relatively long and moderately depressed anteriorly. Rostral projection is poorly developed and has crenate margin. Mouth is terminal and relatively large, with small bony knob at symphysis extending beyond upper jaw. Nasal, supplemental preopercular, rostral, and infraorbital spines are absent. Villiform teeth are arranged in narrow bands in jaws, palatine, and vomer. Gill rakers on first arch number 9 to 11. Measurements are expressed as percent of SL: head length 31%–36%, head depth 16%–20%, snout length 13%–16%, orbit diameter 5%–7%, interorbital width 3%– 4%, upper jaw length 15%–17%, preopercular spine length 5%– 6%, pectoral fin length 19%–27%, first free pectoral fin ray length 15%–19%, body depth 17%–21%. Joined pectoral fin rays number 12 or 13. First dorsal fin has 10 spines, and second dorsal fin has 12 or 13 rays. Anal fin rays number 10 to 12. Body is covered with small ctenoid scales.

Chest is completely scaled. Color is silvery black. Pectoral fin and posterior section of caudal fin are darkly pigmented. This species occurs between 11 and 549 m in the western Atlantic from North Carolina to French Guiana, including the entire Gulf of Mexico. It is most common between 37 and 110 m. This species is associated with soft to semihard bottoms. Food consists largely of shrimps, crabs, and ray-finned fishes, with larger specimens consuming a larger percentage of fishes. Spawning takes place from November or December through May or June. It matures between 80 and 115 mm TL. Maximum known size is about 180 mm SL.

references: Longley and Hildebrand 1941; Ginsburg 1950; Teague 1951; Lewis and Yerger 1976; Hoese and Moore 1977, 1998; Ross 1983; Uyeno and Sato 1983c; Robins and Ray 1986; Cervigón 1991, 1993a; Boschung 1992; Hoff 1992; Russell et al. 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Richards and Miller 2002a.

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TRIGLIDAE 69

Prionotus tribulus Bighead searobin

Cuvier, 1829

Prionotus tribulus has a large, broad head and moderately large and rounded pectoral fins. Snout is relatively long, nearly straight in profile, and depressed anteriorly. Rostral projection is broadly rounded, with crenate margin. Mouth is subterminal and relatively large. Nasal spine is absent, but preopercular spine is well developed. Rostral and infraorbital spines are prominent in small specimens but are rudimentary in large specimens. Villiform teeth are arranged in narrow bands in jaws, palatine, and vomer. Gill rakers on first arch number 8 to 16. Measurements are expressed as percent of SL: head length 39%– 43%, head depth 21%–23%, snout length 17%–19%, orbit diameter 7%–9%, interorbital width 7%– 8%, upper jaw length 19%–20%, preopercular spine length 10%–13%, pectoral fin length 39%– 47%, first free pectoral fin ray length 21%–28%, body depth 20%–23%. Joined pectoral fin rays number 12 to 14. First dorsal fin has 10 spines, and second dorsal fin has 11 or 12 rays. Anal fin has 10 to 12 rays. Body is covered with moderate-sized ctenoid scales. Nape is scaled, and chest is fully scaled. Lateral scale rows number 69 to 84. Color is brown to grayish brown, with three short oblique dark

bands on upper half of body, and white to yellowish white on lower half of body. Dorsal fin has dark blotch near margin between fourth and fifth spines. Pectoral fin is dusky to almost black, often with many irregular cross streaks. Free pectoral fin rays are banded. This species occurs between the shoreline and 183 m in the western Atlantic from New York to Florida and throughout the Gulf of Mexico. It is most common between 9 and 64 m. Juveniles enter estuaries. Food consists of mysids, amphipods, shrimps, crabs, ray-finned fishes, and to a lesser extent polychaetes. Spawning takes place from fall to early spring. Maturity is reached at 85 to 140 mm TL at age one. Maximum known size is about 350 mm SL.

references: Ginsburg 1950; Teague 1951; Springer and Woodburn 1960; Hoese and Moore 1977, 1998; Castro-Aguirre and Márquez-Espinoza 1981; Ross 1983; Robins and Ray 1986; Boschung 1992; Hoff 1992; Russell et al. 1992; HuidobroCampos and Schmitter-Soto 1993; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999; Richards and Miller 2002a.

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70

PERISTEDIIDAE

Armored searobins

Armored searobins are slender to moderately robust, with body covered on each side with four rows of spinous scutes and two lowermost rays of pectoral fins thickened and free of membranes. Head is large and bony, with many ridges and spines. Snout is broad and depressed. Lachrymal bone is moderately projected as flattened rostrum that bears short to very long rostral projections on each side. Eye is moderate to rather small in size and laterally located. Bony ridge (bony stay), formed by infraorbital bones, is generally present below eye and extends from just behind mouth to preoperculum. Mouth is short and slightly inferior or subterminal and does not completely close. Premaxilla is protractile, and maxilla slips under preorbital bone. Teeth are absent in jaws, palatine, and vomer. Tongue is usually absent, but when present, it is rudimentary and bifurcated at tip. Mandibular barbel is present, and lip and chin barbels are usually present. Preoperculum and operculum have well-developed spines. Gill membrane is free of isthmus or narrowly joined to isthmus anteriorly. Gills are present on four arches, with large slit behind fourth arch. Pectoral fin is short and rather narrow, with 11 to 16 attached rays in addition to swollen free rays. First dorsal fin has seven to nine spines, and second dorsal fin has 16 to 23 rays. Pelvic fin is thoracic and has one spine and 5 rays. Anal fin is located below second dorsal fin and has 16 to 23 rays. Caudal peduncle is very slender, and caudal fin is narrow and truncate. Physoclistous gas bladder and pyloric caecae are present. Gas bladder has a single lobe. Armored searobins occur worldwide in tropical to temperate seas. All species are benthic on continental and insular slopes. They usually occur at greater than 180 m. There are about 30 species in four genera, 12 species in a single genus occur in the western Central Atlantic, and 8 to 12 species in a single genus occur in the Gulf of Mexico. references: Washington, Eschmeyer, et al. 1984; Washington, Moser, et al. 1984; Heemstra 1986b; Hureau and Litvinenko 1986b; Eschmeyer 1994; Nelson 1994; Richards 1996; Smith 1997: Richards and Miller 2002b.

Key to the Species of the Gulf of Mexico (Adapted from Richards and Miller 2002b) 1a. Head narrow, perifacial rim barely extending beyond posterior lateral margin of head (Fig. 3). . . . . . . . . . . . . . . . . 2

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PERISTEDIIDAE 71

FIG 3

1b. Head moderately broad to broad, perifacial rim distinctly extending beyond posterior lateral margin of head (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2a. Interorbital width equal to or greater than minimum rostrum width; body depth and width at level of anus about equal to depth and width at head; body rectangular in cross section . . . . . . . . . . . . . . . . . Peristedion gracile p. 76 2b. Interorbital width less than minimum rostrum width; body depth and width at level of anus considerably less than depth and width at head; body not rectangular in cross section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. First (upper) free pectoral fin ray longer than joined section of fin; 17 to 23 (usually 21) chin barbels (on each side) . . . . . . . . . . . . . . . . . . . . . . . . . . Peristedion imberbe p. 78 3b. First (upper) free pectoral fin ray shorter than joined section of fin; 34 to 57 (usually 41 to 43) chin barbels (on each side) . . . . . . . . . . . . . . . . . . . . . . Peristedion sp. p. 84 4a. Perifacial rim posteriorly terminating perpendicular to head (occasionally as flat spinelike projection in P. miniatum; Fig. 5); anterior edge of first ventral scute posterior to anterior edge of bony ridge of pelvic girdle . . . . . . . . . . . . . . . 5 4b. Perifacial rim posteriorly terminating as spine or flat spinelike projection (Fig. 6); anterior edge of first ventral scute equal with or anterior to anterior edge of bony ridge of pelvic girdle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 5a. Dorsal preopercular ridge and second infraorbital spine very large; 8 or 9 chin barbels (on each side), with anterior cluster with 1 large and 2 small barbels, and other clusters with 1 large and 1 small barbel, but last cluster may have only 1 large barbel . . . . . . . . . . Peristedion miniatum p. 80 5b. Dorsal preopercular ridge and second infraorbital spine not large; usually either more or fewer than 8 or 9 chin barbels (on each side) and not arranged into clusters of large and small barbels (P. unicuspis has 8 or 9 chin barbels, but they are arranged in four clusters of 2 equal-length barbels, and the ninth, if present, occurs separately). . . . . . . . . . . . . . . 6

FIG 4

FIG 5

FIG 6

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72 SCORPAENIFORMES

FIG 7

FIG 8

6a. Rostral projection very short, broad, and triangular shaped (Fig. 7) . . . . . . . . . . . . . . . . . Peristedion ecuadorense p. 75 6b. Rostral projection not short and not triangular shaped (Fig. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7a. Rostral projection very slender and longer than snout; 1 lip barbel (on each side); 0 to 1 (usually 0) chin barbel (on each side) . . . . . . . . . . . . . . . . Peristedion antillarum p. 73 7b. Rostral projection slender or not, but shorter than rostrum; more than 1 lip barbel (on each side); more than 1 chin barbel. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8a. Head very broad; lip barbels (on each side) in two clusters of 1 barbel each. . . . . . . . . . . Peristedion longispatha p. 77 8b. Head not very broad; lip barbels (on each side) in two or three clusters, and two clusters have more than 1 barbel .............................................9 9a. Lip barbels (on each side) in three clusters of 2, 1, and 2 barbels; mandibular filamentous barbel barely reaching posterior articulation of jaws; rostral projection short and broad . . . . . . . . . . . . . . . . . . . Peristedion truncatum p. 82 9b. Lip barbels (on each side) in two clusters of 2 barbels each; mandibular filamentous barbel at least reaching posterior end of perifacial rim; rostral projection not short and broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10a. Mandibular filamentous barbel reaching considerably beyond perifacial rim to level of first ventral scute; posterior scutes of supramedian series unicuspid; peritoneum and posterior roof of mouth dark . . . . . . . . . . . . . . . . . . . . . . . . . Peristedion unicuspis p. 83 10b. Mandibular filamentous barbel not extending beyond posterior end of perifacial rim; posterior scutes of supramedian series bicuspid; peritoneum and posterior roof of mouth silvery, white, or translucent . . . . . . . . . . . . . . . . . . . . . . . . . . . Peristedion greyae p. 77 11a. Perifacial rim terminating as strong spine; nasal spine large; more than 30 chin barbels (on each side) . . . . . . . . . . . . . . . . . . . . . . . . Peristedion brevirostre p. 74 11b. Perifacial rim terminating as flat spinelike projection or small spine; nasal spine small or absent; fewer than 24 chin barbels (on each side) . . . . . . . Peristedion thompsoni p. 81

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PERISTEDIIDAE 73

Peristedion antillarum

Teague, 1961

Peristedion antillarum has extremely long and narrow rostral projections and a long mandibular barbel. Head is short, moderately narrow, and depressed, and snout is very long and narrow. Rostral projection is very slightly laterally oriented, with rostral spine very small or absent. Nasal spine is absent. Perifacial rim is not interrupted by large spines or crenulations, and terminates as a moderately acute and flat spine, with posterior margin perpendicular to head. Lip has a single barbel. Chin has zero to one barbel in addition to one long to very long filamentous mandibular barbel. Gill rakers on first arch number 25 to 29. Measurements are expressed as percent of SL: head length 34%– 37%, head depth 16%–17%, rostral projection length 25%–32%, snout length 18%– 19%, mandibular barbel length 16%–22%, orbit length 7%–9%, interorbital width 7%– 8%, joined pectoral fin length 22%–23%, first free pectoral fin ray length 24%–29%, body depth 17%–20%. Joined pectoral fin rays number 12 or 13. First dorsal fin has seven or

eight spines, and second dorsal fin has 16 or 17 rays. Anal fin has 15 to 18 rays. Scutes in dorsal series number 26 or 27, in supramedian series number 21 or 22, in inframedian series number 21 or 22, and in ventral series number 20, with anterior pointed end of first ventral scute posterior to anterior edge of bony ridge of pelvic girdle. Color in preservative is bronze yellow, with black margins on two dorsal fins and anal fin. This species occurs between 329 and 649 m in the western Atlantic from the Antilles off the Florida Straits to Panama, including the eastern Gulf of Mexico, the Florida Keys, and the Greater and Lesser Antilles. It has been captured once in the Gulf of Mexico off southwest Florida at 2429N, 8327W. Juveniles occur between 350 and 400 m, and adults occur from 400 to 650 m. Maximum known size is 180 mm SL. references: Teague 1961; Richards and Miller 2002b.

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74 SCORPAENIFORMES

Peristedion brevirostre

(Günther, 1860)

Peristedion brevirostre has very short and broad rostral projections and numerous chin barbels. Head is moderately high and very broad, and snout is rather short and broad. Rostral projection is triangular shaped, directed anteriorly, and lacking spine on base. Lower jaw has a single serrated ridge. Lip barbels are finely pointed, number 4, and are arranged in two clusters of 2 barbels each. Chin barbels are flattened and sharp pointed, number 31 to 42, and are arranged in seven clusters. Filamentous mandibular barbel is distinctly longer than longest chin barbel. Perifacial rim terminates as rather narrow, acute angled spine. Gill rakers number 2 to 4 on upper limb and 10 to 13 on lower limb. Measurements are expressed as percent of SL: head length 34%– 35%, head depth 16%–19%, rostral projection length 7%–9%, snout length 14%–16%, orbit length 10%–11%, interorbital width 10%–11%, mandibular barbel length 12%– 16%, first free pectoral fin ray length 20%– 21%, joined pectoral fin length 25%–28%,

body depth 16%–19%. Joined pectoral fin rays number 12. Dorsal fin has seven spines and 17 rays. Anal fin has 17 rays. Scutes in dorsal series number 26, in supramedian series number 32, in inframedian series number 23, and in ventral series number 21, with anterior pointed end of first ventral scute on line with or anterior to anterior edge of bony knob of pelvic girdle. Color in preservative is yellowish, with mottled spotting on pectoral fin. This species occurs between 220 and 527 m in the western Atlantic from southern Florida and the Bahamas to Costa Rica, including the Antilles and possibly the eastern Gulf of Mexico. The Gulf of Mexico record is based on Longley and Hildebrand’s (1941) account from the Dry Tortugas. This species is associated with rocky bottoms. Maximum known size is 250 mm SL. references: Longley and Hildebrand 1941 (as P. platycephalum); Teague 1961 (in part as P. platycephalum); Richards and Miller 2002b.

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PERISTEDIIDAE 75

Peristedion ecuadorense

Teague, 1961

Peristedion ecuadorense has short and broad rostral projections and broad perifacial rims. Head is very short and moderately depressed, and snout is relatively long and broad. Rostral projection is broad and triangular shaped and bears stout spine near base. Nasal spine is absent. Lip barbels number 5 and are arranged in three clusters of 2, 1, and 2. Chin barbels number 11 to 13 and are arranged in four or five clusters. Perifacial rim ends as broad rightangled wing. Gill rakers number 3 or 4 on upper limb, 1 in corner, and 23 or 24 on lower limb. Measurements are expressed as percent of SL: head length 34%–36%, head depth 16%–17%, rostral projection length 3%– 5%, snout length 15%–16%, mandibular barbel length 16%–19%, orbit length 7%–9%, interorbital width 8%–9%, joined pectoral fin length 14%–15%, first free pectoral fin ray length 19%–21%, body depth 16%–18%. Pectoral fin has 13 or 14 joined rays. First dorsal fin has eight spines, and second dorsal fin has 16 to 19 rays. Anal fin has 17 to 19 rays.

Scutes in dorsal series number 27 or 28, in supramedian series number 30 to 32, in inframedian series number 27, and in ventral series number 21 or 22, with anterior point of first ventral scute posterior to anterior edge of bony knob of pelvic girdle. Color in preservative is pale yellow, with dusky stripes on body between scute rows, striping and spotting on head, distal half of first dorsal fin black, and distal quarter of second dorsal fin dusky. This species occurs between 324 and 910 m in the western Atlantic from Virginia to Brazil, including the northern Gulf of Mexico, the Florida Keys, and the Lesser Antilles. In the Gulf of Mexico it is known from Louisiana and is common off Florida. It is rarely captured in the Lesser Antilles. Maximum known size is about 200 mm SL.

references: Teague 1961; Miller 1967b; Uyeno and Sato 1983d; Darcy and Gutherz 1984; Richards and Miller 2002b.

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Peristedion gracile Goode and Bean, 1896

Peristedion gracile has long and narrow rostral projections and a broad interorbital space. Head is short, narrow, and depressed, and snout is very long. Rostral projection is slightly to moderately laterally oriented, and rostral and nasal spines are absent. Perifacial rim is very narrow and terminates as a narrow, posteriorly directed spine. Lip barbels number 5 and are arranged in two clusters of 2 and 3. Chin barbels number 26 to 36 and are arranged in seven clusters. Gill rakers number 2 or 3 on upper limb, 1 in corner, and 19 to 26 on lower limb. Measurements are expressed as percent of SL: head length 34%–36%, head depth 13%–15%, rostral projection length 14%–17%, snout length 17%–19%, mandibular barbel length 11%–17%, orbit length 7%– 8%, interorbital width 7%–9%, pectoral fin length 20%–25%, first free pectoral fin ray length 19%–22%, body depth 12%–16%. Joined pectoral fin rays number 12. First dorsal fin has eight spines, and second dorsal fin

has 20 rays. Anal fin has 20 to 21 rays. Scutes in dorsal series number 28 or 29, in supramedian series number 30 to 32, in inframedian series number 26, and in ventral series number 23. Bicuspid spines in supramedian series number six to nine. Color in preservative is bronze yellow. This species occurs between 29 and 475 m in the western Atlantic from Virginia to Suriname, including the Gulf of Mexico. It is most common on the outer continental shelf between 74 and 146 m. In the Gulf of Mexico it is common off Florida and Alabama and also occurs off Texas and in the Bay of Campeche. Maximum known size is about 200 mm SL.

references: Longley and Hildebrand 1941 (as P. imberbe); Fowler 1952 (as P. taeniopterum); Teague 1961 (in part as P. taeniopterum); Hoese and Moore 1977, 1998; Uyeno and Sato 1983d; Darcy and Gutherz 1984; Boschung 1992; Schaldach et al. 1997; Richards and Miller 2002b.

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PERISTEDIIDAE 77

Peristedion greyae

Miller, 1967

Peristedion greyae has long, narrow rostral exsertions and numerous lip and chin barbels. Head is moderately broad, and snout is moderately long. Rostral projection is narrow, is directed slightly laterally, and bears a single spine near base. Nasal spine is absent. Lower jaw has two or three serrated ridges. Lip barbels are slender and arranged in two groups of 2. Chin barbels are slender and number 12 or 13 and are grouped in five clusters of 3, 3, 3, 2, and 1 or 2. Filamentous mandibular barbel is moderately long but fails to reach perifacial rim, with filaments arranged in clusters of two on one side and one on the other side, and terminating as three or four filaments. Perifacial rim terminates as broad right-angled wing, often with concave lateral margin. Margin of perifacial rim bears strong spines and crenulations. Gill rakers on first arch number 5 to 7 on upper limb and 19 to 26 on lower limb. Measurements are expressed as percent of SL: head length 33%– 42%, head depth 14%–17%, rostral projection length 11%–12%, snout length 15%–16%, mandibular barbel length 16%–18%, orbit length 7%– 8%, interorbital width 6%–7%, first free pectoral fin ray length 18%–24%, first joined pectoral fin ray length 18%–21%, body depth 16%–21%. Joined pectoral fin rays number 11 to 13. First dorsal fin has eight spines, and second dorsal fin has

18 to 20 rays. Anal fin has 18 to 20 rays. Scutes in dorsal series number 25 to 27, in supramedian series number 32 to 35, in inframedian series number 23 to 26, and in ventral series number 22 to 24. Posterior scutes of supramedian series are bicuspid, and anterior pointed end of first ventral scute is posterior to anterior edge of bony knob of pelvic girdle. Vertebrae number 33 to 35. Color in life is pink dorsally and white to pink ventrally, with broad bronze band laterally. Fins are bordered with dark red band, and pectoral fin is marked with four pink to red bands. Color in preservative is pale except for dorsal fins and pectoral fin. Distal half of first dorsal fin is black, and margin is white. Margin of second dorsal fin is black, and distal half of pectoral fin is black and margin is white. This species occurs between 179 and 914 m in the western Atlantic from southern New England, off Georges Bank, to northern Brazil, including the Gulf of Mexico, the Florida Keys, and the Bahamas. It occurs throughout the Gulf of Mexico and is common off Florida. Maximum known size is 190 mm SL.

references: Longley and Hildebrand 1941 (as P. longispathum); Teague 1961 (as P. longispatha); Miller 1967b; Richards and Miller 2002b.

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Peristedion imberbe

(Poey, 1861)

Peristedion imberbe has long, narrow rostral projections and a narrow interorbital distance. Head is short, narrow, and relatively depressed, and snout is very long. Rostral projection is narrow, lacks spine at base, and is directed anteriorly. Nasal spine is lacking. Perifacial rim is very narrow and terminates as blunt or rounded protuberance. Lip barbels number 5 to 7 in two clusters of 2 to 4 barbels. Chin barbels number 17 to 23 and are arranged in six clusters of 2 to 5. Gill rakers number 4 or 5 on upper limb and 20 to 23 on lower limb. Measurements are expressed as percent of SL: head length 35%–38%, head depth 15%–17%, rostral projection length 14%–16%, snout length 18%–20%, mandibular barbel length 8%–11%, orbit length about 8%, interorbital width 6%–7%, joined pectoral fin length 19%–24%, first free pectoral fin ray length 25%–27%, body depth 16%–18%. Joined

pectoral fin rays number 11 or 12. First dorsal fin has eight spines, and second dorsal fin has 18 or 19 rays. Anal fin has 19 rays. Scutes in dorsal series number 28, in supramedian series number 30, in inframedian series number 24 or 25, and in ventral series number 22. Bicuspid spines in supramedian series number seven to nine. Color in preservative is pale yellow, with pale dusky markings on upper surface of body. Dorsal fins have submarginal dark stripe, and pectoral fin has dusky reticular pattern. This species occurs between 137 and 457 m in the western Atlantic from Honduras, Cuba, and the southern Lesser Antilles to Brazil. It has not been recorded from the Gulf of Mexico, but it may occur there. Maximum known size is 180 mm SL. references: Teague 1961; Richards and Miller 2002b.

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PERISTEDIIDAE 79

Peristedion longispatha Goode and Bean, 1886

Peristedion longispatha has moderately long rostral projections and broad perifacial rims. Head is broad and moderately depressed, and snout is of moderate length. Rostral projection is narrow and curves laterally. Rostral and nasal spines are present in juveniles but absent in adults. Lower jaw has a single serrated ridge. Lip barbels number two and are separated. Chin barbels number 12 or 13 and are arranged in six clusters of 1, 2, 2 or 3, 3, 2, and 2. Filamentous mandibular barbel is moderately long but fails to reach termination of perifacial rim. Perifacial rim terminates as right-angled wing, with straight to slightly undulating lateral margin free of strong spines or crenulations. Gill rakers number 4 on upper limb and 17 to 19 on lower limb. Measurements are expressed as percent of SL: head length 36%–38%, head depth 16%–19%, rostral projection length 14%–16%; snout length 15%–17%, orbit diameter 8%–9%, interorbital width 9%–10%, mandibular barbel length 11%–16%, joined pectoral fin length 18%–20%, first free pectoral fin ray length

17%–23%, body depth 17%–20%. Joined pectoral fin rays number 12. First dorsal fin has eight spines, and second dorsal fin has 17 rays. Anal fin rays number 17. Scutes in dorsal series number 24 to 26, in supramedian series number 26, in inframedian series number 24 or 25, and in ventral series number 21, with anterior pointed end of first ventral scute posterior to anterior edge of bony knob of pelvic girdle. Color in preservative is pale yellow, with margin of dorsal fin dusky. This species occurs between 101 and 787 m in the western Atlantic in the Gulf of Mexico, the Straits of Yucatán, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs off the Mississippi Delta; however, most of the specimens recorded from the Gulf of Mexico were later redetermined as P. greyae (Miller (1967b). Maximum known size is about 180 mm SL. references: Teague 1961 (as P. bartschi, P. greyae, P. imberbe, and P. longispathum); Miller 1967b; Richards and Miller 2002b.

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Peristedion miniatum Armored searobin

Goode, 1880

Peristedion miniatum has relatively short rostral projections and a long, filamentous mandibular barbel. Head is relatively short and broad, and snout is long and depressed. Rostral projection is narrow, bears a single spine near base, and is directed anteriorly. Nasal spine is well developed. Lower jaw has a single serrated ridge. Lip barbels are slender, number 2, and are separated. Chin barbels number 8 or 9 and are arranged in four or five assemblages usually of 3, 2, 2, and 1 or 2 barbels, with anterior cluster consisting of 1 large and 2 small barbels. Filamentous mandibular barbel is long and generally extends to end of perifacial rim. Perifacial rim terminates as broad right-angled wing or flat spinelike projection, with lateral margin interrupted by spines and crenulations, and posterior margin slightly to moderately concave and possessing one or two spines. Gill rakers number 2 or 3 on upper limb, 1 in corner, and 17 to 21 on lower limb. Measurements are expressed as percent of SL: head length 33%–37%, head depth 15%– 19%, rostral projection length 5%– 6%, snout length 16%–17%, mandibular barbel length 21%–31%, orbit length 6%– 8%, interorbital width 6%– 8%, pectoral fin length 14%–17%,

first free pectoral fin length 20%–21%, body depth 18%–22%. Joined pectoral fin rays number 12 or 13. First dorsal fin has seven or eight spines, and second dorsal fin has 17 or 18 rays. Anal fin has 17 to 19 rays. Scutes in dorsal series number 24 to 26, in supramedian series number 28 to 30, in inframedian series number 23 to 26, and in ventral series number 19 to 21, with anterior pointed end of first ventral scute posterior to anterior edge of bony knob of pelvic girdle. Color in life is orangish red, with dorsal fin margins reddish black. In preservative, body is straw colored, and margins of dorsal fins are black. This species occurs between 64 and 910 m in the western Atlantic from Georges Bank to Brazil, including the entire Gulf of Mexico. In the Gulf of Mexico it is common off the west coast of Florida. Food consists of shrimps, stomatopods, and other small crustaceans. Maximum known size is about 300 mm SL. references: Longley and Hildebrand 1941; Bigelow and Schroeder 1953; Teague 1961; Hoese and Moore 1977, 1998; Darcy and Gutherz 1984; Robins and Ray 1986; Cervigón 1991; Boschung 1992; Richards and Miller 2002b.

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PERISTEDIIDAE 81

Peristedion thompsoni Rimspine searobin

Fowler, 1952

Peristedion thompsoni has very short, broad rostral projections and perifacial rims terminating as flat spines. Head is high and very broad, and snout is rather short and broad. Rostral projection is directed slightly laterally and bears a small spine on base. Nasal spine is present in juveniles but not in adults. Lower jaw has a single serrated ridge. Lip barbels are finely pointed, number 5 or 6 to 12, and are arranged in two clusters of 3 to 6. Chin barbels are flattened and sharp pointed, number 12 to 18, and are arranged in four clusters. Filamentous mandibular barbel is slightly longer than longest chin barbel. Perifacial rim is rather narrow. Gill rakers number 2 on upper limb and 18 to 22 on lower limb. Measurements are expressed as percent of SL: head length 37%– 42%, head depth 19%–21%, rostral projection length 6%–7%, snout length 15%–16%, orbit length about 9%, interorbital width about 10%, mandibular barbel length 10%– 14%, first free pectoral fin ray length 20%–

24%, first joined pectoral fin ray length 20%– 22%, body depth 20%–22%. Joined pectoral rays number 12. First dorsal fin has seven spines, and second dorsal fin has 15 rays. Anal fin has 16 rays. Scutes in dorsal series number 26, in supramedian series number 32, in inframedian series number 23, and in ventral series number 21, with anterior pointed end of first ventral scute on line with or anterior to anterior edge of bony knob of pelvic girdle. Color in preservative is yellowish, with mottled spotting on pectoral fin. This species occurs between 115 and 475 m in the western Atlantic from North Carolina to Brazil, including the northern Gulf of Mexico and the Florida Keys, but it is absent from the Greater and Lesser Antilles. Maximum known size is 250 mm SL. references: Fowler 1952; Teague 1961 (in part as P. schmitti); Miller 1967b; Robins and Ray 1986; Richards and Miller 2002b.

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Peristedion truncatum

(Günther, 1880)

Peristedion truncatum has moderately long and moderately broad rostral projections and short mandibular barbels. Head is moderately long, slightly depressed, and moderately broad, and snout is long and depressed. Rostral projection is very slightly laterally oriented and bears a small spine on base. Nasal spine is small or usually absent. Lower jaw has a single serrated ridge. Lip barbels are very slender and number 5 or 6 in three clusters of 2, 1 or 2, and 2. Chin barbels are very slender, number 17 to 20, and are arranged in six clusters. Filamentous mandibular barbel extends beyond end of perifacial rim. Perifacial rim terminates as obtuseangled wing, with lateral margin undulating and posterior margin convex. Gill rakers number 3 or 4 on upper limb and 21 to 25 on lower limb. Measurements are expressed as percent of SL: head length 34%–37%, head depth 17%–18%, rostral projection length 8%– 10%, snout length about 16%, mandibular barbel length 15%–18%, orbit length 8%– 9%, interorbital width about 8%, pectoral fin length 16%–21%, first free pectoral fin

ray length 23%–25%, body depth 16%–18%. Joined pectoral fin rays number 12. First dorsal fin has eight spines, and second dorsal fin has 21 rays. Anal fin rays number 18. Scutes in dorsal series number 26 or 27, in supramedian series number 31, in inframedian series number 25, and in ventral series number 22, with anterior pointed end of first ventral scute posterior to anterior edge of bony knob of pelvic girdle. Color in preservative is light brown dorsally and tan ventrally, with distal half of first dorsal fin black, second dorsal fin mottled at base, and pectoral fin dusky. This species occurs between 155 and 910 m in the western Atlantic from New Jersey to northern Brazil, including the Gulf of Mexico, the Straits of Florida, and the Lesser Antilles. In the Gulf of Mexico it is common off Florida. Maximum known size is about 170 mm SL. references: Longley and Hildebrand 1941 (as P. springer); Teague 1961 (in part as P. springer); Bullis and Thompson 1965; Miller 1967b; Uyeno and Sato 1983d; Richards and Miller 2002b.

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PERISTEDIIDAE 83

Peristedion unicuspis

Miller, 1967

Peristedion unicuspis has moderately long, narrow rostral projections and numerous lip and chin barbels. Head is moderately broad, and snout is long and relatively depressed. Rostral projection is slender, directed anteriorly, and lacks spine near base. Nasal spine is absent. Lower jaw has one prominent serrated ridge. Lip barbels are slender and arranged in two groups of 2. Chin barbels are slender, number 8 or 9, and are grouped in five clusters of 2, 2, 2, 1 or 2, and 1. Filamentous mandibular barbel is very long, extends beyond first ventral scute, with filaments arranged in clusters of two on one side and one on the other side, and terminates as a single long filament. Perifacial rim terminates as broad right-angled wing. Margin of perifacial rim bears strong spines and crenulations. Gill rakers on first arch number 5 on upper limb and 21 on lower limb. Measurements are expressed as percent of SL: head length 37%–38%, head width 32%– 33%, first free pectoral fin ray length 22%– 24%, first joined pectoral fin ray length 17%– 19%, body depth 19%–21%; and as percent of head length: rostral projection length 34%– 35%, snout length 47%– 48%, orbital length 25%–26%, mandibular barbel length 79%–

92%. Joined pectoral fin rays number 12, and uppermost free fin ray extends beyond joined pectoral fin rays. First dorsal fin has eight spines, and second dorsal fin has 19 to 20 rays. Anal fin has 20 rays. Scutes in dorsal series number 25, in supramedian series number 32, in inframedian series number 24, and in ventral series number 20, with anterior pointed end of first ventral scute posterior to anterior edge of bony knob of pelvic girdle. Supramedian scutes have a single posteriorly directed retrorse cusp. Vertebrae number 33. Color in life is pale gray, with distal half of membrane between first five dorsal fin spines black and small black spot on center of pectoral fin. Color in preservative is pale, with distal half of first dorsal fin black and black spot on center of pectoral fin. This species occurs in the western Atlantic from Cay Sal Bank in the Straits of Florida, Tongue of the Ocean in the Bahamas, and possibly the eastern Gulf of Mexico. Depth of capture is 530 m. Maximum known size is 160 mm SL. references: Miller 1967a; Richards and Miller 2002b.

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Peristedion

sp.

Peristedion sp. has long, narrow rostral exsertions and numerous chin barbels. Head is short, narrow, and relatively depressed, and snout is very long. Rostral projection is slightly to moderately laterally oriented and lacks spine. Nasal spine is lacking. Perifacial rim is very narrow and terminates as narrow, short, posteriorly directed spine. Lip barbels number 5 in two clusters of 2 and 3. Chin barbels number 34 to 57 and are arranged in seven clusters. Gill rakers number 3 to 5 on upper limb and 17 to 26 on lower limb. Measurements are expressed as percent of SL: head length 33%–35%, head depth 12%–14%, rostral projection length 16%–19%, snout length 16%–17%, mandibular barbel length 10%–13%, orbit length 7%– 8%, interorbital width 6%–7%, joined pectoral fin length 21%–23%, first free pectoral fin ray length 19%–22%, body depth

13%–15%. Joined pectoral fin rays number 11 or 12. First dorsal fin has eight spines, and second dorsal fin has 20 or 21 rays. Anal fin has 21 or 22 rays. Scutes in dorsal series number 28, in supramedian series number 30, in inframedian series number 26, and in ventral series number 22. Bicuspid spines in supramedian series number 9 or 10. Color in preservative is pale yellow. Dorsal fins have submarginal dark stripes. This species occurs between 219 and 549 m in the western Atlantic from the Bahamas and the Greater and Lesser Antilles and along the coast of Belize and Honduras. It is not known from the Gulf of Mexico, but it may occur there. Maximum known size is 170 mm SL. references: George C. Miller (pers. comm., August 1, 1995); Richards and Miller 2002b.

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85

PERCIFORMES Perciformes are the most taxonomically and morphologically diverse of the orders of fishes. For these reasons, they remain poorly known and, most likely, do not constitute a monophyletic group. The order consists of 17 suborders (Weitzman 1995), 153 families (Weitzman 1995), about 1,496 genera (Nelson 1994), and about 9,293 species (Nelson 1994). One of the suborders recognized by Weitzman (Mugiloidei), with a single family, Mugilidae, is herein considered a separate order, sister to the Atherinomorpha, following Staissny (1990, 1993), Johnson and Patterson (1993), and Nelson (1994), and was treated in the first volume. Future research may drastically alter the composition of the Perciformes. Thirteen of the 16 suborders, 68 of the 152 families, 286 of the approximately 1,496 genera, and 817 of the approximately 9,293 species occur in the western Central Atlantic, and 67 families, about 234 genera, and about 530 species occur in the Gulf of Mexico. The classification of perciforms and numbers of families, genera, and species follow Nelson (1994), except that herein we recognize Howella and Symphysanodon each in their own family because none exclusively share derived characters with any of the other perciform families.

Key to the Families of the Gulf of Mexico (Adapted in part from characters in Smith and Heemstra 1986c and family key in Smith-Vaniz et al. 1999) 1a. Pelvic fins developed, each with 1 spine and 4 or 5 rays .............................................2 1b. Pelvic fins reduced, each with 0 or 1 spine and fewer than 4 rays, or absent (luvarids have pelvic fins reduced to flaplike structures in front of anus, and microdesmids have reduced pelvic fins, with 1 spine and 2 to 5 rays) . . . . . . 59 2a. Pelvic fins close together, usually united to form sucking disc (pelvic fins separate in western Atlantic gobiid genera Chriolepis, Ptereleotris, Pycnoma, and Varicus) . . . . . . . . 3 2b. Pelvic fins generally moderately to widely separated and not forming sucking disc . . . . . . . . . . . . . . . . . . . . . . . . . 4 3a. Pelvic fins generally consisting of 1 small spine and 4 rays, rarely consisting of 1 small spine and 5 rays; single dorsal fin consisting entirely of 5 to 19 rays [body depressed anteriorly and compressed to subcylindrical posteriorly; dorsal fin located over anal fin; pectoral fins modified and supporting sucking disc] . . . . . . . . . . . . . Gobiesocidae p. 641 3b. Pelvic fins consisting of 1 spine and 5 rays (Ptereleotris has 1 spine and 4 rays, but spines are also present in first dor-

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86 PERCIFORMES

sal fin); 1 or 2 dorsal fins with 5 to 17 spines and 5 to 51 rays [body generally stout; head short and broad; lateral line absent on body] . . . . . . . . . . . . . . . . . Gobiidae p. 661 4a. Pelvic fins subabdominal, pelvic fin base distinctly behind pectoral fin insertion . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4b. Pelvic fins thoracic or jugular, pelvic fin base slightly behind, under, or in front of pectoral fin base . . . . . . . . . 6 5a. Lower 3 to 16 pectoral fin rays filamentous and free of membranes [body oblong and moderately compressed; snout conical and protruding beyond mouth; two widely separated dorsal fins; lateral line extending to posterior margin of caudal fin] . . . . . . . . . . . . . . Polynemidae p. 405 5b. Lower pectoral fin rays not filamentous and connected by membranes [body elongate and subcylindrical to slightly compressed; snout long and pointed; jaw teeth large, sharp, and flattened to conical] . . . . . . . . . . . Sphyraenidae p. 747 6a. Top of head covered with bony plates [body moderately elongate and either depressed or compressed anteriorly; eyes on top of head and directed dorsally] . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uranoscopidae p. 567 6b. Top of head not covered with bony plates . . . . . . . . . . . . 7 7a. Chin with pair of long, slender barbels [body moderately elongate and moderately compressed; mouth low on head; dorsal fins well separated and short] . . . . . Mullidae p. 447 7b. Chin without pair of long, slender barbels . . . . . . . . . . . . 8 8a. Upper jaw elongated to form flat, bony, swordlike or rounded, bony, spearlike bill [body elongate and cylindrical; two widely separated dorsal fins and anal fins in adults] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xiphiidae p. 789

8b. Upper jaw not elongated into bill . . . . . . . . . . . . . . . . . . . 9 9a. Top of head with adhesive disc bearing transverse lamellae [body elongate and subcylindrical; head wide and depressed] . . . . . . . . . . . . . . . . . . . . . . . Echeneidae p. 253 9b. Top of head without adhesive disc . . . . . . . . . . . . . . . . . 10 10a. Body fusiform; series of finlets behind second dorsal and anal fins; two or more keels on caudal peduncle. . . . . . . 11 10b. Body variable in shape; series of finlets lacking behind sec-

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ond dorsal and anal fins; two caudal peduncle keels absent except in ariommatids and tetragonurids . . . . . . . . . . . . 12 11a. Jaw teeth small; lateral line straight to moderately wavy; color bluish gray, generally with dark bars, spots, or other markings [body elongate; snout pointed; finlets behind second dorsal and anal fins; first dorsal fin high anteriorly] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scombridae p. 771 11b. Jaw teeth large, often fanglike; lateral line strongly sinuous; color brown or bluish brown without dark markings [body elongate; snout pointed; finlets behind second dorsal and anal fins; first dorsal fin uniformly low] . . . . . . . . . . . . . . . . . . . . . . . . Gempylidae (in part) p. 753 12a. Body completely naked, free of scales (all chiasmodontids except Dysalotus are naked as adults; juvenile and adult Dysalotus are sparsely covered with spinoid scales) . . . . 13 12b. Body at least partially scaled (scales limited to caudal peduncle in some gempylids) . . . . . . . . . . . . . . . . . . . . . 15 13a. Head rugose and pitted with sensory pores; mouth and stomach extremely distensible [body elongate; snout longer than eye; mouth extending beyond eye; two separated dorsal fins, second one long] . . . . . . . Chiasmodontidae p. 550 13b. Head not rugose and pitted with sensory pores; mouth and stomach not extremely distensible . . . . . . . . . . . . . . . . . 14 14a. Preoperculum with stout spine; operculum and suboperculum without spines [body elongate; head broad and depressed; opercular opening is a small dorsal or sublateral pore] . . . . . . . . . . . . . . . . . . . . . . . . Callionymidae p. 646

14b. Preoperculum without spine; operculum and suboperculum with stout spines [body elongate and rounded; head broad and depressed; gill opening wide] . . . Draconettidae p. 652

15a. Anal fin preceded by 1 (usually 2) free spines (embedded in some adults) [body ranging from elongate and fusiform to deep and strongly depressed]. . . . . . . Carangidae p. 269 15b. Anal fin not preceded by 1 or 2 free spines. . . . . . . . . . . 16 16a. Single dorsal fin, may be notched to half of fin height . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16b. Two separate dorsal fins, or single dorsal fin notched more than half of fin height . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 17a. Dorsal fin base shorter than head length and consisting of

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4 to 7 spines and 7 to 13 rays [body moderately elongate to deep and compressed; snout short; eye very large; anal fin base long]. . . . . . . . . . . . . . . . . . . . Pempheridae p. 453 17b. Dorsal fin base longer than head length . . . . . . . . . . . . . 18 18a. Dorsal fin spines absent or weak and flexible, little differentiated from rays (except first 1 to 3 spines are stout in Bramidae). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 18b. Dorsal fin spines rigid and clearly differentiated from rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 19a. Maxilla covered with scales in adults; scales thick and adhesive; lateral line absent or obsolete [body compressed and ovate to elongate; jaws very oblique; snout and lower jaw free of scales] . . . . . . . . . . . . . . . . . . . Bramidae p. 304 19b. Maxilla not covered with scales; scales cycloid, thin, and deciduous; lateral line well developed. . . . . . . . . . . . . . . 20

20a. Dorsal fin high and sail-like; dorsal and anal fins depressible into membranous sheath along fin bases [body moderately elongate and compressed; pelvic fins very long] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caristiidae p. 311 20b. Dorsal fin variable but not sail-like; dorsal and anal fins not depressible into membranous sheath along fin bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 21a. Dorsal fin is small and originates posterior to midbody [body moderately elongate and very compressed; dorsal profile of body nearly horizontal] . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bathyclupeidae p. 455 21b. Dorsal fin is very long and originates anterior to midbody . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22a. Dorsal fin origin anterior to upper end of gill opening; pelvic fin depressible into groove on belly; jaw teeth in bands [body elongate and compressed; mouth large and with many small teeth] . . . . . . . . . . Coryphaenidae p. 265 22b. Dorsal fin origin over pectoral fin base; pelvic fin not depressible into groove on belly; jaw teeth uniserial [body elongate, shallow to deep, and moderately compressed; jaw teeth fine and uniserial] . . . . . . . . . . Centrolophidae p. 796

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23a. Nostril single on each side of snout; lateral line in two sections, one following dorsal contour and other on midlateral aspect of caudal peduncle [body moderately elongate to elliptical; mouth small and with a single row of incisiform teeth] . . . . . . . . . . . . . . . . . . . . Pomacentridae p. 487

23b. Nostril paired on each side of snout; lateral line, if present, not as above (23a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 24a. Caudal peduncle with forwardly directed spine or keeled bony plate on each side [body oval in profile and compressed; mouth terminal and low on head] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acanthuridae p. 740 24b. Caudal peduncle without forwardly directed spine or keeled bony plate on each side . . . . . . . . . . . . . . . . . . . . 25 25a. Scales on body cycloid, lacking spiny projections (smooth to touch) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 25b. Scales on body ctenoid, possessing spiny projections (rough to touch) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 26a. Upper jaw very protrusible and premaxilla arched ventrally when protracted; scales deciduous [body oblong or oval and compressed; bases of dorsal and anal fins covered by scaly sheath]. . . . . . . . . . . . . . . . . . . . . . . Gerreidae p. 344 26b. Upper jaw not very protrusible (except in labrid Lachnolaimus) and premaxilla not arched ventrally when protracted; scales not markedly deciduous. . . . . . . . . . . . . . 27 27a. Jaw teeth united at bases or fused into beak; upper jaw not protrusible; anal fin rays 9 [body oblong and moderately compressed; scales large and cycloid] . . . . . Scaridae p. 527 27b. Jaw teeth separate; upper jaw moderately protrusible; anal fin rays usually 10 to 12 (9 in Doratonotus) [body moderately elongate to elongate and slightly to very compressed; scales variable in size and cycloid] . . . . . . . Labridae p. 504

28a. Pelvic fins with 1 spine and 5 rays, with outer 2 rays stout and unbranched [body moderately elongate and tapering distally; mouth very large] . . . . . . . Opistognathidae p. 199 28b. Pelvic fin with 1 spine and 5 branched rays . . . . . . . . . . 29 29a. Lower 5 to 7 pectoral fin rays unbranched and enlarged, with connecting membranes deeply incised; cirri behind

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tips of dorsal fin spines [body oblong and moderately compressed; posterior margin of anterior naris fringed with cirri] . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cirrhitidae p. 485 29b. Lower pectoral fin rays usually branched and not enlarged; cirri present or absent behind tips of dorsal fin spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 30a. Jaws with outer series of incisiform teeth [body oblong to elliptical and moderately compressed; snout short] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosidae p. 482 30b. Jaws without outer series of incisiform teeth (sparids have incisiform teeth along anterior sections of jaws only) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 31a. Teeth in posterior sections of jaws molariform [body oblong, moderately deep, and compressed] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sparidae p. 385 31b. Teeth in posterior sections of jaws not molariform . . . . . 32 32a. Maxilla and lower jaw covered with scales; pelvic fins broadly joined to body by membranes [body deep and compressed; eyes very large; mouth strongly oblique] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Priacanthidae p. 211 32b. Maxilla and lower jaw not covered with scales (except in some serranids); pelvic fins not broadly joined to body by membranes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 33a. Jaw teeth bristlelike and arranged in bands; body deep and very compressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 33b. Jaw teeth not bristlelike; body variably shaped. . . . . . . . 35 34a. Preoperculum with prominent spine at angle; axillary process on pelvic fin base absent [body oblong to oval and deep; snout short and not produced; eyes dorsally located] . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pomacanthidae p. 471

34b. Preoperculum without prominent spine at angle; axillary process on pelvic fin base present [body oval and deep; snout produced; eyes slightly above body axis] . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodontidae p. 458

35a. Caudal fin broadly rounded; posterior lobelike sections of dorsal and anal fins similar in size and shape and extending posteriorly about half the length of caudal fin [body oblong, deep, and compressed; preoperculum finely to coarsely serrate]. . . . . . . . . . . . . . . . . . . . Lobotidae p. 342

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35b. Caudal fin variably shaped but not broadly rounded; posterior lobes of dorsal and anal fins variably shaped but not extending posteriorly half the length of caudal fin . . . . . 36 36a. Dorsal margin of maxilla fully exposed when mouth closed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 36b. Dorsal margin of maxilla partially covered by lachrymal bone when mouth closed . . . . . . . . . . . . . . . . . . . . . . . . 39 37a. Teeth absent in vomer and palatines; dorsal fin with 4 to 7 spines; anal fin with 1 or 2 spines [body oblong or elongate and moderately deep or slender; snout blunt or rounded] . . . . . . . . . . . . . . . . . . . . . Malacanthidae p. 243 37b. Teeth present in vomer and palatines; dorsal fin with 9 to 13 spines (except 3 or 4 in serranid Rypticus); anal fin with 0 or 3 spines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 38a. Three flat spines on posterior margin of operculum, with middle spine prominent; lateral line complete [body variable, deep to elongate and little compressed to compressed; preoperculum margin usually serrate] . . . Serranidae p. 119

38b. One or no flat spines on posterior margin of operculum; lateral line interrupted or absent [body moderately elongate and compressed; eyes large] . . . . Grammatidae p. 194

39a. Front of jaws with one or two pairs of prominent canine teeth; vomer with tooth patch [body oblong and moderately compressed; maxilla concealed by lachrymal bone when mouth closed] . . . . . . . . . . . . . . . . Lutjanidae p. 317 39b. Front of jaws without canine teeth; vomer without tooth patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 40a. Upper surface of lower jaw with pronounced, angular step; only uppermost margin of maxilla covered by lachrymal bone when mouth closed [body slender to moderately deep and moderately compressed; lower jaw usually with large teeth; most of head covered with scales] . . . . . . . . . . . . . . . . . . . . . . . Symphysanodontidae p. 338 40b. Upper surface of lower jaw without pronounced, angular step; most of maxilla covered by lachrymal bone when mouth closed [body oblong, moderately deep, and compressed; chin with two or six pores; posterior margin of preoperculum slightly concave and serrate] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haemulidae p. 359

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41a. Upper jaw very protrusible; jaw teeth either absent or small, conical, and limited to front of jaws; rayed dorsal and anal fins with scaly basal sheaths that broaden distally to cover last 2 or 3 rays . . . . . . . . . . . . . . . . . . . . . . . . . 42 41b. Character combinations not as above (41a) . . . . . . . . . . 43 42a. Maxilla broadly expanded posteriorly, covered with scales, and free of lachrymal bone when mouth closed; first dorsal fin with 7 to 13 spines [body fusiform and moderately slender; upper and lower profiles of head symmetrical; caudal fin deeply forked] . . . . . . . . . . . . . Emmelichthyidae p. 313 42b. Maxilla not broadly expanded posteriorly, naked, and covered by lachrymal bone when mouth closed; first dorsal fin with 9 or 10 or 14 to 17 spines [body elongate and fusiform; dorsal profile of head straight to slightly convex; caudal fin deeply forked] . . . . . . . . . . . . Inermiidae p. 382 43a. Two separate, short dorsal fins, first with 6 to 10 spines, second with 7 to 14 rays . . . . . . . . . . . . . . . . . . . . . . . . 44 43b. Two dorsal fins, one of which is long (containing 14 or more elements), or a single deeply notched dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 44a. Bases of second dorsal and anal fins covered with scales (in epigonids only Epigonus has scales on fins, but other genera do not occur in Gulf of Mexico) . . . . . . . . . . . . . . . . 45 44b. Bases of second dorsal and anal fins free of scales . . . . . 46 45a. Eye diameter equal to or greater than snout length (Gulf of Mexico species); first dorsal fin with 6 to 8 spines; operculum with 1 or 3 spines on posterior margin; no auxillary scales above or below lateral line scales on caudal fin [body short or elongate and subcylindrical or compressed; mouth large and oblique] . . . . . . . . . . . . . . . . . Epigonidae p. 235 45b. Eye diameter less than snout length; first dorsal fin with 8 to 10 spines; operculum with 2 spines on posterior margin; row of auxillary scales above and below lateral line scales on caudal fin [body oblong and slightly compressed; caudal peduncle rather deep]. . . . . . . . . . . . Moronidae p. 105 46a. Operculum without spines; anal fin with 1 to 3 spines and 6 to 12 rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 46b. Operculum with 1 to 3 spines; anal fin with 2 or 3 spines and 6 to 18 rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 47a. Lateral line extending to posterior margin of caudal fin; axillary process on pelvic fin base [body moderately elongate and moderately compressed; snout long and acute] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centropomidae p. 97

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47b. Lateral line not extending onto caudal fin; no axillary process on pelvic fin base [body stout; head short and broad; head usually with series of sensory canals, pores, and papillae; lateral line absent] . . . . . . . . . . . Eleotridae p. 654 48a. Operculum with 1 spine on posterior margin; anal fin with 2 spines and 8 to 18 rays [body ovate to elongate and moderately compressed; posterior margin of preoperculum double edged and smooth to serrate] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apogonidae p. 216 48b. Operculum with 2 or 3 spines on posterior margin; anal fin with 2 or 3 spines and 6 to 8 rays . . . . . . . . . . . . . . . . . 49 49a. Jaw teeth minute; no teeth in vomer or palatine; lateral line interrupted [body elongate, slender, and compressed; snout short and convex; eyes large] . . . . . . . . . Howellidae p. 117 49b. Jaw teeth large and caninelike (minute in Bathysphyraenops); teeth in vomer and palatine (except Bathysphyraenops); lateral line complete [body oblong, slender, and moderately compressed; snout is short and convex] . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acropomatidae p. 111 50a. First dorsal fin base longer than second dorsal fin base, or spinous section of dorsal fin longer than rayed section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 50b. First dorsal fin base shorter than second dorsal fin base, or spinous section of dorsal fin shorter than rayed section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 51a. Lower limb of first gill arch with slender and flat gill rakers; jaw teeth very small; caudal peduncle almost square in cross section [body very elongate and nearly round in cross section; caudal peduncle long, with two scaly keels on each side]. . . . . . . . . . . . . . . . . . . . . . . . . Tetragonuridae p. 815 51b. Lower limb of first gill arch with denticulate or tubercular gill rakers, or gill rakers absent; jaw teeth large, slender, and sharp pointed; caudal peduncle compressed . . . . . . 52 52a. Lower limb of first gill arch with four or five denticulate gill rakers; pectoral fins longer than head length origin [body moderately elongate and compressed; eyes very large; pectoral fins large and nearly reaching anal fin origin] . . . . . . . . . . . . . . . . . . . . . . . . . Scombrolabracidae p. 745 52b. Lower limb of first gill arch without gill rakers, or with gill rakers represented by tubercles; pectoral fin shorter than head length [body elongate and compressed; fanglike teeth in jaws] . . . . . . . . . . . . . . . . . . Gempylidae (in part) p. 753

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53a. Spines of first dorsal fin considerably shorter than rays of second dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 53b. Spines of first dorsal fin not considerably shorter than rays of second dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 54a. Spines of first dorsal fin not joined by membrane and not depressible into groove; jaw teeth small and arranged in bands [body elongate and subcylindrical in cross section; head broad and depressed] . . . . . . . Rachycentridae p. 263 54b. Spines of first dorsal fin connected by membrane and depressible into groove; jaw teeth sharp and compressed [body robust and moderately compressed; head large] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pomatomidae p. 251 55a. Caudal peduncle with fleshy keel at base of upper and lower caudal fin lobes; body with symmetrical dorsal and ventral profiles [body elliptical to elongate and compressed; caudal peduncle short and slender; eyes large and located on horizontal axis of body] . . . . . . . . Ariommatidae p. 810 55b. Caudal peduncle without keels; body shape variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 56a. Lateral line extends to caudal fin margin; barbels and sensory pores often well developed on snout and chin [body moderately elongate and moderately compressed; enlarged teeth in outer row of upper jaw and inner row of lower jaw; base of rayed section of dorsal fin longer than base of anal fin]. . . . . . . . . . . . . . . . . . . . . . . . . . Sciaenidae p. 410

56b. Lateral line extends to base of caudal fin; barbels and sensory pores not developed on snout and chin. . . . . . . . . . 57 57a. Body deep and compressed; gill rakers on upper limb of first arch comblike [body oblong or orbicular and deep and compressed; head short; eyes moderately large and located above horizontal axis] . . . . . . . . Ephippidae p. 735 57b. Body elongate and moderately compressed or depressed anteriorly; gill rakers on upper limb of first arch lathlike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58 58a. Body with symmetrical dorsal and ventral profiles; maxilla partially to totally covered by lachrymal bone; lateral line following dorsal contour [body moderately elongate, slender to deep, and compressed; caudal peduncle rather short, deep, and compressed] . . . . . . . . . . . . . . Nomeidae p. 801

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58b. Body depressed anteriorly and with asymmetrical dorsal and ventral profiles; maxilla free of lachrymal bone; lateral line running along body axis [body elongate; eyes large above horizontal axis] . . . . . . . . . . . . . Percophidae p. 563 59a. Pelvic fins absent or represented by scalelike spine or flaplike structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 59b. Pelvic fins reduced to 0 or 1 spine and 1 to 3 rays (microdesmids with 1 spine and 2 to 5 rays) . . . . . . . . . 64 60a. Body oval in profile and strongly compressed; a single nostril on each side; fleshy keel on caudal peduncle [head moderately elevated; snout blunt; caudal peduncle slender; caudal fin lunate] . . . . . . . . . . . . . . . . . . . Luvaridae p. 738 60b. Combination of characters not as above (60a) . . . . . . . . 61 61a. Body elongate; head oval or flattened; gill openings restricted; dorsal, anal, and caudal fins confluent [body eel-like; nares single] . . . . . . . . . . . . . . . . Zoarcidae p. 545 61b. Body elongate and compressed or deep and compressed; head compressed; gill openings unrestricted; dorsal, anal, and caudal fins not confluent . . . . . . . . . . . . . . . . . . . . . 62 62a. Body deep and compressed; dorsal fin single and similar in size and shape to anal fin; eye covered with adipose eyelid [caudal peduncle short, deep, and very compressed; eyes rather small and located on horizontal axis; pelvic fins absent] . . . . . . . . . . . . . . . . . . . . . . . . Stromateidae p. 817 62b. Body elongate to very elongate and compressed; dorsal fin single or double and considerably longer than anal fin; eyes lack adipose eyelid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 63a. Caudal fin well developed; spinous section of dorsal fin longer than rayed section [body elongate and compressed; teeth in front of upper jaw fanglike] . . . . . . . . . . . . . . . . . . . . . . . . Gempylidae (in part) p. 753 63b. Caudal fin very small or absent; spinous section of dorsal fin shorter than rayed section [body very elongate and compressed; nares single] . . . . . . . . . . . Trichiuridae p. 764 64a. Dorsal fin divided into three distinct segments, first two with spines and third with rays [body slender; jaw teeth conical and arranged in broad bands; pelvic fins with 2 or 3 rays and 1 embedded spine] . . . . . . Tripterygiidae p. 572 64b. Dorsal fin single or divided into one spinous and one rayed component . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 65a. Pelvic fins are thoracic and insert below pectoral fin bases; dorsal fin is single and very long; lateral line is absent;

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scales are absent or embedded [body elongate and very compressed; eyes large and laterally located] . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microdesmidae p. 730 65b. Combination of characters not as above (65a) . . . . . . . . 66 66a. Mouth extremely oblique; lips usually fringed; upper section of gill cover subdivided into fingerlike elements [body elongate, tapering, and compressed distally; head broad and deep; eyes superior]. . . . . . . . . Dactyloscopidae p. 578 66b. Mouth not extremely oblique; lips not fringed; upper section of gill cover not subdivided into fingerlike elements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 67a. Lateral line absent; maxilla not visible externally [body elongate, moderately slender, and moderately compressed] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaenopsidae p. 610 67b. Lateral line usually complete and always consists of more than three pores; maxilla visible externally. . . . . . . . . . . 68 68a. Body naked; dorsal fin with fewer spines than rays [body moderately elongate to elongate; jaw teeth incisor-like and in a single row] . . . . . . . . . . . . . . . . . . . . Blenniidae p. 624 68b. Body with cycloid scales (except scales absent in five of six Stathmonotus species); dorsal fin with more spines than rays [body elongate, moderately slender, and moderately compressed; head with cirri] . . . . . . . Labrisomidae p. 586

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CENTROPOMIDAE

Snooks

Snooks are moderately elongate and moderately compressed, with a large mouth, two completely or partially separated dorsal fins, and a lateral line that extends to posterior margin of caudal fin. Snout is long and acute, with lower jaw projecting beyond upper jaw. Head is large and straight to concave in profile. Nostril is paired and located just anterior to eye. Teeth are villiform and in bands in jaws, vomer, palatine, and occasionally ectopterygoid. Eye is large and laterally located. Lachrymal bone is serrated along ventral margin. Preoperculum is serrated along posterior and ventral margins and usually possesses ridges parallel to posterior and ventral margins. Operculum lacks spines except in one genus that does not occur in the Gulf of Mexico. Branchiostegal rays number 7. First dorsal fin has seven or eight spines, and second dorsal fin has one spine and 5 to 11 rays. Pelvic fin is thoracic and has one spine and 5 rays. Anal fin has three spines and 5 to 9 rays. Caudal fin is rounded, truncate, or deeply forked and has 17 principal rays. Scales are ctenoid. Axillary scale is present on pelvic fin base. Vertebrae number 24 or 25: 10 or 11 precaudal and 14 caudal. Snooks occur worldwide in tropical marine, brackish, and, occasionally, freshwaters. One Indo-West Pacific species is catadromous. Eggs are pelagic and about 0.7 mm in diameter. There are 22 species in three genera, and six species in a single genus occur in the western Central Atlantic and the Gulf of Mexico. references: Greenfield 1975b; Greenwood 1976; Fritzsche and Fuiman 1982; Nelson 1994; Smith 1997; Orrell 2002a.

Key to the Species of the Gulf of Mexico (Adapted from Greenfield 1975b; Rivas 1986; and R. E. Matheson, pers. comm., December 20, 2001) 1a. Second anal fin spine, when depressed, extending to or beyond caudal fin base; 49 to 59 (usually 57 or fewer) scales in horizontal series from posttemporal region to caudal fin base; 6 to 9 (usually fewer than 9) scales between second dorsal fin origin and lateral line . . . . . . . . . . . . . . . . . . . . . . . Centropomus ensiferus p. 99 1b. Second anal fin spine, when depressed, usually not extending to caudal fin base and never extending beyond caudal fin base; 61 to 92 (usually 64 or more) scales in horizontal series from posttemporal region to caudal fin base; 9 to 16 (usually more than 9) scales between second dorsal fin origin and lateral line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Gill rakers on lower limb of first arch, excluding rudiments, 14 to 17 (usually 15); pectoral fin distinctly shorter than

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98 PERCIFORMES

2b.

3a.

3b.

4a.

4b.

5a.

5b.

pelvic fin; anal fin rays seven or eight (usually seven); 61 to 72 (usually 64 to 71) scales in horizontal series from posttemporal region to caudal fin base . . . . . . . . . . . . . . . . . . . . . Centropomus pectinatus p. 102 Gill rakers on lower limb of first arch, excluding rudiments, 8 to 12; pectoral fin equal to or longer than pelvic fin; anal fin rays five to seven (usually six); 67 to 92 scales in horizontal series from posttemporal region to caudal fin base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Scales in horizontal series from posttemporal region to caudal fin base 79 to 89 (usually 80 or more); 13 to 18 (usually 15 or more) scales between anal fin origin and lateral line; second anal fin spine, when depressed, may reach caudal fin base, and when erect, extends beyond third anal fin spine . . . . . . . Centropomus parallelus p. 101 Scales in horizontal series from posttemporal region to caudal fin base 68 to 80 (usually 78 or fewer); 10 to 16 (usually fewer than 15) scales between anal fin origin and lateral line; second anal fin spine, when depressed, not extending to caudal fin base, and when erect, may or may not extend beyond third anal fin spine . . . . . . . . . . . . . . . 4 Gill rakers on lower limb of first arch, excluding rudiments, 8 to 10 (usually 9 or fewer); 10 to 14 (usually 11 to 13) scales between anal fin origin and lateral line; dorsal fin rays 9 to 11 (usually 10) . . . . . . . . . . . . . . . . . . . . Centropomus undecimalis p. 104 Gill rakers on lower limb of first arch, excluding rudiments, 9 to 12 (usually 10 or more); 12 to 16 (usually 12 to 15) scales between anal fin origin and lateral line; dorsal fin rays 9 or 10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Dorsal fin rays 9; second anal fin spine, when erect, may or may not extend to tip of third anal fin spine . . . . . . . . . . . . . . . . . . . . . . . . . Centropomus poeyi p. 103 Dorsal fin rays 9 or 10 (usually 10); second anal fin spine, when erect, extends beyond third anal fin spine . . . . . . . . . . . . . . . . . . . . . Centropomus mexicanus p. 100

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CENTROPOMIDAE 99

Centropomus ensiferus Swordspine snook

Poey, 1860

Centropomus ensiferus has pelvic fins that reach beyond anus, very long second anal fin spines, and relatively large scales. Upper jaw extends to or slightly beyond middle of eye. Gill rakers on first arch, including rudiments, number 24 to 30. Measurements are expressed as percent of SL: head length 37%– 42%, snout length 10%–12%, upper jaw length 12%– 14%, lower jaw length 19%–21%, pectoral fin length 23%–28%, pelvic fin length 21%– 24%, anal fin origin to vent 12%–15%; and as percent of head length: body depth 65%– 72%. Pectoral fin extends about to anus and has 14 to 17 rays. Second dorsal fin has slender spine and 10 (rarely 9) rays. Anal fin has three spines, with second very broad, and 6 (rarely 5 or 7) rays. Lateral line scales number 48 to 49, and scales in horizontal series from posttemporal region to caudal fin base number

49 to 59 (usually 57 or fewer). Scales from origin of second dorsal fin to lateral line number 6 to 9 (usually 6 to 8), and scales from lateral line to origin of anal fin number 9 to 11. Color is yellow brown to brownish green dorsally and silvery on sides and ventrally, with black or dark line along course of lateral line. Pelvic fin has large dark blotch near tip. This species occurs in the western Atlantic from southern Florida to Rio de Janeiro, Brazil, including the southern Gulf of Mexico, off Veracruz, Tabasco, Campeche, and Yucatán, and the Antilles. It is rare in the Gulf of Mexico. Maximum known size is 350 mm SL. references: Fraser 1968, 1978b; Greenfield 1975b; Rivas 1986; Robins and Ray 1986; Cervigón 1991, 1993a; Smith 1997; Castro-Aguirre et al. 1999; Orrell 2002a.

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100 PERCIFORMES

Centropomus mexicanus

Bocourt, 1868

Centropomus mexicanus has pelvic fins that reach to or beyond anus, moderately long second anal fin spines, and relatively small scales. Upper jaw extends to about center of eye. Gill rakers on first arch, including rudiments, number 21 to 24. Measurements are expressed as percent of SL: head length 35%–39%, snout length 9%–12%, upper jaw length 14%– 16%, lower jaw length 20%–22%, pectoral fin length 19%–22%, pelvic fin length 20%– 24%, anal fin origin to anus 13%–14%. Pectoral fin has 14 to 16 rays. Second dorsal fin has one slender spine and 10 (rarely 9) rays. Anal fin has three spines, with second spine very broad and longer than third, and 6 rays. Pored lateral line scales number 62 to 66, and scales in horizontal series from posttemporal region to caudal fin base number 68 to 78.

Scales from origin of second dorsal fin base to lateral line number 10 to 15, and scales from lateral line to origin of anal fin number 12 to 16 (usually 12 to 15). Color is yellow brown to brownish green dorsally and silvery on sides and belly, with dark line along course of lateral line. Pelvic fin is pale. This species occurs in the western Atlantic from the southern Gulf of Mexico to southern Brazil, including the Greater and Lesser Antilles. It inhabits coastal waters and estuaries. Some authors consider this species to be synonymous with C. parallelus (Greenfield 1975b). Maximum known size is 430 mm FL. references: Fraser 1968, 1978b; Rivas 1986; Cervigón 1991, 1993b; Castro-Aguirre et al. 1999; Orrell 2002a.

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CENTROPOMIDAE 101

Centropomus parallelus Fat snook

Poey, 1860

Centropomus parallelus has pelvic fins that reach to or beyond anus, very long second anal fin spines, and relatively small scales. Upper jaw extends to anterior margin of orbit. Gill rakers on first arch, including rudiments, number 21 to 25. Measurements are expressed as percent of SL: head length 35%– 40%, snout length 10%–12%, upper jaw length 13%– 16%, lower jaw length 19%–22%, pectoral fin length 20%–23%, pelvic fin length 20%–23%, anal fin origin to anus 13%–16%; and as percent of head length: body depth 67%– 81%. Pectoral fin extends to tip of pelvic fin but fails to reach anus and has 14 to 16 rays. Second dorsal fin has slender spine and 10 (rarely 9 or 11) rays. Anal fin has three spines and 6 (rarely 7) rays, with second spine very broad. Scales in horizontal series from posttemporal region to caudal fin base number 79 to 92. Scales from origin of second dorsal fin base to lateral line number 11 to 16, and scales from lateral line to origin of anal fin number 13 to 18 (usually 15 or more). Color is yellow brown to brownish

green dorsally and silvery on sides and belly, with dark line along course of lateral line. Pelvic fin is pale. This species occurs in the western Atlantic from southern Florida to Brazil, including the Gulf of Mexico and the Antilles. In the Gulf of Mexico it is known from southwestern Florida as far north as Sarasota, from south Texas, and from Veracruz, Tabasco, and Campeche. This species inhabits coastal waters and estuaries and frequently occurs in freshwater streams and lakes with connections to the ocean. Food consists of crustaceans and ray-finned fishes. Spawning takes place between April and December. Some authors consider C. mexicanus to be a synonym of this species. Maximum known size is 580 mm SL. references: Chávez 1963; Fraser 1968, 1978b; Greenfield 1975b; Lee et al. 1980; Rivas 1986; Robins and Ray 1986; Cervigón 1991, 1993b; Martin and King 1991; Smith 1997; CastroAguirre et al. 1999; Orrell 2002a.

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Centropomus pectinatus Tarpon snook

Poey, 1860

Centropomus pectinatus has pelvic fins that reach to or beyond anus, relatively short pectoral fins, and relatively small scales. Upper jaw extends slightly beyond anterior rim of orbit. Gill rakers on first arch, including rudiments, number 22 to 28. Measurements are expressed as percent of SL: head length 33%– 37%, snout length 10%–11%, upper jaw length 13%–15%, lower jaw length 19%– 22%, pectoral fin length 18%–21%, pelvic fin length 23%–26%, anal fin origin to anus 12%–16%; and as percent of head length: body depth 71%– 81%. Pectoral fin fails to reach tip of pelvic fin and anus and has 13 to 15 rays. Second dorsal fin has slender spine and 10 (rarely 11) rays. Anal fin has three spines, with third longer than second, and 7 (rarely 8) rays. Pored lateral line scales number 59 to 71, and scales in horizontal series from posttemporal region to caudal fin base number 61 to 72 (usually 64 to 71). Scales from origin of second dorsal fin to lateral line number 9 to 13, and

scales from lateral line to origin of anal fin number 11 to 14. Color is yellow brown to brownish green dorsally and silvery on sides and ventrally, with black or dark line along course of lateral line. Fins are dusky, and pelvic fin has large, dark, diffuse blotch near tip. This species occurs in the western Atlantic and eastern Pacific. In the western Atlantic it occurs from southern Florida to Bahia, Brazil, including the eastern and southwestern Gulf of Mexico. It occurs near the coast, in estuaries, and in freshwater streams with connections to the sea. Food consists of crustaceans and rayfinned fishes. Spawning is thought to take place from May to December. Maximum known size is 510 mm SL. references: Chávez 1963; Fraser 1968, 1978b; Greenfield 1975b; Rivas 1986; Robins and Ray 1986; Cervigón 1991, 1993b; Smith 1997; CastroAguirre et al. 1999; Orrell 2002a.

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CENTROPOMIDAE 103

Centropomus poeyi

Chávez, 1961

Centropomus poeyi has pelvic fins that fail to reach anus, a relatively short second anal fin spine, and relatively small scales. Upper jaw extends to or beyond posterior margin of orbit. Gill rakers on first arch, including rudiments, number 22 to 25. Measurements are expressed as percent of SL: head length 36%–38%, snout length 9%–10%, upper jaw length 14%–16%, lower jaw length 21%–23%, pectoral fin length 19%–22%, pelvic fin length 20%–22%, anal fin origin to anus 10%–11%; and as percent of head length: body depth 67%–70%. Pectoral fin fails to reach tip of pelvic fin and anus and has 15 to 17 rays. Second dorsal fin has slender spine and 9 rays. Anal fin has three spines, with second very broad and slightly shorter or longer than third, and 6 (rarely 7) rays. Pored lateral line scales number 69 to 71, and scales in horizontal series from posttem-

poral region to caudal fin base number 73 to 80. Scales from second dorsal fin origin to lateral line number 10 to 13, and scales from lateral line to anal fin origin number 12 to 15. Color is yellowish brown to brownish green dorsally and silvery on sides and ventrally, with dark line along course of lateral line. Fins are dusky. This species occurs in the western Atlantic from Tampico, Mexico, to Belize. It occurs in coastal waters, estuaries, and freshwater streams. Food consists of crustaceans and rayfinned fishes. Spawning takes place in brackish or freshwater from May through September. Maximum known size is 900 mm SL. references: Chávez 1961, 1963; Greenfield 1975b; Fraser 1978b; Rivas 1986; Smith 1997; Castro-Aguirre et al. 1999; Orrell 2002a.

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104 PERCIFORMES

Centropomus undecimalis Snook

(Bloch, 1792)

Centropomus undecimalis has pelvic fins that fail to reach anus, a moderately long second anal fin spine, and relatively small scales. Upper jaw extends to near posterior margin of orbit. Gill rakers on first arch, including rudiments, number 18 to 24. Measurements are expressed as percent of SL: head length 35%– 39%, snout length 9%–11%, upper jaw length 13%–15%, lower jaw length 20%–23%, pectoral fin length 18%–22%, pelvic fin length 18%–23%, anal fin origin to anus 9%–12%; and as percent of head length: body depth 59%– 64%. Pectoral fin fails to reach tip of pelvic fin and has 14 to 16 rays. Second dorsal fin has a single spine and 10 (rarely 9 or 11) rays. Pelvic fin fails to reach anus in specimens longer than 120 mm SL. Anal fin has three spines, with second very broad and longer than third, and 6 (rarely 5 or 7) rays. Pored lateral line scales number 67 to 72, and scales in horizontal series from posttemporal region to caudal fin base number 67 to 77. Scales from lateral line to anal fin origin number 10 to 14

(usually 11 to 13). Color is yellowish brown to brownish green dorsally and silvery on sides and ventrally, with dark line along course of lateral line and fins dusky. Pelvic fin lacks dark tip. This species occurs in the western Atlantic from North Carolina to Rio de Janeiro, Brazil, including the Gulf of Mexico and the Antilles. It is common in the southern but not the northern Gulf of Mexico. This species is associated with coastal waters to 20 m and estuaries and freshwater streams. Food consists of crustaceans and ray-finned fishes. Spawning takes place from May to October in brackish to freshwater. Maximum known size is 1,000 mm SL.

REFERENCES:

Longley and Hildebrand 1941; Chávez 1961, 1963; Fraser 1968, 1978b; Greenfield 1975b; Hoese and Moore 1977, 1998; Rivas 1986; Robins and Ray 1986; Cervigón 1991, 1993b; Boschung 1992; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999; Orrell 2002a.

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105

MORONIDAE

Temperate basses

Temperate basses are oblong and slightly compressed, with two opercular spines and a rather deep caudal peduncle. Dorsal profile of head is straight to slightly concave above orbits. Snout is slightly acute and about equal to or longer than eye length. Nostril is paired, and nares are close set. Mouth is terminal, with lower jaw even with upper jaw or slightly projecting, and upper jaw extending posterior to anterior margin of orbit. Upper jaw is slightly protrusible and lacks supramaxilla. Jaw teeth are small and conical. Villiform teeth are arranged in bands in vomer, palatines, and tongue. Posterior and ventral margins of preoperculum are serrated, and serrae are largest at corner. Opercular spines are pointed or rounded. Gill membranes are separate and joined to isthmus far anteriorly. Branchiostegal rays number 7. Gill rakers are moderately long and slender. Pectoral fin is small, with upper rays longer than lower rays. Dorsal fin is divided to base into anterior spinous and posterior rayed fins. First dorsal fin has 8 to 10 rather stout spines, and second dorsal fin has 1 spine and 10 to 14 rays, with last ray divided to base. Pelvic fin is thoracic and has 1 spine and 5 rays. Anal fin has 3 stout spines and 9 to 13 rays, with last ray divided to base. Caudal fin is emarginate to forked and has 9 branched rays in upper section and 8 branched rays in lower section. Lateral line is complete and extends to near posterior margin of caudal fin. Body and head, with exception of snout and lips, are completely covered with ctenoid scales. Scales cover medial section of second dorsal fin and anal fin, and caudal fin has auxillary row of scales on either side of lateral line scales. Pelvic axillary process is absent. Gas bladder is present and extends into hollow first anal fin pterygiophore. Vertebrae number 25: 12 precaudal and 13 caudal. Maximum known size is 180 cm TL. Temperate basses occur in the Northern and Southern Hemisphere in freshwater, brackish water, and inshore marine waters. Species are either stenohaline in freshwater or euryhaline and anadromous. There are about six species in four genera, and three species in a single genus occur in the western North Atlantic and the Gulf of Mexico. references: Johnson 1984; Page and Burr 1991; Nelson 1994.

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Key to the Species of the Gulf of Mexico (After Etnier and Starnes 1993; Jenkins and Burkhead 1994; Heemstra 2002a) 1a. Second anal fin spine greatly thickened and longer than length of anal fin base, extending beyond base when depressed; tip of lower jaw not extending beyond upper jaw. . . . . . . . . . . . . . . . . . . Morone mississippiensis p. 108 1b. Second anal fin spine slightly thickened and shorter than length of anal fin base, not extending beyond base when depressed; tip of lower jaw extending beyond upper jaw .............................................2 2a. Median tooth patches on tongue elongate and distinctly divided; lateral line scales 57 to 65; second dorsal fin with 10 to 14 rays . . . . . . . . . . . . . . . . . Morone saxatilis p. 109 2b. Median tooth patches on tongue oval shaped and single or with slight medial division; lateral line scales 54 to 58; second dorsal fin with 12 to 14 rays . . . . . . . . . . . . . . . . . . . . . . . . . . . Morone chrysops p. 107

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MORONIDAE 107

Morone chrysops White bass

(Rafinesque, 1820)

Morone chrysops has a lower jaw that projects beyond upper jaw and a second anal fin spine that is relatively short and slender. Dorsal profile of head is distinctly arched. Snout is acute and about equal to eye diameter. Anterior and posterior nares are close set, with short, broad flap on posterior margin of anterior naris. Mouth is terminal, with upper jaw extending to level of center of pupil. Jaw teeth are villiform and in bands that narrow posteriorly. Vomerine and palatine teeth are granular and in patches or bands; vomerine patch lacks distal extension. Tongue has teeth along lateral margins and one or two medial tooth patches, with length of medial patch(es) half the length of lateral patches. Preoperculum is angular, with posterior margin serrated, corner serrae enlarged, and ventral margin mostly smooth. Operculum has blunt, flat upper spine and pointed, flat lower spine. Gill rakers on first arch are long and moderately slender and number 20 to 25. Measurements are expressed as percent of SL: head length 32%–33%, snout length 7%– 8%, eye diameter 6%– 8%, body depth 32%–35%. Pectoral fin is pointed and has 15 to 17 rays. First dorsal fin has nine spines, and second has one spine and 12 to 14 rays. Anal fin has three spines, with third spine 50%–75% of fin length, and 11 to 13

rays. Caudal fin is shallowly forked. Lateral line scales number 54 to 58, scales between first dorsal fin base and lateral line number 7 to 11, and scales between lateral line and anal fin base number 5 to 17. Color in preservative is dark to dusky dorsally, grading to pale ventrally, with four to seven dark grayish brown stripes, often interrupted, along flanks. Pectoral, dorsal, and anal fins are dusky, and pelvic fin is pale. In life, color is bluish, dark green, or gray dorsally and silvery on sides and ventrally. Lateral stripes are grayish olive to dark olive. Dorsal, anal, and caudal fins are pale medially and dusky grayish blue distally. Paired fins are milky white. This species occurs naturally in the Great Lakes–St. Lawrence, Mississippi, and western Gulf of Mexico drainages. In the Gulf of Mexico it occurs in brackish waters of the Mississippi drainage to the Rio Grande. Food consists of zooplankton, insects, and larval rayfinned fishes for juvenile specimens and rayfinned fishes for subadults and adults. Maximum known size is 440 mm TL.

references: Lee et al. 1980; Page and Burr 1991; Etnier and Starnes 1993; Jenkins and Burkhead 1994.

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108 PERCIFORMES

Morone mississippiensis Yellow bass

Jordan and Eigenmann, 1887

Morone mississippiensis has a lower jaw that is even with the upper jaw, and a second anal spine that is distinctly thickened and longer than anal fin base. Profile of head is distinctly arched. Snout is acute and about equal to eye diameter. Nares are close set, with short, broad flap on posterior margin of anterior naris. Jaw teeth are in villiform bands that narrow posteriorly. Vomer has V-shaped patch of granular teeth, and palatine teeth are similarly shaped and in narrow bands. Tongue has teeth on lateral margins but lacks medial tooth patch. Preoperculum is angular, with serrae on posterior and ventral margins, and those at angle are slightly enlarged. Operculum has small, flat upper spine and large, flat lower spine. Gill rakers on first arch are long and moderately slender and number 18 to 22. Measurements are expressed as percent of SL: head length 32%– 33%, snout length 8%–9%, eye diameter 7%– 8%, body depth 30%–36%. Pectoral fin is

short and has 15 to 17 rays. First dorsal fin has nine spines, and second dorsal fin has one spine and 12 rays. Anal fin has three spines and 9 or 10 rays. Color in preservative is olive gray dorsally, grading to silver yellow on side and ventrally, with five to seven black stripes on sides. Stripes are broken and offset on lower side. Medial fins are clear to blue gray. This species occurs naturally in Lake Michigan and the Mississippi drainages, northward to Wisconsin and Minnesota, westward to Indiana, eastward to Tennessee, and southward to the Gulf of Mexico drainages between Mobile Bay and Galveston Bay. It is considered a freshwater species common in lowland habitats in pools and backwaters, but it does enter low-salinity estuaries. Maximum known size is 460 mm TL. references: Lee et al. 1980; Page and Burr 1991; Etnier and Starnes 1993.

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MORONIDAE 109

Morone saxatilis Striped bass

(Walbaum, 1792)

Morone saxatilis has a lower jaw that extends beyond upper jaw and a second anal fin spine that is relatively slender and short. Profile of head is not distinctly arched. Snout is acute and about equal to eye diameter. Nares are close set, with short, broad flap extending from posterior margin of anterior naris. Teeth in jaws are in villiform bands that narrow posteriorly. Vomer has V-shaped patch of granular teeth, and palatine teeth are similarly shaped and in narrow bands. Tongue has teeth along lateral margins and two medial tooth patches, with length of medial patches greater than half the length of lateral patches. Preoperculum is angular, with serrae on posterior and ventral margins and those at corner enlarged. Operculum has flat, blunt upper spine and flat, sharp lower spine. Gill rakers on first arch are long and moderately slender and number 19 to 29, with 6 to 12 on upper limb and 12 to 15 on lower limb. Measurements are expressed as percent of SL: head length 29%–34%, snout length 8%–9%, eye diameter 6%–9%, body depth 26%–28%. Pectoral fin is small and subacute and has 15 to 18 rays. First dorsal fin has 9 or 10 spines, and second has 1 spine and 10 to 14 rays. Anal fin has concave posterior margin and 3 spines and 9 to 12 rays, with third spine 40%– 60% of fin length. Caudal fin

is moderately to weakly forked. Lateral line scales number 57 to 65, scales between first dorsal fin base and lateral line number 9 to 11, and scales between anal fin base and lateral line number 14 to 16. Vertebrae number 25: 12 precaudal and 13 caudal. Color in preservative is dark to dusky dorsally and dusky to pale ventrally, with six to nine dark gray stripes, largely uninterrupted, on flank. Dorsal and anal fins are dusky with pale bases, pelvic fin is pale, and pectoral fin is slightly dusky. In life, dorsal surface is dark olive to steel blue, and side and ventral surfaces are silver with brassy sheen. Lateral stripes are dark olive to black. Dorsal, anal, and caudal fins are pale green to dusky green; pelvic fin is white; and pectoral fin is greenish. Juveniles lack dark stripes and have dusky bars on side. This species occurs along the Atlantic coast of North America from the St. Lawrence River to the St. Johns River in Florida, and along the Gulf slope from the Suwannee River in Florida to eastern Texas. The native populations in the Gulf of Mexico may no longer exist because of the introduction of stocks from the Atlantic coast of the United States. This species is anadromous, occupying inshore marine waters as juveniles and ascending rivers to spawn in freshwater as adults. Food consists

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110 PERCIFORMES

of polychaetes, crustaceans, insects, and rayfinned fishes for young juveniles, and clams, squids, shrimps, crabs, lobsters, and ray-finned fishes for large juveniles and adults. Maximum known size is about 1.8 m TL.

references: Hildebrand and Schroeder 1928; Hoese and Moore 1977, 1998; Hardy 1978; Robins and Ray 1986; Page and Burr 1991; Etnier and Starnes 1993; Jenkins and Burkhead 1994; Heemstra 2002a.

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111

ACROPOMATIDAE

Temperate ocean-basses

Temperate ocean-basses are oblong and moderately compressed, with dorsal fin divided to base or separated into two fins and canine teeth usually present in jaws. Snout is short and convex in profile. Interorbital region is broad and flat. Nostril is paired, and nares are close set. Eye is large and laterally located. Mouth is terminal, moderately large, and moderately oblique, and lower jaw projects slightly beyond upper jaw. Upper jaw is protrusible. Maxilla is mostly exposed when mouth is closed, is expanded posteriorly, and extends to below or beyond pupil, with supramaxilla present on upper margin. Jaw teeth consist of villiform bands and canines, and vomerine and palatine teeth are villiform. Preoperculum is angular and has serrae or spines at angle. Operculum has 2 flat spines on posterior margin. Upper margin of operculum is free. Gill membranes are separate and free of isthmus. Branchiostegal rays number 6 or 7. Gill rakers on first gill arch are well developed. Pectoral fin is longer than pelvic fin and located on lower flank. First dorsal fin consists of 7 to 10 slender spines, and second consists of 1 spine and 9, 10, or 14 rays, with last ray split to base. Pelvic fin inserts under pectoral fin base, has 1 spine and 5 rays, and lacks axillary process of fused scales on base. Anal fin has 2 or 3 slender spines and 6 to 9 rays or 12 rays. Caudal fin is moderately to deeply forked and has 9 branched rays in upper section and 8 in lower section. Scales are cycloid or occasionally ctenoid and cover body, cheek, and occasionally operculum. Lateral line is complete and consists of 25 to 55 tubed scales. Vertebrae number 25 or 26: 10 precaudal and 15 or 16 caudal. Maximum known size is 800 mm TL for western Atlantic species. Acropomatids occur worldwide in the water column between 87 and 910 m. There are about 25 species in seven genera, 6 species in three genera in the western Central Atlantic, and 4 species in two genera in the Gulf of Mexico. references: Schultz 1940; Johnson 1984; Nelson 1994; Heemstra 2002b.

Key to the Species of the Gulf of Mexico (Adapted from Heemstra 2002b) 1a. Branchiostegal rays 6; body covered with adherent ctenoid scales; anal fin with three spines and 7 rays; pectoral fin rays 13 or 14; gill rakers on first arch 7 on upper limb and 21 on lower limb . . . . . . Bathysphyraenops simplex p. 113 1b. Branchiostegal rays 7; body covered with mostly cycloid, rather deciduous scales; anal fin with two or three spines and 7 to 9 rays; pectoral fin rays 14 to 18; gill rakers

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112 PERCIFORMES

2a.

2b.

3a.

3b.

on first arch 4 to 7 on upper limb and 12 to 18 on lower limb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 First dorsal fin with eight spines; anal fin with three spines and 8 or 9 rays; lateral line scales 46 to 51; body depth 22%–27% of SL . . . . . . . . . . Synagrops trispinosus p. 116 First dorsal fin with nine spines; anal fin with two spines and 7 or 9 rays; lateral line scales 29 to 35; body depth 26%–30% of SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Anterior edge of pelvic fin spine and second spines of first dorsal and anal fins serrate; gill rakers on first arch 17 to 20 . . . . . . . . . . . . . . . . . . . . . . . Synagrops spinosus p. 115 Anterior edge of pelvic fin spine and second spines of first dorsal and anal fins smooth; gill rakers on first arch 13 to 17 . . . . . . . . . . . . . . . . . . . . . . . Synagrops bellus p. 114

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ACROPOMATIDAE 113

Bathysphyraenops simplex

Parr, 1933

Bathysphyraenops simplex has entire head covered with scales and has spines on the interoperculum and suboperculum. Dorsal profile of head is moderately convex, and snout is very short. Nares are close set, with posterior naris slitlike and about 4 times size of anterior naris. Maxilla extends to almost middle of eye. Jaw teeth are villiform and arranged in a single row. Teeth in vomer and palatine consist of rows of denticles. Preoperculum has double edge and is smooth along posterior margin but has series of small spines at corner. Suboperculum has two spines, and interoperculum has one spine. Branchiostegal rays number 6. Gill rakers on first arch number 7 on upper limb and 21 on lower limb. Measurements are expressed as percent of SL: head length 31%–33%, snout length 6%–7%, eye diameter 10%–11%, interorbital width 6%–7%, upper jaw length 12%–

13%, body depth 27%–29%. Pectoral fin is relatively long and has 13 or 14 rays. First dorsal fin originates just posterior to pectoral fin base and has eight spines. Second dorsal fin has one spine and 9 or 10 rays. Anal fin has three spines and 7 rays. Fin spines lack serrations. Scales are ctenoid, with ctenii attached to scale. Lateral line is continuous to interrupted and has 30 to 35 scales. Color is darkish brown. This species occurs worldwide in tropical and subtropical seas. In the western Atlantic it occurs from North Carolina to northern Brazil, including the southern Gulf of Mexico, the Bahamas, and the Caribbean Sea. Maximum known size is 88 mm SL.

references: Parr 1933; Schultz 1940; Heemstra 2002b.

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Synagrops bellus Blackmouth bass

(Goode and Bean, 1896)

Synagrops bellus has canine teeth in jaws, two short dorsal fins, and a pelvic spine with a smooth anterior edge. Dorsal profile of head is moderately convex, and snout is very short. Nares are close set, with posterior naris slitlike and about 4 times size of anterior naris. Lower jaw projects beyond upper jaw, and upper jaw extends to middle of eye. Teeth occur in jaws, vomer, and palatine. Upper jaw has a single canine on either side of symphysis, followed by a wide band of villiform teeth. Lower jaw has a pair of canines on either side of symphysis, followed by a band of villiform teeth in concave area adjacent to canines and a lateral row of canines and median band of villiform teeth. Teeth in vomer and palatine are villiform and arranged in bands. Preoperculum has two or three small ridges on lower posterior margin. Suboperculum and interoperculum are serrated along margins. Branchiostegal rays number 7. Gill rakers on first arch number 2 or 3 on upper limb, 1 in corner, and 10 to 13 on lower limb. Measurements are expressed as percent

of SL: head length 31%–34%, snout length 7%– 8%, eye diameter 9%–11%, interorbital width 9%–10%, upper jaw length 14%–15%, body depth 23%–27%. Pectoral fin is relatively long and has 16 or 17 rays, with upper rays longer than lower rays. First dorsal fin originates just posterior to pectoral fin base and has nine spines. Second dorsal fin has one spine and 9 rays. Anal fin has two spines and 7 rays. Fin spines lack serrations. Scales are cycloid, large, and deciduous. Lateral line is complete and consists of 29 to 31 scales. Color is brownish gray. This species occurs in the tropical and warm temperate Atlantic. In the western Atlantic it occurs between 277 and 806 m, from Canada and Bermuda to Suriname, including the northern and southern Gulf of Mexico and the Antilles. Maximum known size is 360 mm SL. references: Schultz 1940; Bullis and Thompson 1965; Fujii 1983a; Boschung 1992; Heemstra 2002b.

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ACROPOMATIDAE 115

Synagrops spinosus Keelcheek bass

Schultz, 1940

Synagrops spinosus has canine teeth in jaws, two short dorsal fins, and a pelvic fin spine with a serrated anterior edge. Dorsal profile of head is moderately convex, and snout is very short. Nares are close set, with posterior naris slitlike and several times larger than anterior naris. Lower jaw projects beyond upper jaw, and upper jaw extends to middle of eye. Upper jaw has a single canine on either side of symphysis, followed by a wide band of villiform teeth. Lower jaw has one or two canines on either side of symphysis, a band of villiform teeth in concave area adjacent to canines, and a lateral row of canines and a median band of villiform teeth posterior to concave area. Teeth in vomer and palatine are villiform and arranged in bands. Preoperculum has two or three small ridges on lower posterior margin. Suboperculum and interoperculum are serrated along margins. Branchiostegal rays number 7. Gill rakers on first arch number 17 to 20. Measurements are expressed as percent of SL: head length 31%–36%, snout length 7%– 8%, eye

diameter 9%–12%, upper jaw length 16%– 17%, interorbital width 8%–9%, body depth 25%–29%. Pectoral fin is relatively long and has 16 rays, with upper rays longer than lower rays. First dorsal fin originates slightly posterior to pectoral fin base and has nine spines, with serrations along leading edge of second spine. Second dorsal fin has one spine and 9 rays. Anal fin has two spines and 7 rays. Scales are cycloid, large, and deciduous. Lateral line scales number 29 to 31. Color is brownish gray. This species occurs in the western Atlantic and western Pacific. In the western Atlantic it occurs between 75 and 412 m, from North Carolina to Suriname, including the Gulf of Mexico. It has been captured throughout the Gulf of Mexico over its depth range. Maximum known size is about 130 mm TL.

references: Schultz 1940; Bullis and Thompson 1965; Fujii 1983a; Boschung 1992; Heemstra 2002b.

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Synagrops trispinosus

Mochizuki and Sano, 1984

Synagrops trispinosus has canine teeth in jaws, three anal fin spines, and a pelvic fin spine with a serrated anterior edge. Dorsal profile of head is moderately convex, and snout is moderately short. Nares are close set, with small flap on anterior naris. Lower jaw projects slightly beyond upper jaw, and upper jaw extends to anterior margin of pupil. Upper jaw has a single canine on either side of symphysis, followed by a wide band of villiform teeth. Lower jaw has one canine on either side of symphysis, followed by a series of about five canines and a median band of villiform teeth. Teeth in vomer and palatine are villiform and arranged in a V or triangular patch and a row, respectively. Preoperculum has two small ridges on lower posterior margin and serrations along its angle. Branchiostegal rays number 7. Gill rakers on first arch number 3 to 5 on upper limb and 12 to 16 on lower limb. Measurements are expressed as percent of SL: head length 34%– 38%, body depth 25%–29%; and as percent

of head length: snout length 25%–32%, eye diameter 26%–34%, upper jaw length 44%– 52%, interorbital length 21%–26%. Pectoral fin is moderately long and has 16 to 18 rays. First dorsal fin originates slightly posterior to pectoral fin base and has eight spines. Second dorsal fin has one spine and 10 rays. Anal fin has three spines and 8 or 9 rays. Second dorsal and anal fin spines are smooth. Lateral line scales number 46 to 51. Color in preservative is yellowish to dark purplish brown. This species occurs in the western Atlantic from the northeastern Gulf of Mexico to Suriname, including Jamaica, Puerto Rico, and the Virgin Islands. It has been captured once in the Gulf of Mexico off the Florida Panhandle (2910N, 8555W). Maximum known size is about 120 mm TL.

references: Mochizuki and Sano 1984; Heemstra 2002b.

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117

HOWELLIDAE

Howellids are slender and compressed, with strong opercular, subopercular, and interopercular spines and an interrupted lateral line. Snout is short and convex in profile. Eye is large and laterally located. Mouth is moderately large and moderately oblique, with lower jaw even with upper jaw. Upper jaw is protrusible. Nostril is paired, and nares are moderately separated. Jaws have a single row of small conical teeth. Small teeth are either present or absent in vomer and palatine. Maxilla is exposed when mouth is closed and lacks supramaxilla. Preoperculum is slightly extended as crest at angle bearing serrae. Operculum has two spines on upper angle and cluster of spines at lower angle. Upper margin of operculum is free. Suboperculum has a single strong spine or cluster of spines. Interoperculum has one to three spines. Gill membrane is free of isthmus. Branchiostegal rays number seven. Gill rakers on first arch are long and slender. Pectoral fin is long and located on lower flank. Dorsal fins are widely separated. Pelvic fin is thoracic, has one spine and five rays, and lacks axillary process of fused scales. Anal fin has three slender spines and seven rays. Caudal fin is deeply forked and has nine branched rays in upper section and eight in lower section. Scales are ctenoid and adherent. Vertebrae number 26: 10 precaudal and 16 caudal. Howellids occur worldwide in tropical and warm temperate waters and are either pelagic or benthopelagic over outer continental shelves and slopes. There are five species in one genus, and one species occurs in the western Central Atlantic, including the Gulf of Mexico. references: Johnson 1984; Roberts 1993; Nelson 1994.

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Howella brodiei Ogilby, 1899

Howella brodiei has two widely separated dorsal fins and strong opercular, subopercular, and interopercular spines. Dorsal profile of head is moderately convex, and snout is very short. Nares are moderately separated, with posterior naris larger than anterior naris and located on anterodorsal margin of eye, in front of supraorbital spine. Maxilla extends to pupil. Jaws have a single row of minute conical teeth. Vomer has a small triangular patch of granular teeth, and palatine has a long row of small conical teeth. Upper opercular spine is either simple or branched, and lower spine is branched. Subopercular spine is simple and very long. Interopercular spine is simple and shorter than subopercular spine. Gill rakers on first arch number 7 to 9 on upper limb and 19 to 22 on lower limb. Measurements are expressed as percent of SL: head length 31%–34%, snout length 7%–9%, eye diame-

ter 10%–11%, interorbital width 7%–9%, upper jaw length 14%–15%, body depth 27%– 29%. Pectoral fin is long and pointed and has 14 or 15 rays. First dorsal fin has eight spines, and second has one spine and 9 to 11 rays, with last ray split to base. Anal fin has three spines and 7 or 8 rays. Lateral line scales number 35 to 39. Color is dark brown, with head slightly lighter. This species occurs in the Atlantic and western Pacific Oceans in tropical to warm temperate waters. Juveniles are pelagic between 26 and 300 m, and adults are bathypelagic or possibly benthic. It is widespread in the Gulf of Mexico between 102 and 1,134 m. This species was previously known as Howella sherborni. Maximum known size is 120 mm TL. references: Fedoryako 1976 (as H. sherborni); Fujii 1983a; Tortonese 1986; Heemstra 2002b.

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119

SERRANIDAE

Seabasses

Seabasses are robust to moderately compressed and oblong-oval to rather elongate, with a continuous dorsal fin and three anal fin spines. Snout is short to moderately long, and generally convex in profile. Nostril is paired and located anterior to eye. Mouth is moderate to large and either terminal or with lower jaw projecting beyond upper jaw. Upper jaw is more or less protrusible, and maxilla is mostly exposed when mouth is closed and extends to below or slightly beyond eye. A single, well-developed supramaxilla is present on dorsal margin of maxilla (except absent in Serraninae). Teeth are small, slender, conical, and depressible and occur in jaws, vomer, and palatines (absent in Schultzea). Some species also have several canine teeth in jaws. Eyes are large and laterally located. Lachrymal bone lacks serrations. Preoperculum has serrate posterior margin and serrate or undulate lower margin. Upper margin of operculum is free (except in Rypticus). Rear margin of operculum has 3 (rarely 2) flat spines, with upper and lower spines often inconspicuous and covered with skin. Gill membranes are separate but joined to isthmus anteriorly. Pseudobranch is well developed. Branchiostegal rays number 7 (rarely 6). Gill rakers on first arch are short to long and occasionally bear small teeth. Pectoral fin is rounded to slightly pointed, and usually longer than pelvic fin. Dorsal fin is often notched, rarely divided to base, and has 2 to 11 spines and 10 to 27 rays. Pelvic fin is thoracic, has 1 spine and 5 branched rays, and either lacks or has a rudimentary axillary scale. Anal fin has 0 or 3 spines and 6 to 17 rays. Caudal fin is usually rounded, truncate, or lunate, rarely forked, and has 13 to 16 branched rays. Lateral line is single and does not extend onto caudal fin but is lacking in one genus. Scales are generally ctenoid but occasionally cycloid, and cover body and part of head. Vertebrae number 24 to 26: 10 or 11 precaudal and 14 to 16 caudal. Serranids occur worldwide in tropical to temperate seas and occasionally in freshwater. Most species are benthic and associated with hard bottoms to depths of 200 m, but some reach depths of 500 m and others occur on soft bottoms and sea-grass beds on continental shelves. Food consists of zooplankton, polychaetes, cephalopods, crustaceans, and ray-finned fishes. Many species are protogynous hermaphrodites, which function first as females and later the ovaries transform to testes and the fish functions as a male. Eggs and larvae are pelagic. Eggs are 0.7 to 1.2 mm in diameter, and larvae are distinguished by their greatly elongated second dorsal spine and pelvic spines. There are about 449 species in 62 genera; 89 species and 25 genera occur in the western Central Atlantic, and 61 or 62 species in 20 genera are present in the Gulf of Mexico.

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references: Johnson 1983, 1984; Kendall 1984; Heemstra and Randall 1993; Nelson 1994; Smith 1997; Heemstra 2002c.

Key to the Species of the Gulf of Mexico (Adapted from Heemstra 2002c) 1a. Dorsal fin with 2 to 4 spines and 18 to 27 rays; no anal fin spine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Dorsal fin with 8 to 11 spines and 9 to 20 rays; 3 anal fin spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 2a. Background color pale, with dark markings. . . . . . . . . . . 3 2b. Background color dark, with or without pale markings .............................................4 3a. Dorsal fin with 2 spines and 24 to 27 rays . . . . . . . . . . . . . . . . . . . . . . . . Rypticus bistrispinus p. 176 3b. Dorsal fin with 3 or 4 spines and 20 to 24 rays . . . . . . . . . . . . . . . . . . . . . . Rypticus subbifrenatus p. 180 4a. Head and body uniformly dark brown except belly slightly lighter; snout length 5%– 6% of SL . . . . . . . . . . . . . . . . . . . . . . . . . . . Rypticus randalli p. 178 4b. Body and head with pale-colored blotches or spots; snout length 6%–9% of SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5a. Dorsal fin spines 3; body with pale blotches; jaws and belly mottled; pores on lower jaw and preopercular margin numerous, small, and clustered into patches . . . . . . . . . . . . . . . . . . . . . . . . Rypticus saponaceus p. 179 5b. Dorsal fin spines usually 2; body with pale spots; jaws and belly uniformly pale; pores on lower jaw and preopercular margin distinct and single, not clustered into patches . . . . . . . . . . . . . . . . . . . . . . . . . Rypticus maculatus p. 177 6a. Upper jaw distinctly protrusible; jaws without teeth; branchiostegal rays 6 . . . . . . . . . . . . . . . . Schultzea beta p. 181 6b. Upper jaw not or only slightly protrusible; jaws with teeth; branchiostegal rays 7 (rarely 6) . . . . . . . . . . . . . . . . . . . . 7 7a. Head length 26%–29% of SL; caudal fin deeply forked, middle rays less than half the length of upper and lower rays. . . . . . . . . . . . . . . . . . . . . . . Paranthias furcifer p. 172 7b. Head length 30%– 43% of SL; caudal fin usually rounded, truncate, or emarginate (deeply forked only in Anthias and Hemanthias). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8a. Dorsal fin with 8 spines and 13 rays; preoperculum with large antrorse spine on lower edge (under scales and skin); middle opercular spine extending to edge of opercular margin . . . . . . . . . . . . . . . . Gonioplectrus hispanus p. 152 8b. Dorsal fin with 7 to 11 spines and 9 to 20 rays; preoperculum generally without large antrorse spine; middle opercular spine not extending to opercular margin . . . . . . . . . . . 9

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9a. Anterior naris distinctly tubular and located closer to upper lip than to posterior naris; dorsal fin with 7 to 9 spines and 9 to 14 rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9b. Anterior naris variously shaped but always closer to posterior naris than to upper lip; dorsal fin with 9 to 11 spines and 10 to 20 rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 10a. Dorsal fin with 8 spines and 13 or 14 rays, with only shallow notch between spinous and rayed sections . . . . . . . . . . . . . . . . . . . . . . . Bathyanthias cubensis p. 132 10b. Dorsal fin with 7 or 8 spines and 9, 12, or 13 rays, with spinous and rayed sections completely separate . . . . . . . 11 11a. Head and body with five red, reddish brown, or orange stripes. . . . . . . . . . . . . . . . . . . . . Liopropoma rubre p. 164 11b. Head and body with broad, yellow-edged, dark reddish brown, midlateral stripe. . . . . Liopropoma eukrines p. 163 12a. Supramaxilla (thin movable bone) on upper rear margin of maxilla; most jaw teeth depressible; scaly membrane joining upper section of pectoral fin base to side of body; rayed sections of dorsal and anal fins mostly covered with thick skin and scales; scales in horizontal series 69 to 146; dorsal fin with 9 to 11 spines and 13 to 20 rays . . . . . . . . . . . . 13 12b. Supramaxilla (thin movable bone) on upper rear margin of maxilla absent; jaw teeth fixed; no scaly membrane joining upper section of pectoral fin base to side of body; rayed sections of dorsal and anal fins not covered with thick skin and scales; scales in horizontal series 39 to 55; dorsal fin with 10 spines and 10 to 16 rays . . . . . . . . . . . . . . . . . . 33 13a. Corner of preoperculum with large antrorse spine (under skin); snout length equal to or less than eye diameter; dorsal fin with 11 spines and 17 to 19 rays; anal fin with 9 rays . . . . . . . . . . . . . . . . . . . . . . . . Alphestes afer p. 128 13b. Corner of preoperculum without large antrorse spine (enlarged, ventrally directed serrae may be present); snout longer than eye diameter; dorsal fin with 9 to 11 spines and 13 to 20 rays; anal fin with 8 to 13 rays . . . . . . . . . 14 14a. Dorsal fin spines 9; caudal fin rounded or convex . . . . . 15 14b. Dorsal fin spines 10 or 11; caudal fin rounded, truncate, or emarginate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 15a. Head, body, and fins pale gray, brown, or olive green, with orangish brown or reddish spots; four black or white spots below dorsal fin; anal fin with 8 rays; pectoral fin with 16 rays . . . . . . . . . . . . . . . Cephalopholis cruentata p. 137 15b. Head and body with small, dark-edged pale blue spots; two black spots at tip of lower jaw and two black spots near dorsal margin of caudal peduncle; anal fin with 9 rays; pectoral fin with 17 to 19 rays . . . . . . . . . . . . . . . . . . . . . . . . Cephalopholis fulva p. 138

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16a. Body depth greater than head length and 40%– 45% of SL; dorsal fin with 11 spines and 18 to 20 rays . . . . . . . . . . . . . . . . . . . . . . . Dermatolepis inermis p. 139 16b. Body depth usually not greater than head length and 28%– 40% of SL; dorsal fin with 10 or 11 spines and 13 to 19 rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 17a. Anal fin with 10 to 13 rays. . . . . . . . . . . . . . . . . . . . . . . 18 17b. Anal fin with 7 to 9 (rarely 10) rays . . . . . . . . . . . . . . . . 24 18a. Gill rakers on first arch 48 to 55; body depth 31%–39% of SL . . . . . . . . . . . . . . . . Mycteroperca acutirostris p. 165 18b. Gill rakers on first arch 11 to 41; body depth 28%–34% of SL. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 19a. Preopercular margin with notch above serrate lobe at angle (in specimens longer than 30 cm SL) . . . . . . . . . . . . . . . 20 19b. Preopercular margin without notch above serrate lobe at angle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 20a. Median fins of adults without exserted rays; lateral line scales 88 to 96; scales in longitudinal series 128 to 146 . . . . . . . . . . . . . . . . . . . . . Mycteroperca microlepis p. 168 20b. Some but not all fin rays produced beyond fin membranes in large adults only; lateral line scales 70 to 82; scales in longitudinal series 108 to 128 . . . . . . . . . . . . . . . . . . . . 21 21a. Gill rakers on first arch 23 to 27; caudal fin rays equally exserted . . . . . . . . . . . . . Mycteroperca interstitialis p. 167 21b. Gill rakers on first arch 27 to 41; caudal fin rays exserted in adults only and uneven . . . Mycteroperca phenax p. 169 22a. Gill rakers on lower limb of first arch (excluding rudiments) 9 to 11; four to eight pale oblique lines on body; median fin rays exserted in large adults . . . Mycteroperca tigris p. 170 22b. Gill rakers on lower limb of first arch (excluding rudiments) 9 to 16; color pattern not as above (22a); median fin rays not exserted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23a. Distal third of pectoral fin bright yellow and sharply demarcated from remainder of fin in adults; posterior naris about twice size of anterior naris; gill rakers on first arch 24 to 27 . . . . . . . . . . . . . . . Mycteroperca venenosa p. 171 23b. Pectoral fin of adults with narrow orange or white margin shading into remainder of fin; posterior naris about same size as anterior naris; gill rakers on first arch 17 to 24 . . . . . . . . . . . . . . . . . . . . . . . . Mycteroperca bonaci p. 166 24a. Dorsal fin with 13 to 15 rays; anal fin with 9 rays; pelvic fin origin in front of pectoral fin insertion; pelvic fin as long as or longer than pectoral fin (in specimens between 10 and 50 cm SL) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 24b. Dorsal fin with 14 to 18 rays; anal fin with 8 to 10 rays; pelvic fin origin below or behind pectoral fin insertion; pelvic fin not longer than pectoral fin (usually 90% of pectoral fin length). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

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25a. Anal fin with 8 to 10 rays; caudal fin truncate or concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 25b. Anal fin with 8 rays; caudal fin convex or rounded . . . . 27 26a. Dorsal fin membranes distinctly incised between spines, third or fourth spine longest; head and body reddish brown and densely covered with small white spots (spots about size of pupil) . . . . . . . . Epinephelus drummondhayi p. 143 26b. Dorsal fin membrane not incised between spines, second or third spine longest; head and body dark reddish brown and covered with irregular white spots and blotches . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus morio p. 147 27a. Third to eleventh dorsal fin spines about equal in length and distinctly shorter than anterior dorsal fin rays; greatest body width at least half body depth . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus itajara p. 146 27b. Second, third, or fourth dorsal fin spine longest, and longest spine longer than anterior dorsal fin rays; body width usually less than half of body depth . . . . . . . . . . . . . . . . 28 28a. Head and body with dark orangish brown or dark red spots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 28b. Head and body without dark orangish brown or dark red spots . . . . . . . . . . . . . . . . . . . . Epinephelus striatus p. 151 29a. Caudal peduncle with black saddle; base of dorsal fin with three to five dark blotches; rayed section of dorsal fin without blackish margin; pectoral fin with 18 to 20 rays . . . . . . . . . . . . . . . . . . . . Epinephelus adscensionis p. 142 29b. Caudal peduncle without black saddle; base of dorsal fin without dark blotches; rayed section of dorsal fin with blackish margin; pectoral fin with 16 to 18 rays . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus guttatus p. 145 30a. Dorsal fin has 10 spines, with second spine longest and twice the length of tenth spine; juveniles (shorter than 20 cm SL) without black bar or saddle on caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus nigritus p. 149 30b. Dorsal fin has 11 spines, with third or fourth longest but not much longer than tenth spine; juveniles (shorter than 20 cm SL) with black bar or saddle on caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 31a. Body with eight or nine dark bars (last two may be joined into one wide band on caudal peduncle) . . . . . . . . . . . . . . . . . . . . . Epinephelus mystacinus p. 148 31b. Body without dark bars . . . . . . . . . . . . . . . . . . . . . . . . . 32 32a. Membranes between dorsal fin spines not deeply incised, margin of membranes yellow or yellowish green in juveniles and small adults; nares subequal; pearly blue line runs from eye to corner of operculum; black saddle on caudal peduncle does not extend below lateral line . . . . . . . . . . . . . . . . . . . . Epinephelus flavolimbatus p. 144

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32b. Membranes between dorsal fin spines deeply incised, margin of membranes black; posterior naris 3 to 5 times size of anterior naris (except in juveniles less than 15 cm SL); no blue line between eye and operculum; black saddle on caudal peduncle extends below lateral line . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus niveatus p. 150 33a. Preoperculum expanded posteriorly and bearing many large, diverging serrae . . . . . . . . . . . . . . . . . . . . . . . . . . 34 33b. Preoperculum not expanded posteriorly, and serrae only slightly enlarged at angle . . . . . . . . . . . . . . . . . . . . . . . . 35 34a. Preoperculum with two clusters of enlarged serrae or spines; cheek scale rows (from eye to lower corner of preoperculum) 10 to 15; scales in horizontal series 66 to 88 . . . . . . . . . . . . . . . . . . . . . . . Diplectrum formosum p. 141 34b. Preoperculum with one cluster of enlarged serrae or spines; cheek scale rows 7 to 9; scales in horizontal series 54 to 75 . . . . . . . . . . . . . . . . . . . . . . . Diplectrum bivittatum p. 140 35a. Body depth 42%– 48% of SL; dorsal fin with 10 spines and 14 to 16 rays, membranes between dorsal fin spines not incised; lateral line scales 48 to 53; gill rakers on first arch 17 to 23 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 35b. Body depth 27%– 43% of SL; dorsal fin with 10 spines and 10 to 16 rays, membranes between dorsal fin spines distinctly incised; lateral line scales 28 to 64; gill rakers on first arch 9 to 48. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 36a. Body with whitish bars . . . . . . Hypoplectrus puella p. 160 36b. Body without whitish bars . . . . . . . . . . . . . . . . . . . . . . . 37 37a. Head and body pale bluish. . . Hypoplectrus gemma p. 158 37b. Head and body not uniformly blue . . . . . . . . . . . . . . . . 38 38a. Head and body buff . . . . . . . Hypoplectrus unicolor p. 162 38b. Head and body not buff . . . . . . . . . . . . . . . . . . . . . . . . . 39 39a. Head and body tan to brown; caudal fin not yellow . . . . . . . . . . . . . . . . . . . . . . Hypoplectrus nigricans p. 159 39b. Head and body dark brown, blue, or black; caudal fin yellow. . . . . . . . . . . . . . . . . Hypoplectrus chlorurus p. 157 40a. Branchiostegal rays 6; dorsal fin with 10 spines and 10 or 11 rays; lateral line scales 40 to 46; gill rakers on first arch 9 to 13 . . . . . . . . . . . . . . . . . . Serraniculus pumilio p. 182 40b. Branchiostegal rays 7; dorsal fin with 10 spines and 10 to 16 rays; lateral line scales 28 to 64; gill rakers on first arch 14 to 48 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 41a. Lateral line scales 27 to 30; dorsal fin with 10 spines and 15 to 17 rays; lower margin of preoperculum with 1 to 3 antrorse spines; gill rakers on first arch 14 to 18 . . . . . . . . . . . . . . . . . . . . . Plectranthias garrupellus p. 173 41b. Lateral line scales 39 to 64; dorsal fin with 10 spines and 10 to 16 rays; lower margin of preoperculum without antrorse spines; gill rakers on first arch 14 to 44 . . . . . . 42 42a. Branched caudal fin rays 13 to 15; dorsal fin with 10 spines

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and 10 to 14 rays; gill rakers on first arch 14 to 31; precaudal vertebrae 10 and caudal vertebrae 14 . . . . . . . . . 43 42b. Branched caudal fin rays 13; dorsal fin with 10 spines and 13 to 16 rays; gill rakers on first arch 34 to 44; precaudal vertebrae 10 or 11 and caudal vertebrae 15 or 16 . . . . . 56 43a. Caudal fin of adults double emarginate or triple lobed (upper middle and lower middle rays longer than other rays); dorsal fin rays 10 to 12; membranes between dorsal fin spines elongated into filaments at tips of spines; pectoral fin with 16 to 20 rays; lateral line scales 46 to 49. . . . . . 44 43b. Caudal fin of adults truncate or emarginate; dorsal fin rays 10 to 14; membranes between dorsal fin spines not elongated into filaments; pectoral fin with 13 to 18 rays; lateral line scales 39 to 63 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 44a. Adults dark, with light spot on each scale, forming series of stripes on side; dorsal fin spines silvery blue, membranes black with silvery streaks and spots; juveniles mottled with brown blotches and white spots, with dark brown or black stripe from eye to caudal fin base; gill rakers on first arch 21 to 29 . . . . . . . . . . . . . . . . . . Centropristis striata p. 135 44b. Adults and juveniles pale, with seven or fewer distinct dark vertical bars; dorsal and caudal fins pale, with several dark spots and several white spots; gill rakers on first arch 17 to 22 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 45a. Dorsal fin spines with filaments rarely extending past spine tips; no black spot at base of last three dorsal fin spines; distinct blackish brown bars on side; black spots on dorsal and caudal fins; interorbital width 6%–9% of SL . . . . . . . . . . . . . . . . . . . . . . . Centropristis ocyurus p. 133 45b. Dorsal fin spines with filaments often extending past spine tips; large black spot at base of last three dorsal fin spines; indistinct dark brownish bars on side; reddish or golden brown spots on dorsal and caudal fins; interorbital width 4%– 6% of SL . . . . . . . . Centropristis philadelphica p. 134 46a. Middle opercular spine enlarged and curved upward; snout length 12%–14% of SL; caudal fin forked, with upper rays longer than lower rays; lateral line scales 50 to 55 . . . . . . . . . . . . . . . . . . . . . . Serranus luciopercanus p. 187 46b. Middle opercular lobe not curved upward; snout length 7%–12% of SL; caudal fin truncate or emarginate; lateral line scales 39 to 52 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 47a. Top of head scaled posterior to rear end of interorbital region; opercular spines poorly developed; scales weakly ctenoid and often deciduous. . . . . . . . . . . . . . . . . . . . . . 48 47b. Top of head naked; opercular spines well developed; scales strongly ctenoid and not deciduous . . . . . . . . . . . . . . . . 49 48a. Inner surface of operculum with black mark anterior to and just below middle spine; no dark blotch on rayed section of dorsal fin. . . . . . . . . . Serranus atrobranchus p. 184

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126 PERCIFORMES

48b. Inner surface of operculum without black mark; dark blotch on rayed section of dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Serranus notospilus 49a. Dorsal fin rays 12 to 14; pectoral fin rays 15 to 17; gill rakers on first arch 15 to 19; lateral line scales 42 to 46 . . . . . . . . . . . . . . . . . . . . . . . . Serranus subligarius p. 190 49b. Dorsal fin rays 10 to 12 (rarely 13); pectoral fin rays 13 to 16; gill rakers on first arch 14 to 31; lateral line scales 39 to 52 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 50a. Caudal fin with submarginal black stripes converging at base of fin to form V; dorsal fin notched between spinous and rayed section, with fourth and fifth spines longer than tenth spine; pectoral fin rays 15; gill rakers on first arch 21 to 25; lateral line scales 50 to 52 . . . . . . . . . . . . . . . . . . . . . . . . . Serranus tabacarius p. 191 50b. Caudal fin without submarginal black stripes converging at base; dorsal fin not notched between spinous and rayed sections, with fourth and fifth spines not much longer than tenth spine; pectoral fin rays 13 to 16; gill rakers on first arch 14 to 20; lateral line scales 42 to 51 . . . . . . . . . . . . 51 51a. Head and front of body with several irregular blue stripes; upper and lower margins of caudal fin with series of six to seven dark brown to blackish spots; anal fin with brown spots; pectoral fin rays 13 or 14; lateral line scales 45 to 47; caudal peduncle scales 26 to 30 . . . . . . . . . . . . . . . . . . . . . . . . . Serranus chionaraia p. 186 51b. Color not as above (51a); pectoral fin rays 13 to 16; lateral line scales 42 to 51; caudal peduncle scales 23 to 39. . . . 52 52a. Body with six or seven irregular dark brown to black bars and longitudinal rows of small dark blotches; snout longer than eye diameter; gill rakers on first arch 15 to 19; lateral line scales 48 to 51 . . . . . . . . . . . . Serranus tigrinus p. 192 52b. Body without six or seven irregular dark brown bars and longitudinal rows of small dark blotches; snout not longer than eye diameter; gill rakers on first arch 17 to 31; lateral line scales 42 to 51 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 53a. Body with vertical white bar just before anus; pectoral fin rays 15 or 16; lateral line scales 47 to 51 . . . . . . . . . . . . . . . . . . . . . . . . . . . Serranus phoebe p. 189 53b. Body without vertical white bar just before anus; pectoral fin rays 13 to 15; lateral line scales 42 to 51. . . . . . . . . . 54 54a. Body with seven pale blue or greenish bars above lateral line between nape and caudal peduncle; gill rakers on first arch 27 to 31; lateral line scales 46 to 50; cheek scale rows 5 or 6 . . . . . . . . . . . . . . . . . . . Serranus tortugarum p. 193 54b. Body without seven pale blue or greenish bars above lateral line; gill rakers on first arch 14 to 18; lateral line scales 42 to 50; cheek scale rows 6 to 12 . . . . . . . . . . . . . . . . . . . 55

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55a. Lower side of body with four square black blotches, each with red or yellow bar below it; caudal fin base with four dark spots in vertical series; cheek scale rows 6 or 7; lateral line scales 42 to 48; caudal peduncle scales 23 to 26 . . . . . . . . . . . . . . . . . . . . . . . . . . Serranus baldwini p. 185 55b. Lower side of body without square black blotches and red or yellow bars; caudal fin base without dark spots; cheek scale rows 10 to 12; lateral line scales 46 to 50; caudal peduncle scales 31 to 36 . . . . . . . Serranus annularis p. 183 56a. Maxilla and interorbital region naked . . . . . . . . . . . . . . 57 56b. Maxilla and interorbital region scaly . . . . . . . . . . . . . . . 59 57a. Lateral line scales 54 to 62 . . . . . Hemanthias leptus p. 154 57b. Lateral line scales 42 to 53 . . . . . . . . . . . . . . . . . . . . . . . 58 58a. Adults with long filaments at tips of some dorsal fin spines; dorsal fin rays 13 or 14; pectoral fin rays 16 to 21 . . . . . . . . . . . . . . . . . . . . . . . . Hemanthias vivanus p. 155 58b. Adults with no filaments at tips of dorsal fin spines; dorsal fin rays 14 to 16; pectoral fin rays 15 to 19 . . . . . . . . . . . . . . . . . . . . . Hemanthias aureorubens p. 153 59a. Vomerine tooth patch elongated; tongue with oval tooth patch; dorsal fin rays 13 to 16; pectoral fin rays 16 to 18; lateral line scales 35 to 41 . . . . . . . . . . . . . . . . Pronotogrammus martinicensis p. 174 59b. Vomerine tooth patch triangular or V-shaped; tongue usually without teeth; dorsal fin rays 14 to 16; pectoral fin rays 16 to 21; lateral line scales 31 to 57 . . . . . . . . . . . . 60 60a. Lateral line interrupted below dorsal fin insertion; lateral line scales 51 to 57; anterior naris with long filament extending beyond posterior naris; body depth 27%–32% of SL; pectoral fin rays 19 to 21; gill rakers on first arch 34 to 39 . . . . . . . . . . . . . . . . . . . . . . Anthias tenuis p. 130 60b. Lateral line continuous; lateral line scales 31 to 48; anterior naris without long filament; body depth 34%– 43% of SL; pectoral fin rays 16 to 21; gill rakers on first arch 38 to 48 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 61a. Lateral line scales 42 to 48; caudal fin lobes produced, with upper lobe about 50% of SL . . . . . . Anthias woodsi p. 131 61b. Lateral line scales 31 to 34; caudal fin forked and lobes rounded in specimens longer than 12 cm SL and not produced . . . . . . . . . . . . . . . . . . . . Anthias nicholsi p. 129

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128 PERCIFORMES

Alphestes afer (Bloch, 1793) Mutton hamlet

Alphestes afer has a large eye, a short snout, and cryptic coloration. Profile of head is nearly straight. Maxilla extends beyond posterior margin of orbit, with ventral margin smoothly curved and lacking step or bony knob. Supramaxilla is well developed. Jaw teeth are small, slender, and arranged in three to five rows. A pair of small canine teeth is located at front of upper and lower jaws. Vomerine and palatine teeth are similar to jaw teeth but smaller. Preoperculum is rounded, with posterior margin serrated and bearing a large, anteroventrally directed spine at corner. Gill rakers on first arch number 22 to 25, with 6 to 8 on upper limb and 16 or 17 on lower limb. Measurements are expressed as percent of SL: head length 39%– 42%, caudal peduncle depth 12%–14%, body depth 32%– 42%; and as percent of head length: eye diameter 19%–24%. Pectoral fin is broadly rounded and symmetrically shaped and has 16 or 17 rays. Dorsal fin has 11 spines and 17 to 19 rays. Anal fin has 9 rays. Caudal fin is rounded. Scales are cycloid and cover body and rayed sections of dorsal and anal fins.

Lateral line scales number 55 to 61, and scales in horizontal series range from 68 to 77. Color is olivaceous to light brown, with irregular blotches and bars of dark brown and orange spots. Body and fins are covered with small white spots, and pectoral fin is orange to yellow, with faint brown reticulations. This species occurs to a depth of 35 m in the western Atlantic from southern Florida and Bermuda to southern Brazil, including the eastern Gulf of Mexico off southwestern Florida, the Bahamas, and the Antilles. During the day it hides in crevices or in vegetation, and during the night it is an active predator. Food consists of crustaceans. Fecundity ranges from 157,512 to 223,706 eggs. Maximum known size is 330 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Smith 1971 (in part), 1997; Robins and Ray 1986 (as Epinephelus afer); Cervigón 1991 (as Epinephelus [Alphestes] afer), 1993; Heemstra and Randall 1993; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 129

Anthias nicholsi Yellowfin bass

Firth, 1933

Anthias nicholsi has most of head scaled, an uninterrupted lateral line, and a deeply forked caudal fin. Anterior and posterior nares are close set, with posterior margin of anterior naris produced into short flap. Eye length is greater than interorbital width. Upper jaw extends to center of pupil. Premaxilla is slightly protractile. Supramaxilla is absent. Jaw teeth are arranged into outer conical row and inner villiform or cardiform band. Teeth are present in palatine, and vomerine tooth patch is chevron to triangular shaped and lacks posterior extension. Teeth are usually absent on tongue. Preoperculum has serrated posterior margin. Gill rakers on first arch number 39 to 44, with 12 or 13 on upper limb and 27 to 31 on lower limb. Measurements are expressed as percent of SL: head length 34%–39%, snout length 6%– 8%, orbit diameter 9%–13%, upper jaw length 17%–19%, pectoral fin length 30%– 32%, body depth 34%– 44%. Pectoral fin has 18 to 21 rays. Dorsal fin is continuous and not incised and has 10 spines and 15 (occasionally 14) rays, with long filament trailing third spine and short filaments trailing other spines. Anal fin has 6 to 8 rays. Caudal fin is deeply forked, lobes are rounded, and upper lobe is slightly longer than lower lobe. Scales are ctenoid and occur on most of head, including maxilla.

Scales encircling caudal peduncle number 17 or 18. Lateral line is continuous, and tubed scales number 31 to 34. Color is reddish lavender dorsally and silvery white ventrally, with three or four bright yellow stripes on side and yellow patches on fins and on body. Head has two additional stripes; one runs from below eye to pectoral fin base, and the other runs through eye to opercular margin. Dorsal and anal fins are bright yellow. Pelvic fin is yellow with pink outer margin. Caudal fin is whitish with red margin. This species occurs between 55 and 430 m in the western Atlantic from Nova Scotia to northeastern Brazil, including the Gulf of Mexico and the Caribbean Sea. It occurs in the northeastern Gulf of Mexico. Food consists of zooplankton such as pteropods and copepods. This species may be a protogynous hermaphrodite, with females ranging from 71 to 139 mm SL and males ranging from 106 to 149 mm SL. Spawning takes place in the spring in the eastern Gulf of Mexico. Maximum known size is 200 mm SL. references: Anderson and Heemstra 1980; Williams and Shipp 1980; Robins and Ray 1986; Bullock and Smith 1991; Smith 1997; Anderson 2002b.

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130 PERCIFORMES

Anthias tenuis Nichols, 1920 Threadnose bass

Anthias tenuis has most of head scaled, an interrupted lateral line, and a deeply forked caudal fin. Anterior and posterior nares are relatively far apart, with posterior margin of anterior naris developed into a long, slender filament that usually extends to orbit. Upper jaw extends to about center of pupil. Premaxilla is slightly protractile. Supramaxilla is absent. Eye length is greater than interorbital width. Jaw teeth are arranged into outer conical row and inner villiform or cardiform band. Villiform teeth are present on palatine, and vomerine tooth patch lacks posterior extension. Teeth are absent on tongue. Preoperculum has serrated posterior margin. Gill rakers on first arch number 34 to 39, with 9 to 11 on upper limb and 24 to 28 on lower limb. Measurements are expressed as percent of SL: head length 28%–32%, snout length 5%– 6%, orbit diameter 8%–11%, upper jaw length 13%– 15%, pectoral fin length 27%–31%, body depth 27%–32%. Pectoral fin has 19 to 21 rays. Dorsal fin is continuous and notched between spinous and rayed sections and has 10 spines and 15 (rarely 14) rays, with short filaments trailing spines. Anal fin has 7 to

9 rays. Caudal fin lobes are pointed, and upper is slightly longer than lower. Scales are ctenoid and occur on most of head, including maxilla. Scales encircling caudal peduncle number 25 to 28 (usually 27). Lateral line is interrupted for several rows below posterior end of dorsal fin, and pored scales number 51 to 57. Color is red orange to rosy dorsally, pallid anteroventrally, and purplish pink posteroventrally. Iris is yellow. Dorsal and anal fin membranes have oblong to elliptical greenish spots, pectoral and pelvic fins are pallid, and posterior sections of upper and lower caudal fin lobes are dark red. This species occurs between 55 and 915 m in the western Atlantic from North Carolina and Bermuda to Venezuela, including the Gulf of Mexico. In the Gulf of Mexico it has been captured off western Florida and northern Yucatán. Maximum known size is 90 mm SL. references: Beebe and Tee-Van 1933; Anderson and Heemstra 1980; Bullock and Godcharles 1982; Bullock and Smith 1991; Smith 1997; Smith-Vaniz et al. 1999; Anderson 2002b.

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SERRANIDAE 131

Anthias woodsi

Anderson and Heemstra, 1980

Anthias woodsi has most of head scaled, a continuous lateral line, and a deeply forked caudal fin. Anterior and posterior nares are close set, with posterior margin of anterior naris produced into short flap. Upper jaw extends to center of pupil. Premaxilla is slightly protractile. Supramaxilla is absent. Eye length is greater than interorbital width. Upper jaw has outer row of small conical teeth and inner band of villiform teeth, with two or three pairs of enlarged canines on either side of symphysis. Lower jaw has a row of conical teeth, with several on either side of symphysis enlarged and directed posteriorly or curved posteriorly. Vomerine and palatine teeth are granular or villiform. Vomerine tooth patch is chevron shaped, without posterior prolongation. Teeth are absent on tongue. Preoperculum has smooth posterior margin, small spines at corner, and fine serrations on ventral margin. Gill rakers on first arch are long and slender and number 38 to 40, with 11 or 12 on upper limb and 26 to 28 on lower limb. Measurements are expressed as percent of SL: head length 36%– 40%, snout length 8%–11%, orbit diameter 11%–14%, upper jaw length 17%–19%, pectoral fin length 31%–35%, pelvic fin length 27%–31%, body depth 34%– 41%. Pectoral fin has 16 to 18 rays. Dorsal fin is continuous and unnotched and has 10 spines and 14 or 15

(usually 14) rays, with fourth spine longest and fin membranes produced into filaments behind spines. Pelvic fin is moderately to greatly produced. Anal fin has 7 or 8 (usually 7) rays. Caudal fin is deeply forked, and lobes are greatly produced. Scales are ctenoid and occur on body, head (including maxilla), and bases of rayed section of dorsal and anal fins. Scales encircling caudal peduncle number 21 to 24. Tubed lateral line scales number 42 to 48. Cheek has 9 or 10 scale rows. Color is reddish, with yellow bar posterior to orbit and broad yellow or vermilion band running from operculum to caudal peduncle, but narrowing below insertion of dorsal fin. Iris of eye is reddish orange. Pectoral fin is pale pink to bright red. Dorsal fin is largely yellow. Caudal fin is red, with lateralmost fin rays and proximal sections of other rays yellow, and distal margin yellow or yellowish green. Body is straw colored in alcohol. This species occurs between 175 and 475 m in the western Atlantic from North Carolina to the Straits of Florida, including the eastern Gulf of Mexico. In the Gulf of Mexico it is known from the Dry Tortugas. Maximum known size is 250 mm SL. references: Anderson and Heemstra 1980; Anderson 2002b.

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132 PERCIFORMES

Bathyanthias cubensis Yellowtail bass

(Schultz, 1958)

Bathyanthias cubensis has scales on head, an unnotched dorsal fin, and scales covering basal section of dorsal, anal, pectoral, and caudal fins. Head is moderately compressed. Nares are widely separated, with anterior naris tubular and posterior naris near anterodorsal margin of orbit. Upper jaw extends to about middle of pupil. Eye diameter is about equal to interorbital width. Jaw teeth and vomerine and palatine teeth are villiform. Posterior margin of interoperculum is finely serrate. Gill rakers on first arch are slender and number 18 to 23. Measurements are expressed as percent of SL: head length 33%–38%, snout length 7%–9%, eye diameter 8%–9%, interorbital width 4%– 5%, body depth 25%–30%. Pectoral fin is long and pointed and has 14 or 15 rays. Dorsal fin has eight spines and 14 or 15 rays, with first spine very short. Anal fin has 8 rays. Caudal fin is truncate or slightly emarginate. Scales are ctenoid and extend anterior to eyes on head. Lateral line is arched, and pored scales number 45 to 47. Color is pale red on body

and head, darker dorsally than ventrally, with yellow spots. Narrow yellow stripe extends from tip of snout across cheek, and from posterior margin of eye to posterior margin of operculum. Dorsal and caudal fins are yellow, with upper and lower margins of caudal fin rays black tipped. Anal fin is pale yellow to reddish. This species occurs between 28 and 377 m in the western Atlantic from the east coast of Florida to Venezuela, including the Gulf of Mexico and Cuba. It occurs throughout the Gulf of Mexico. Individuals are sequential hermaphrodites; they first mature as females and later function as males. Maximum known size is 150 mm SL.

references: Longley and Hildebrand 1941 (as Liopropoma aberrans); Schultz 1958b; Chittenden and Moore 1977; Hoese and Moore 1977, 1998; Matsuura 1983a; Darcy and Gutherz 1984; Robins and Ray 1986; Bullock and Smith 1991; Cervigón 1991 (all above as Pikea mexicana); Boschung 1992 (as Bathyanthias mexicanus); Heemstra 2002c.

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SERRANIDAE 133

Centropristis ocyurus (Jordan and Evermann, 1887) Bank seabass

Centropristis ocyurus has short dermal flaps on dorsal fin spines and a large dark blotch on fourth vertical bar on body. Anterior and posterior nares are close set, with posterior margin of anterior naris developed into large flap. Upper jaw extends to about center of eye. Eye diameter is greater than interorbital width. Jaw teeth are in wide bands, with inner and outer teeth slightly larger than median teeth. Vomerine teeth are arranged in a broad, triangular patch. Teeth on palatine are in narrow band. Tongue lacks teeth. Posterior margin of preoperculum is finely serrate, and angle and ventral margin are coarsely serrate. Gill rakers on first arch are long and slender and number 17 to 22 (usually 19 to 21). Measurements are expressed as percent of SL: head length 37%– 44%, snout length 7%–13%, orbital length 8%–12%, interorbital width 6%–9%, body depth 28%–38%. Pectoral fin rays number 16 to 18. Dorsal fin has 10 spines and 10 to 12 rays. Anal fin has 7 rays. Caudal fin is rounded in juveniles and trilobed or double concave in adults. Scales are ctenoid, scales between dorsal fin origin and lateral line number 10 to 14, and scales between anal fin origin and lateral line number 14 to 19. Lateral line scales number 46 to 48. Top of head, suborbital region,

maxilla, lower jaw, and snout are naked. Vertebrae number 24. Color is pale cream to yellowish white, with six or seven dark bars from dorsal fin base to flank. Each bar has dark rectangular blotch at or just below lateral line, and the one below front part of dorsal fin is darker than the others. Dorsal and caudal fins are covered with black spots. This species occurs between 11 and 100 m in the western Atlantic from Cape Hatteras to Quintana Roo, including the Gulf of Mexico. In the western Gulf of Mexico it is not known from south of Tampico. Adults are most common at depths greater than 30 to 35 m on sandy-shell substrates near reefs. It is a protogynous hermaphrodite, with mature females ranging from 80 to 158 mm SL. Food consists mainly of crustaceans and, to a lesser extent, mollusks, polychaetes, echinoderms, and ray-finned fishes. Maximum known size is 300 mm SL.

references: Miller 1959; Smith et al. 1975; Hastings et al. 1976; Bortone 1977a; Hoese and Moore 1977, 1998; Hastings 1979; Darcy and Gutherz 1984; Robins and Ray 1986; Bullock and Smith 1991; Boschung 1992; Castro-Aguirre et al. 1999; Heemstra 2002c.

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134 PERCIFORMES

Centropristis philadelphica (Linnaeus, 1758) Rock seabass

Centropristis philadelphica has long fleshy filaments on dorsal fin spines and a dark blotch at base of last three dorsal fin spines. Anterior and posterior nares are close set, with posterior margin of anterior naris developed into large flap. Upper jaw extends to center of eye. Eye length is greater than interorbital width. Jaw teeth are in wide bands, with outer and inner teeth slightly to moderately larger than median teeth. Vomerine teeth are arranged in narrow chevron-shaped patch. Teeth on palatine are in narrow band. Tongue lacks teeth. Posterior margin of preoperculum is finely serrated, and angle and ventral margin are coarsely serrated. Gill rakers on first arch are long and slender and number 17 to 22. Measurements are expressed as percent of SL: head length 35%– 43%, snout length 7%–11%, orbital length 7%–13%, interorbital width 4%– 6%, body depth 25%–33%. Pectoral fin rays number 15 to 20 (usually 18). Dorsal fin has 10 spines and 11 rays. Anal fin has 7 rays. Caudal fin is rounded in juveniles and trilobed or double concave in adults. Scales are ctenoid, scales between dorsal fin origin and lateral line number 11 to 18, and scales between anal fin origin

and lateral line number 14 to 21. Lateral line scales number 46 to 49. Vertebrae number 22 to 24 (usually 24). Top of head, suborbital region, maxilla, lower jaw, and snout are naked. Color is olive gray to brown dorsally, with rusty spots that are most dense on head, and pale to whitish ventrally. Seven obscure brown bars run obliquely forward on upper part of body. Dorsal and caudal fins are covered with reddish brown to golden brown spots. This species occurs between 10 and 172 m in the western Atlantic from Virginia to the southern Gulf of Mexico. It is most common on soft mud and sandy bottoms. This species is a protogynous hermaphrodite; females mature in their first year of life and change to males in the second year. Maximum life span is three or four years. Food consists of mollusks, crustaceans, and ray-finned fishes. Maximum known size is 209 mm SL. references: Longley and Hildebrand 1941; Gunter 1945; Hildebrand 1954; Miller 1959; Bortone 1977a; Hoese and Moore 1977, 1998; Robins and Ray 1986; Bullock and Smith 1991; Boschung 1992; Schaldach et al. 1997; Heemstra 2002c.

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SERRANIDAE 135

Centropristis striata Black seabass

(Linnaeus, 1758)

Centropristis striata has very short dermal flaps on dorsal fin spines and a darkly colored body. Anterior and posterior nares are close set, with posterior margin of anterior naris developed into large flap. Upper jaw extends to center of eye. Eye diameter is greater than interorbital width. Jaw teeth are in wide bands, with inner and outer teeth slightly enlarged. Vomerine teeth are arranged in wedge-shaped patch. Teeth on palatine are arranged in long, narrow patch. Tongue lacks teeth. Posterior margin of preoperculum is finely serrate, and angle and ventral margin are coarsely serrate. Gill rakers on first arch are long and slender and number 21 to 29 (usually 22 or 23). Measurements are expressed as percent of SL: head length 39%– 44%, snout length 8%–14%, orbital length 6%–12%, interorbital width 5%–9%, body depth 31%–38%. Pectoral fin rays number 14 to 20 (usually 16 to 19). Dorsal fin has 10 spines and 10 to 12 rays. Anal fin has 7 rays. Caudal fin is rounded in juveniles and trilobed or double concave, with uppermost ray often greatly elongated in adults. Scales are ctenoid, scales between dorsal fin origin and lateral line number 10 to 15, and scales between anal fin origin and lateral line number 12 to 16. Lateral line scales number 46 to 49. Vertebrae number

24. Top of head, suborbital region, maxilla, lower jaw, and snout are naked. Color is dark to nearly black, and dorsal fin has obscure bars, white stripes, or horizontal rows of stripes. Body often has narrow pale stripes formed by pale spots on center of each scale. Upper and lower margins of caudal fin are white. Juveniles are pale gray, with broad dark stripe along side, large black spot near end of spinous section of dorsal fin, and turquoise streaks below eye. This species occurs between the shoreline and 75 m in the western Atlantic from Maine to the Florida Keys and the eastern Gulf of Mexico. In the Gulf of Mexico it is known only from the west coast of Florida, from Pensacola to Placida. Some researchers consider specimens in the Gulf of Mexico to be a separate subspecies, C. striata melanus. This species is associated with hard bottoms. Food consists of mollusks, caridean and penaeid shrimps, crabs, and ray-finned fishes. It is a protogynous hermaphrodite; females mature between one and four years at lengths up to 190 mm SL, and change to males at three to seven years at lengths of 230 mm SL. In the eastern Gulf of Mexico spawning takes place between fall and spring over the inner continental shelf. Maxi-

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136 PERCIFORMES

mum known age is about 20 years, and maximum known size is 600 mm TL. references: Hildebrand and Schroeder 1928; Ginsburg 1952a (as Centropristis melanus); Miller

1959; Springer and Woodburn 1960 (as Centropristis melanus); Bortone 1977a; Hoese and Moore 1977, 1998; Darcy and Gutherz 1984; Robins and Ray 1986; Wenner et al. 1986; Sedberry 1988; Bullock and Smith 1991; Boschung 1992; Heemstra 2002c.

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SERRANIDAE 137

Cephalopholis cruentata Graysby

(Lacepède, 1802)

Cephalopholis cruentata has a moderately long snout; dorsal fin membranes distinctly incised; and head, body, and fins covered with reddish spots. Interorbital region is flat. Upper jaw extends beyond posterior margin of orbit and bears bony knob on posteroventral corner. Supramaxilla is well developed. Teeth in front of jaws are small canines. Preoperculum has rounded, finely serrate margin, with shallow notch above angle. Upper edge of operculum is distinctly convex. Gill rakers on first arch number 18 to 21. Measurements are expressed as percent of SL: head length 38%– 42%, body depth 34%– 40%. Pectoral fin is broadly rounded and symmetrically shaped and has 16 rays. Dorsal fin has nine spines, with fourth or fifth spine longest, and 13 to 15 rays. Anal fin has 8 rays. Caudal fin is rounded. Scales on lateral body are ctenoid. Lateral line scales number 47 to 51, and scales in horizontal series number 69 to 81. Color of head, body, and fins is pale gray, brown, or olive green, with dense covering of orangish brown or reddish spots. Four black or white spots are located in arch at base of dorsal fin, and mid-dorsal white

stripe is occasionally present between tip of lower jaw and nape. This species occurs between 27 and 170 m in the western Atlantic from North Carolina and Bermuda to northern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is common in turtle-grass (Thalassia sp.) beds and on coral reefs. Food of adults consists of ray-finned fishes, and feeding takes place at dawn and dusk. During the day it is secretive. Females change sex between 200 and 230 mm TL at ages four or five. Maximum known size is 330 mm TL. references: Longley and Hildebrand 1941 (as Petrometopon cruentatus); Böhlke and Chaplin 1968 (as Petrometopon cruentatum); Smith 1971 (as Epinephelus cruentatus); Bright and Cashman 1974 (as E. [Cephalopholis] cruentatus); Smith et al. 1975 (as E. cruentatus); Hoese and Moore 1977, 1998 (as E. cruentatus); Matsuura 1983a; Robins and Ray 1986 (as E. cruentatus); Cervigón 1991 (as E. [Cephalopholis] cruentatus); Boschung 1992 (as E. cruentatus); Heemstra and Randall 1993; Smith 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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138 PERCIFORMES

Cephalopholis fulva (Linnaeus, 1758) Coney

Cephalopholis fulva has a moderately long snout, distinctly incised dorsal fin membranes, two small black spots at tip of lower jaw, and two small spots on dorsal surface of caudal peduncle. Interorbital region is flat. Upper jaw extends beyond posterior margin of orbit and bears bony knob on posteroventral margin. Supramaxilla is well developed. Anterior jaw teeth are small canines. Preoperculum has rounded margin, with shallow notch above angle, serrae along upper margin, and moderately enlarged serrae at corner. Upper edge of operculum is distinctly convex. Gill rakers on first arch number 23 to 27, with 7 to 9 on upper limb and 16 to 18 on lower limb. Measurements are expressed as percent of SL: head length 40%– 43%, body depth 34%–38%. Pectoral fin is broadly rounded and symmetrically shaped and has 17 to 19 rays. Dorsal fin has nine spines and 14 to 16 rays. Anal fin has 9 rays. Caudal fin has convex posterior margin and angular corners. Scales on lateral body are ctenoid. Lateral line scales number 46 to 54, and scales in horizontal series number 90 to 97. Color varies with habitat occupied, but all specimens have two large black spots at tip of lower jaw and two black spots on dorsal margin of caudal peduncle. Specimens from deep water are red, those from shallow water are either uniformly orangish brown or orangish brown dorsally and pale tan laterally and ven-

trally, and those ranging from shallow to deep water are yellow. Red or orangish brown specimens are densely covered with small, darkedged pale blue spots, but yellow specimens are sparsely covered with small, dark-edged blue spots on head and anterior part of body. This species occurs to a depth of 45 m in the western Atlantic from South Carolina and Bermuda to southern Brazil, including the Gulf of Mexico and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico and is associated with clear water and frequently occurs on coral reefs. All individuals are protogynous hermaphrodites; females mature at 160 mm SL and transform into males at about 200 mm SL. Food consists of crustaceans and small ray-finned fishes. Eggs are 0.95 mm in diameter, and fecundity ranges from 150,000 to 282,000 eggs per female. Maximum known size is 390 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Smith 1971 (as Epinephelus fulvus); Smith et al. 1975 (as E. fulvus); Hoese and Moore 1977 (as E. fulvus); Matsuura 1983a; Cordova 1986 (as E. fulvus); Robins and Ray 1986 (as E. fulvus); Dennis and Bright 1988a (as E. fulvus); Cervigón 1991 (as E. [Cephalopholis] fulvus); Boschung 1992 (as E. fulvus); Heemstra and Randall 1993; Smith 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 139

Dermatolepis inermis Marbled grouper

(Valenciennes, 1833)

Dermatolepis inermis has inconspicuous opercular spines, a long anal fin, and weakly ctenoid or cycloid scales. Profile of head is steep and nearly straight. Anterior and posterior nares are close set, and posterior naris is 2 or 3 times size of anterior naris. Maxilla extends to about center of eye, with ventral margin smooth and lacking step or bony knob. Supramaxilla is well developed. Jaw teeth are small. Preoperculum is subangular, posterior margin bears fine serrae covered with skin, and serrae at angle are enlarged. Lower opercular spine is rudimentary, and upper edge of operculum is convex. Gill rakers on first arch number 19 to 22. Measurements are expressed as percent of SL: head length 36%– 40%, pectoral fin length 29%– 35%, body depth 40%– 45%. Pectoral fin is broadly rounded and symmetrically shaped and has 18 or 19 rays. Dorsal fin has 11 spines and 18 to 20 rays, and fin membrane is slightly incised between spines. Anal fin has 8 to 10 rays. Caudal fin is rounded in juveniles but truncate to concave in adults. Scales are deeply

embedded. Color of adults is mottled grayish brown, with white speckling and small black spots loosely arranged into rings. Color of juveniles is black to dark brown, with irregular white blotches on head, body, and fins. This species occurs between 21 and 250 m in the western Atlantic from North Carolina to Rio de Janeiro, Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico, although it is rare there, as it is throughout its range. Maximum known size is 900 mm TL. references: Randall 1968a (as Epinephelus inermis), 1996; Smith 1971 (as E. inermis); Bright and Cashman 1974 (as E. [Dermatolepis] inermis); Smith 1997 (as E. inermis); Matsuura 1983a; Robins and Ray 1986 (as E. inermis); Dennis and Bright 1988a (as E.[Dermatolepis] inermis); Cervigón 1991 (as E. [Dermatolepis] inermis); Boschung 1992 (as E. inermis); Heemstra and Randall 1993; Schaldach et al. 1997 (as E. inermis); Heemstra 2002c.

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140 PERCIFORMES

Diplectrum bivittatum Dwarf seabass

(Valenciennes, 1828)

Diplectrum bivittatum has preoperculum bearing numerous strong spines diverging from a single cluster. Posterior margin of anterior naris is developed as large flap. Upper jaw extends to center of eye. Jaw teeth are villiform except for outer row of 5 to 10 canine teeth. Villiform teeth in vomer are in wedge- or Vshaped patch and in palatine are in narrow band. Preoperculum forms right angle between upper and lower limbs, and posterior margin is serrated. Gill rakers on first arch number 20 to 25. Measurements are expressed as percent of SL: head length 32%–37%, snout length 8%– 11%, orbital length 8%–10%, interorbital width 6%– 8%, body depth 24%–30%. Pectoral fin has 14 to 16 rays. Dorsal fin is slightly indented between spinous and rayed sections and has 10 spines and 12 rays. Anal fin has 6 to 8 rays. Caudal fin is truncate to slightly forked, with upper lobe occasionally extended as long filament. Scales are ctenoid, with seven to nine rows on cheek and up to 9 scales between dorsal fin origin and lateral line. Lateral line scales number 54 to 75. Color is brown with copper tinge on dorsal surface, tan on side, and white ventrally. Adults have triangular black spot on upper section of operculum, two broad dark stripes intersected by short, dark vertical bars

on upper flank, and distinct dark spot on caudal fin. Juveniles have two broad, dark lateral stripes extending from operculum to base of caudal fin and from head to dorsal aspect of caudal peduncle. This species occurs from near shore to 120 m in the western Atlantic from southeastern Florida to northern Brazil, including the Gulf of Mexico and the Florida Keys. In the Gulf of Mexico it occurs from Pensacola, FL, westward and southward to Campeche Bay. It does not occur in the Bahamas or the Antilles. It apparently is most common on muddy silt bottoms. Food consists of polychaetes, amphipods, natanian shrimps, brachyuran crabs, and ray-finned fishes. This species is a synchronous hermaphrodite. Maximum known size is 250 mm SL.

references: Longley and Hildebrand 1941; Hildebrand 1955; Stark 1968; Bortone 1977b; Chittenden and Moore 1977; Hoese and Moore 1977, 1998; Ogren and Brusher 1977; Bortone et al. 1981; Matsuura 1983a (as D. radiale); Darcy and Gutherz 1984; Robins and Ray 1986; Bullock and Smith 1991; Cervigón 1991; Boschung 1992; Randall 1996; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Heemstra 2002c.

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SERRANIDAE 141

Diplectrum formosum Sand perch

(Linnaeus, 1766)

Diplectrum formosum has spines on preoperculum radiating from two clusters. Posterior margin of anterior naris is developed as large flap. Maxilla extends to center of eye. Jaw teeth are villiform except for row of 5 to 10 canine teeth. Villiform teeth on vomer are in wedge- or V-shaped patch and on palatine are in narrow band. Preoperculum forms right angle between upper and lower limbs, and posterior margin is serrated. Gill rakers on first arch number 17 to 24. Measurements are expressed as percent of SL: head length 33%–39%, snout length 9%–13%, orbital length 6%–9%, interorbital width 6%– 8%, body depth 25%– 32%. Pectoral fin has 15 to 17 rays. Dorsal fin is slightly indented between spinous and rayed sections and has 10 spines and 11 to 13 rays. Anal fin has 6 to 8 rays. Caudal fin is truncate to moderately forked, with upper lobe slightly longer than lower lobe and frequently extended as filament. Scales are ctenoid, with 10 to 15 rows on cheeks and 8 to 12 scales between dorsal fin origin and lateral line. Pored lateral line scales number 46 to 55, and scales in horizontal series number 66 to 88. Color is light brown on dorsal surface, tan on flanks, and white on belly. Body and dorsal fins are patterned with five to seven dark brown bars alternating with narrow blue and orange stripes.

Dark blotch is present on caudal peduncle. Head is patterned with many wavy blue lines. Juveniles have two dark brown stripes. This species occurs from near shore to 73 m in the western Atlantic from Chincoteague, VA, and Bermuda to São Paulo, Brazil, including the Gulf of Mexico, the Florida Keys, the Bahamas, Cuba, and the Virgin Islands. It is rare in the Antilles and the Bahamas, and in the Gulf of Mexico it is most abundant off the coasts of Florida and Campeche. It is most commonly associated with sandy to shelly bottoms or limestone and coral substrates. Food consists of bivalves; polychaetes; mysid, penaeid, and sergestid shrimps; crabs; chaetognaths; cephalochordates; and ray-finned fishes. Crustaceans are the dominant prey. This species is a synchronous hermaphrodite. Maximum known size is 30 cm SL. references: Longley and Hildebrand 1941; Joseph and Yerger 1956; Springer and Woodburn 1960; Tabb and Manning 1961; Böhlke and Chaplin 1968; Stark 1968; Bortone 1977b; Hoese and Moore 1977, 1998; Bortone et al. 1981; Matsuura 1983a; Darcy and Gutherz 1984; Robins and Ray 1986; Bullock and Smith 1991; Cervigón 1991; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Heemstra 2002c.

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142 PERCIFORMES

Epinephelus adscensionis Rock hind

(Osbeck, 1765)

Epinephelus adscensionis has dorsal fin with fourth or fifth spine longest, ctenoid scales, and three to five dark blotches on body at base of dorsal fin. Maxilla extends beyond posterior margin of orbit, with ventral margin smoothly curved and lacking step or bony knob. Supramaxilla is well developed. Teeth in front of jaws are small canines. Preoperculum is angular, with posterior margin evenly serrate. Dorsal margin of operculum is convex. Gill rakers number 23 to 28, with 7 to 9 on upper limb and 16 to 19 on lower limb. Measurements are expressed as percent of SL: head length 40%– 48%, body depth 29%–36%. Pectoral fin is broadly rounded and has 18 to 20 rays. Dorsal fin has 11 spines and 16 to 18 rays, with membrane between spines distinctly incised. Anal fin has 8 rays. Caudal fin has convex posterior margin. Scales have strong ctenii, and numerous auxillary scales are present. Lateral line scales number 48 to 53, and scales in horizontal series number 82 to 95. Color is buff to pale green, with reddish brown spots and pale blotches scattered over head, body, and fins. Most specimens have three to five dark blotches

at base of dorsal fin, a single dark blotch on dorsal side of caudal peduncle, and dark pigment along posterior margin of caudal fin. This species occurs in the tropical and warm temperate Atlantic. In the western Atlantic it occurs between 2 and 100 m, from Massachusetts and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico and is associated with rocky substrates and coral reefs. Food consists of crustaceans, ray-finned fishes, and young sea turtles. Maximum known size is 60 cm TL in the western Atlantic but reaches about 100 cm at Ascension Island.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Smith 1971, 1997; Smith et al. 1975; CastroAguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Cervigón 1991; Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 143

Epinephelus drummondhayi Goode and Bean, 1878 Speckled hind Epinephelus drummondhayi has pelvic fin origin behind pectoral fin origin and pectoral fin longer than pelvic fin. Profile of head is slightly convex to straight. Maxilla extends beyond middle of eye, with ventral margin smoothly curved and lacking step or bony knob. Supramaxilla is well developed. Teeth in front of jaws are small canines. Preoperculum is angular, with posterior margin evenly serrated. Dorsal margin of operculum is convex. Gill rakers on first arch number 26 to 28, with 9 or 10 on upper limb and 17 or 18 on lower limb. Measurements are expressed as percent of SL: head length 39%– 42%, body depth 38%– 42%. Pectoral fin is broadly rounded and has 18 rays. Dorsal fin has 11 spines and 15 or 16 rays, with membrane incised between anterior spines. Anal fin has 9 rays. Caudal fin is truncate to slightly emarginate. Scales have strong ctenii, and numerous auxillary scales are present. Lateral line scales number 72 to 76, and scales in horizontal series number about 125. Color of adults (greater than 300 mm SL)

is dark reddish brown, with dense covering of small pearly white spots. Distal section of pectoral fin is yellowish. Color of juveniles is bright yellow with numerous small bluish white spots. This species occurs from 25 to 183 m in the western Atlantic from North Carolina and Bermuda to Quintana Roo, including the Florida Keys and the entire Gulf of Mexico. It is associated with rocky bottoms. Food consists of crustaceans and ray-finned fishes. Spawning takes place between July and September. Fecundity ranges up to 2,000,000 eggs per female. Females mature between four and five years of age at 45 to 60 cm SL. Maximum age is about 15 years, and maximum known length is 110 cm TL. references: Smith 1971, 1997; Smith et al. 1975; Robins and Ray 1986; Bullock and Smith 1991; Cervigón 1991; Boschung 1992; Heemstra and Randall 1993; Smith-Vaniz et al. 1999; Heemstra 2002c.

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144 PERCIFORMES

Epinephelus flavolimbatus Yellowedge grouper

Poey, 1865

Epinephelus flavolimbatus has dorsal fin membranes only slightly incised between spines, and pelvic fin origin is anterior to lower end of pectoral fin base. Snout length is about equal to eye diameter in large specimens but shorter than eye diameter in specimens less than 45 cm SL. Maxilla extends to or slightly beyond posterior margin of orbit, with ventral margin smoothly rounded and lacking step or bony knob. Supramaxilla is well developed. Teeth in front of jaws are small canines. Preoperculum is angular, with posterior margin serrate and serrae at angle large and forming lobe in large specimens. Dorsal margin of operculum is convex. Gill rakers on first arch number 23 to 25, with 8 or 9 on upper limb and 15 to 17 on lower limb. Measurements are expressed as percent of SL: head length 38%– 42%, body depth 34%–37%. Pectoral fin is broadly rounded and has 17 or 18 rays. Dorsal fin has 11 spines and 13 to 15 rays, with third or fourth spine longest. Anal fin has 9 rays. Caudal fin is truncate to slightly concave in specimens larger than 30 cm SL and convex in specimens less than 30 cm SL. Lateral body scales are ctenoid. Lateral line scales number about 65, and scales in horizontal series number 82 to 99. Color of head and body is buff to grayish brown dorsally and on sides, and white ventrally. Margins of dorsal and pectoral fins, and occasionally of anal and

caudal fins, are yellow except in largest specimens. Pale blue line extends from eye to corner of preoperculum. Juveniles (between 5 and 10 cm SL) have pearly white spots arranged in four longitudinal rows and seven vertical columns, broad yellow margin on dorsal fin, white caudal fin, blackish anal and pelvic fins, and black saddle on caudal peduncle that does not reach lateral line. This pattern is occasionally displayed in adult specimens. This species occurs between 64 and 275 m in the western Atlantic from North Carolina to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico and is associated with both rocky and soft sand and mud bottoms. Food consists of a large variety of invertebrates, especially brachyuran crabs, and ray-finned fishes. This species is a protogynous hermaphrodite; females mature at about 53 to 60 cm SL and change sex at about 75 cm SL. Life span is about 20 years, and maximum known size is 115 cm TL.

references: Smith 1971, 1997; Moore 1975a; Hoese and Moore 1977, 1998; Matsuura 1983a; Robins and Ray 1986; Jones et al. 1989; Bullock and Smith 1991; Cervigón 1991; Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; Heemstra 2002c.

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SERRANIDAE 145

Epinephelus guttatus Red hind

(Linnaeus, 1758)

Epinephelus guttatus has pelvic fin origin below or behind pectoral fin base, pectoral fin longer than pelvic fin, and head and body covered with dark orangish brown or red spots. Posterior naris is larger than anterior naris. Maxilla extends about to posterior margin of orbit, with ventral margin smoothly curved and lacking step or bony knob. Supramaxilla is well developed. Teeth in front of jaws are small canines. Preoperculum is angular, and posterior margin is evenly serrated. Dorsal margin of operculum is convex. Gill rakers on first arch number 24 to 26, with 8 or 9 on upper limb and 16 to 18 on lower limb. Measurements are expressed as percent of SL: head length 42%– 43%, body depth 32%–37%. Pectoral fin is broadly rounded and has 16 to 18 rays. Dorsal fin has 11 spines and 15 or 16 rays, with third or fourth spine longest and membrane between spines deeply incised and produced into small flag behind tip of each spine. Anal fin has 8 rays. Caudal fin is rounded. Lateral body scales are ctenoid. Scales in horizontal series number 92 to 104. Color is buff, greenish white, or light reddish brown, with head and body covered with bright orangish brown to dark red spots, spinous dorsal fin olive, and flags at tips of spines yellow. Caudal and rayed sections of dorsal and anal fins are olivaceous, with sub-

marginal black band and narrow pale margin. Pectoral fin is light orangish red, with darker red spots on base, and pelvic fin is colored like body. This species occurs between 2 and 100 m in the western Atlantic from North Carolina and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico and is generally associated with shallow coral reefs and rocky areas. Food consists of octopuses, crabs and other crustaceans, and ray-finned fishes. This species is a protogynous hermaphrodite; females mature at 22 to 24 cm SL and change sex at 28 cm SL. Most specimens larger than 40 cm SL are males. Fecundity ranges from 90,000 to 3,000,000 eggs per female. Maximum life span is at least 22 years, and maximum known size is 76 cm TL.

references: Menzel 1960; Randall 1968a, 1996; Smith 1971, 1997; Bright and Cashman 1974; Smith et al. 1975; Thompson and Munro 1978; Robins and Ray 1986; Colin et al. 1987; Bullock and Smith 1991; Cervigón 1991; Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Heemstra 2002c.

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146 PERCIFORMES

Epinephelus itajara (Lichtenstein, 1822) Jewfish

Epinephelus itajara has most dorsal fin spines shorter than dorsal fin rays, pelvic fin origin below or posterior to pectoral fin base, and pectoral fin longer than pelvic fin. Interorbital region is flat and very broad. Eye is relatively small. Maxilla is scaly and extends beyond posterior margin of orbit, with ventral margin smoothly rounded and lacking step and bony knob. Supramaxilla is well developed. Jaw teeth are small, and those in midlateral part of lower jaw are arranged in three to five rows. Preoperculum is subangular, with posterior margin evenly serrate. Operculum has convex dorsal margin. Gill rakers on first arch number 21 to 24, with 8 or 9 on upper limb and 13 to 15 on lower limb. Measurements are expressed as percent of SL: head 34%– 43%, body depth 29%–37%. Pectoral fin is broadly rounded and has 18 or 19 rays. Dorsal fin has 11 spines and 15 or 16 rays. Anal fin has 8 rays. Caudal fin is rounded. Body scales are ctenoid. Lateral line scales number 61 to 64, and scales in horizontal series number 89 to 110. Color is brown, brownish yellow, or green, with head, upper body, and fins covered with small black spots. Specimens less than 100 cm SL have three or four indistinct bars

on upper part of body. Specimens larger than 100 cm SL are darker than smaller specimens and lack bars. This species occurs from the shoreline to about 30 m in the tropical and warm temperate Atlantic and eastern Pacific Oceans. In the western Atlantic it occurs from Florida and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico. Juveniles occur inshore in mangrove swamps and bays. Adults are found on coral reefs and other areas with high relief. Food consists of octopods, shrimps, crabs, lobsters, rayfinned fishes, and young sea turtles; however, it feeds largely on crustaceans. Fecundity ranges up to 58 million eggs. Males mature at 110 to 115 cm SL, and females mature at 120 to 135 cm SL. Maximum known size is 250 cm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Smith 1971, 1997; Robins and Ray 1986; Bullock and Smith 1991; Cervigón 1991 (as E. [Promicrops] itajara); Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 147

Epinephelus morio Red grouper

(Valenciennes, 1828)

Epinephelus morio has pelvic fin origin below or behind pectoral fin base, pelvic fin shorter than pectoral fin, and dorsal fin membrane unnotched between spines. Maxilla extends to posterior margin of orbit, with ventral margin smoothly curved and lacking step or bony knob. Supramaxilla is well developed. Teeth in front of jaws are small canines. Preoperculum is subangular, with posterior margin serrate and serrae at angle enlarged. Dorsal margin of operculum is straight. Gill rakers on first arch number 23 to 25, with 8 or 9 on upper limb and 15 or 16 on lower limb. Measurements are expressed as percent of SL: head length 40%– 43%, body depth 33%–38%. Pectoral fin is broadly rounded and has 16 to 18 rays. Dorsal fin has 11 spines and 16 or 17 rays, with second or third spine longest. Anal fin has 8 to 10 rays. Caudal fin is convex in specimens less than 15 cm SL and truncate to slightly concave in larger specimens. Lateral body scales are ctenoid and have auxillary scales. Lateral line scales number 60 to 68, and scales in horizontal series number 112 to 128. Head and body are dark reddish brown dorsally and laterally, and pink to reddish ventrally. Rayed section of dorsal fin and anal and caudal fins are dark distally, with white margin. Inside of mouth is bright reddish brown.

Body occasionally has irregular white spots or blotches. This species occurs between the shore and 300 m in the western Atlantic from North Carolina and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico. Juveniles are found in shallow sea-grass beds and inshore reefs. Adults occur in 50 to 300 m on soft bottoms. Food consists of invertebrates and rayfinned fishes. Spawning takes place from April to May in the eastern Gulf of Mexico. Fecundity ranges from 312,000 to 5,735,700 eggs. Females mature at about 45 cm SL, at 4 to 6 years of age, and transform to males at about 50 cm SL, at 7 to 14 years of age. Maximum known age is 25 years, and maximum known size is 90 cm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Moe 1969; Smith 1971, 1997; Hastings et al. 1976; Hastings 1979; Matsuura 1983a; Robins and Ray 1986; Bullock and Smith 1991; Cervigón 1991 (as E. [Epinephelus] morio); Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Heemstra 2002c.

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148 PERCIFORMES

Epinephelus mystacinus Misty grouper

(Poey, 1852)

Epinephelus mystacinus has pelvic fin origin in front of pectoral fin base, pelvic fin nearly as long as pectoral fin, and body patterned with dark bars. Posterior naris is 3 to 5 times larger than anterior naris. Maxilla extends to posterior margin of orbit, with ventral margin smoothly curved and lacking step or bony knob. Supramaxilla is well developed. Teeth in front of jaws are small canines. Preoperculum is angular, with posterior margin serrate, serrae at angle enlarged, and lower margin with one or two serrae. Dorsal margin of operculum is convex. Gill rakers on first arch number 22 to 25, with 8 or 9 on upper limb and 14 to 16 on lower limb. Measurements are expressed as percent of SL: head length 40%– 43%, body depth 34%– 42%. Pectoral fin is broadly rounded and has 18 or 19 rays. Dorsal fin has 11 spines and 14 or 15 rays, with third or fourth spine longest and interspinous membranes distinctly incised. Anal fin has 9 or (rarely) 8 rays. Caudal fin is rounded or truncate. Body scales are ctenoid and lack auxillary scales. Lateral line scales number 58 to 69, and scales in horizontal series number 99 to 112. Color of head and body is buff, with eight or nine dark subvertical bars on body and last

two bars darker than others. Last two bars on caudal peduncle are darker than remainder and may be fused into broad band. Dark brown mark is located on cheek at upper edge of maxilla. Pelvic fin is darkly pigmented. Juveniles less than 20 cm SL have black bar or saddle on caudal peduncle. This species occurs in the western Atlantic and eastern Pacific. In the western Atlantic it occurs between 30 and 400 m, from North Carolina and Bermuda to Belize and Trinidad, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is absent along the coast of Central America south of Belize and the northern coast of South America. Juveniles occasionally occur as shallow as 30 m. Food consists of squids, crustaceans, and ray-finned fishes. Females reach 100 cm SL. Maximum known size is 115 cm TL.

references: Longley and Hildebrand 1941; Robins 1967b; Böhlke and Chaplin 1968; Smith 1971, 1997; Thompson and Munro 1978; Robins and Ray 1986; Bullock and Smith 1991; Cervigón 1991; Heemstra and Randall 1993; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 149

Epinephelus nigritus Warsaw grouper

(Holbrook, 1855)

Epinephelus nigritus has pelvic fin origin anterior to pectoral fin base, pelvic fin nearly as long as pectoral fin, and second dorsal fin spine longer than remainder of spines. Interorbital region is convex. Maxilla extends distinctly posterior to rear margin of orbit, with ventral margin smoothly rounded and lacking step and bony knob. Teeth in front of jaws are small canines. Preoperculum is rounded, with posterior margin serrated, serrae at corner slightly enlarged, and one or two serrae on lower margin. Gill rakers on first arch number 23 to 25, with 9 to 11 on upper limb and 14 to 16 on lower limb. Measurements are expressed as percent of SL: head length 40%– 48%, body depth 38%– 42%. Pectoral fin is broadly rounded and has 18 or 19 rays. Dorsal fin has 10 spines and 13 to 15 rays, with second spine largest and membranes deeply incised between spines. Anal fin has 9 rays. Caudal fin is convex to truncate with rounded corners. Midlateral body scales are ctenoid. Lateral line scales number 62 to 71, and scales in horizontal se-

ries number 99 to 107. Color is dark reddish brown, brownish gray, or nearly black dorsally and on sides, and dull reddish gray ventrally. Juveniles have a few whitish spots on body and a yellow caudal fin. This species occurs in the tropical to warm temperate Atlantic. In the western Atlantic it occurs between 55 and 525 m, from Massachusetts to southern Brazil, including the entire Gulf of Mexico and the coasts of Cuba, Haiti, and Trinidad. Juveniles occasionally occur around jetties and on shallow reefs. Adults are generally associated with rough bottoms. Food consists of crabs, shrimps, lobsters, and rayfinned fishes. Maximum known age is 41 years, and maximum known size is 235 cm TL. references: Smith 1971; Hastings et al. 1976; Matsuura 1983a; Robins and Ray 1986; Bullock and Smith 1991; Cervigón 1991; Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; Castro-Aguirre et al. 1999; Heemstra 2002c.

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150 PERCIFORMES

Epinephelus niveatus Snowy grouper

(Valenciennes, 1828)

Epinephelus niveatus has dorsal fin membranes between spines deeply incised and pelvic fin origin below or anterior to pectoral fin base. Posterior naris is 3 to 5 times diameter of anterior naris. Maxilla extends beyond posterior margin of orbit, with ventral margin smoothly curved and lacking step or bony knob. Supramaxilla is well developed. Teeth in front of jaws are small canines. Preoperculum is angular, with posterior margin serrate, enlarged serrae at angle, and occasionally a small antrorse spine on lower margin just anterior to angle and covered with skin. Dorsal margin of operculum is distinctly convex. Gill rakers on first arch number 22 to 26, with 7 to 10 on upper limb and 15 to 17 on lower limb. Measurements are expressed as percent of SL: head length 42%– 45%, body depth 36%– 40%. Pectoral fin is broadly rounded and has 18 or 19 rays. Dorsal fin has 11 spines and 13 to 15 rays, with second or third spine longest. Anal fin has 9 rays. Caudal fin is straight to slightly concave in adults and convex in juveniles. Midlateral scales are ctenoid. Lateral line scales number 64 to 73, and scales in horizontal series number 102 to 128. Color is dark brown, with margin of spinous part of dorsal fin black edged in all but largest specimens. Ju-

veniles are dark brown, with white spots arranged in about 11 vertical series and five or six longitudinal rows, black saddle on caudal peduncle extending below lateral line, and yellow pectoral and caudal fins. This species occurs between 10 and 400 m in the western Atlantic from Massachusetts and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, Cuba, and the Caribbean coast of Central and South America. It is found throughout the Gulf of Mexico. Juveniles occasionally occur inshore, and adults are associated with rocky bottoms. Food consists of gastropods, cephalopods, brachyuran crustaceans, and ray-finned fishes. Spawning occurs off the Florida Keys from April to July. Females mature between ages 4 and 5 and change to males by age 6 to 8. Maximum known age is 27, and maximum known size is 120 cm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Smith 1971, 1997; Matsuura 1983a; Robins and Ray 1986; Bullock and Smith 1991; Cervigón 1991; Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 151

Epinephelus striatus Nassau grouper

(Bloch, 1792)

Epinephelus striatus has pelvic fin origin below or behind pectoral fin base, pelvic fin shorter than pectoral fin, and dark tuning fork– shaped mark between eyes. Posterior naris is slightly larger than anterior naris and is comma shaped in adults. Maxilla extends to posterior margin of orbit, with ventral margin smoothly curved and lacking step or bony knob. Supramaxilla is well developed. Teeth in front of jaws are small canines. Preoperculum is rounded, with evenly serrate posterior margin. Dorsal margin of operculum is nearly straight. Gill rakers on first arch number 23 to 26, with 8 or 9 on upper limb and 15 to 17 on lower limb. Measurements are expressed as percent of SL: head length 38%– 42%, body depth 34%– 38%. Pectoral fin is broadly rounded and has 17 to 19 rays. Dorsal fin has 11 spines and 16 to 18 rays, with third or fourth spine longest and membrane between spines incised. Anal fin has 8 rays. Caudal fin is convex in adults and rounded in juveniles. Scales are ctenoid. Lateral line scales number about 50, and scales in horizontal series number 98 to 106. Color is generally buff, with dark band extending from upper jaw through eye to midline of nape; several dark dots below and behind eye; five broad, dark, irregular vertical bars on body; and black saddle on caudal peduncle. Submissive adults or juveniles display bicolored color pattern in which dorsolateral parts of head and

body are dark brown, and ventral, lateral, and ventral parts of body are white, with oblique white band running from upper jaw to origin of dorsal fin. This species occurs between the shoreline and 90 m in the western Atlantic from northern Florida and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs off Texas, southwestern Florida, and the west coast of Yucatán. Adults are associated with coral reefs, and juveniles occur in sea-grass beds. Food consists of mollusks, crustaceans (especially crabs), and ray-finned fishes. Spawning takes place from December through February near time of full moon along the outer reef edge. Assemblages of 3 to over 200,000 adults take part in group spawning, and most of the individuals have the bicolored color pattern. Larvae are pelagic for 37 to 45 days before settling on reef habitats. Maximum known size is 100 cm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Smith 1971, 1972; Hoese and Moore 1977, 1998; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Colin et al. 1987, 1997; Cervigón 1991; Heemstra and Randall 1993; Smith-Vaniz et al. 1999; Heemstra 2002c.

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152 PERCIFORMES

Gonioplectrus hispanus Spanish flag

(Cuvier, 1828)

Gonioplectrus hispanus has a large antrorse spine at corner of preoperculum and a very large knifelike middle opercular spine. Profile of head is convex, and top of head is rugose and not covered with skin or scales. Interorbital region is convex. Maxilla extends to posterior margin of pupil, with prominent bony knob on ventral margin (hidden by upper lip). Supramaxilla is well developed. Jaws have band of very small, slender, depressible teeth, and one to three fixed canines along midlength of lower jaw. Near symphysis, teeth are enlarged in inner row and short and curved in outer row. Preoperculum is rounded, and ventral margin is fleshy. Middle spine of operculum extends to or beyond edge of opercular margin, and dorsal margin of operculum is convex. Gill rakers on first arch number 20 to 24, with 5 to 7 on upper limb and 14 to 16 on lower limb. Measurements are expressed as percent of SL: head length 42%– 45%, body depth 40%– 43%. Pectoral fin is broadly rounded and has 16 or 17 rays, with upper rays longer than lower rays. Dorsal fin has eight spines and 13 rays, with third or fourth spine longest and membrane deeply incised between spines. Proximal

one-third of inner pelvic fin ray is connected to body by membrane. Anal fin has 7 rays. Caudal fin is truncate or rounded. Scales are large and cycloid, and auxillary scales are lacking. Lateral line scales number 47 to 49, and scales in horizontal series number 78 to 83. Scales are lacking on top of head, preorbital region, snout, maxilla, and lower jaw. Color of head, body, dorsal fin, and caudal fin is yellow, with six or seven salmon-colored horizontal stripes on body, bright red blotch on anterior section of anal fin, and white blotch on side of belly. First stripe runs along dorsal fin, and second runs from upper jaw through eye to end of dorsal fin. This species occurs between 60 and 365 m in the western Atlantic from North Carolina to southern Brazil, including the Gulf of Mexico, Cuba, and Jamaica. It is found throughout the Gulf of Mexico and is associated with rocky bottoms. Maximum known size is 27 cm TL. references: Colin 1974; Moore 1975a; Robins and Ray 1986; Bullock and Smith 1991; Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; Heemstra 2002c.

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SERRANIDAE 153

Hemanthias aureorubens Streamer bass

(Longley, 1935)

Hemanthias aureorubens has top of head free of scales and caudal fin lobes long and filamentous. Anterior and posterior nares are close set. Orbit length is greater than interorbital width. Mouth is very oblique, with upper jaw indented at symphysis. Supramaxilla is lacking. Teeth in upper jaw are in villiform band except for several moderate-sized canines on either side of indentation at symphysis. Teeth in lower jaw are weak and in a single series except for several canines on either side of symphysis. Vomerine tooth patch is chevron shaped to triangular and only rarely has posterior prolongation. Teeth are absent on tongue. Preoperculum is angular, with serrae on ventral and posterior margins. Suboperculum and interoperculum are serrated along free margins near their articulation. Operculum has small, blunt upper spinea; large, pointed median spinea; and small lower spines. Gill rakers on first arch are long and slender and number 36 to 43 (usually 37 to 41), with 10 or 11 on upper limb and 26 to 31 on lower limb. Measurements are expressed as percent of SL: head length 31%– 41%, snout length 6%–7%, orbit length 11%–19%, interorbital width 8%–9%, body depth 31%– 40%. Pectoral fin is long and

pointed and has 15 to 19 (usually 16 or 17) rays. Dorsal fin is continuous and notched and has 10 spines and 13 to 16 (usually 15) rays. Anal fin has 7 to 9 rays. Caudal fin is very deeply forked. Scales are large and ctenoid and cover body and head, with exception of snout, maxilla, anterior interorbital region, anterior section of lower jaw, and branchiostegal region. Scales encircling caudal peduncle number 22 to 27 (usually 23 to 26). Lateral line runs high along flank and has 42 to 50 (usually 44 to 47) tubed scales. Color is reddish orange dorsally and silvery laterally and ventrally. Iris of eye is yellow and red. Pectoral fin is pink. Dorsal fin and upper and lower lobes of caudal fin are yellowish. Anal fin, midsection of caudal fin, and pelvic fin are pallid. This species occurs between 91 and 610 m in the western Atlantic from New Jersey to Suriname, including the northern Gulf of Mexico and the Caribbean coast of South America. Maximum known size is 220 mm SL. references: Longley and Hildebrand 1940, 1941 (as Pronotogrammus aureorubens); Robins and Ray 1986; Bullock and Smith 1991; Cervigón 1991; Boschung 1992; Anderson 2002b.

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154 PERCIFORMES

Hemanthias leptus Longtail bass

(Ginsburg, 1952)

Hemanthias leptus has maxilla and top of head free of scales and a continuous lateral line. Anterior and posterior nares are close set. Orbit diameter is greater than interorbital width. Mouth is oblique, with upper jaw indented at symphysis. Supramaxilla is lacking. Upper jaw has outer row of conical teeth and inner band of villiform teeth, with two pairs of large canines, directed posteriorly or anteriorly, on either side of groove at symphysis. Lower jaw has series of conical teeth, with two or three pairs of canines on either side of symphysis projecting posteriorly or outward. Vomerine tooth patch is chevron shaped. Preoperculum is angular, with posterior margin finely serrate, ventral margin smooth to coarsely serrate, and angle with enlarged serrae, spinous processes, or one to several spines. Gill rakers on first arch are long and slender and number 34 to 39, with 9 to 12 on upper limb and 24 to 28 on lower limb. Measurements are expressed as percent of SL: head length 30%–36%, snout length 7%– 8%, orbit diameter 6%–13%, interorbital width 6%–7%, body depth 27%–38%. Pectoral fin is of moderate length, is pointed, and has 17 to 19 rays. Dorsal fin is continuous and has 10 spines and 13 to 15 rays, with third spine bearing long filament in adults. Pelvic fin is produced in specimens longer than 190 mm SL. Anal fin has 8 (rarely 7) rays. Caudal fin is deeply forked and bears short filaments in small

specimens (200 mm SL) and is truncate with median notch in large specimens (310 mm SL). Scales are large and ctenoid and cover body and head except for snout, maxilla, most of interorbital region, lower jaw, and gular and branchiostegal regions. Scales encircling caudal peduncle number 40 to 46. Lateral line runs high along flank and has 54 to 62 tubed scales. Color is red dorsally and pink to silvery laterally and ventrally, with yellow or golden stripe running below eye from tip of snout to pectoral fin base. Fins are yellow, with reddish bases. This species occurs between 35 and 640 m in the western Atlantic from North Carolina to Suriname, including the Gulf of Mexico. It is found in both the northern and southern Gulf of Mexico and appears to be more common in the western than in the eastern Gulf. There is some evidence that large females transform into functional males (protogyny), but small males are also known, suggesting that this species has both primary and secondary males. Food consists of zooplankton such as copepods. Maximum known size is 456 mm SL.

references: Ginsburg 1952a, 1954 (both as Anthiasicus leptus); Briggs et al. 1964; Moore 1975a; Hoese and Moore 1977, 1998; Robins and Ray 1986; Cervigón 1991; Boschung 1992; Bullock and Smith 1992; Schaldach et al. 1997; Anderson 2002b.

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SERRANIDAE 155

Hemanthias vivanus Red barbier

(Jordan and Swain, 1885)

Hemanthias vivanus has longest pelvic fin ray and caudal fin lobes filamentous and maxilla free of scales. Anterior and posterior nares are close set, and posterior margin of anterior naris is produced into short flap. Orbit diameter is greater than interorbital width. Upper jaw is indented at symphysis. Supramaxilla is lacking. Upper jaw has outer row of small, slightly curved conical teeth and inner band of villiform teeth, with two pairs of large, backward- or forward-directed canines on either side of symphysis. Lower jaw has row of small, slightly curved conical teeth and short band of villiform teeth on either side of symphysis, with two or more pairs of large, backward- or forward-directed teeth on either side of symphysis. Vomerine tooth patch is chevron shaped to triangular without posterior prolongation. Preoperculum is angular, with serrae on posterior margin, several serrae on ventral margin, and large, flat spines or spinelike processes at angle. Gill rakers on first arch are long and slender and number 38 to 43 (usually 39 to 42), with 10 to 13 on upper limb and 27 to 31 on lower limb. Measurements are expressed as percent of SL: head length 28%–32%, snout length 6%– 8%, orbit diameter 7%–11%, interorbital width 6%–7%, body depth 26%– 35%. Pectoral fin is acutely pointed and has 16

to 21 (usually 18 or 19) rays. Dorsal fin is continuous and has 10 spines and 14 (occasionally 13) rays, with long filaments extending from spines. Filament on fourth spine is usually longest, especially in males. Anal fin has 3 spines and 8 (occasionally 9) rays. Caudal fin is deeply forked, and upper and lower lobes bear long filaments. Scales are large and ctenoid and cover body and head except for snout, maxilla, all or most of interorbital region, lower jaw, and gular and branchiostegal regions. Scales encircling caudal peduncle number 24 to 29 (usually 26 to 29). Lateral line is continuous, runs high along flank, and consists of 44 to 53 (usually 46 to 51) tubed scales. Color is deep red, with yellowish to violet spots on back and flanks and two gold-colored stripes on head. Anal fin is bright yellow in males and mottled blue and olive in females. This species occurs between 20 and 430 m in the western Atlantic from North Carolina to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it occurs from the Dry Tortugas to Alabama and in Campeche Bay. Food consists of calanoid and cyclopoid copepods, amphipods, and ostracods. Individuals form large schools. This species is a protogynous hermaphrodite, and spawning takes place during the winter in

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156 PERCIFORMES

the eastern Gulf of Mexico. Maximum known size is 150 mm SL. references: Longley and Hildebrand 1941; Hoese and Moore 1977, 1998; Hastings 1981;

Darcy and Gutherz 1984; Robins and Ray 1986; Bullock and Smith 1991; Cervigón 1991; Boschung 1992; Schaldach et al. 1997; Anderson 2002b.

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SERRANIDAE 157

Hypoplectrus chlorurus Yellowtail hamlet

(Cuvier, 1828)

Hypoplectrus chlorurus has body uniformly dark brown or bluish brown and caudal fin bright yellow. Head is compressed and of moderate length. Small canine teeth occur in jaws. Vomerine and palatine teeth are villiform and arranged in narrow bands. Preoperculum is angular, with serrae on ventral and posterior margins and several small antrorse spines on ventral margin near angle. Gill rakers on first arch are moderately long and slender and number 6 to 8 on upper limb and 12 or 13 on lower limb. Measurements are expressed as percent of SL: head length 38%–39%, snout length 13%–14%, eye length 9%–10%, upper jaw length 17%–19%, pectoral fin length 31%– 32%, pelvic fin length 24%–27%, body depth 45%– 47%. Pectoral fin is long and rounded, with 13 or 14 rays. Dorsal fin is unnotched between spinous and rayed sections, originates anterior to pectoral fin base, and has 10 spines and 14 or 15 rays. Anal fin has 7 rays. Cau-

dal fin is moderately concave, with lower lobe about equal to upper lobe. Scales are relatively small and ctenoid. Lateral line is gently arched anteriorly and has 48 to 53 pored scales. Color in life is dark bluish or brownish black, except caudal fin is bright yellow or turquoise. Pectoral fins are generally transparent but occasionally yellow. This species occurs in shallow water in the western Atlantic from the northwestern and southern Gulf of Mexico, the Greater and Lesser Antilles, the coast of Panama, and Venezuela. It is considered to be a simultaneous hermaphrodite and to mimic the pomacentrid Microspathodon chrysurus. Food consists mostly of crustaceans. Maximum know size is about 150 mm TL. references: Woods 1942; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Thresher 1978; Domeier 1994; Lobel 2002.

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158 PERCIFORMES

Hypoplectrus gemma

Goode and Bean, 1882

Hypoplectrus gemma has dark upper and lower margins of caudal fin. Head is compressed and of moderate length. Small canine teeth occur in jaws. Vomerine and palatine teeth are villiform and arranged in narrow bands. Preoperculum is angular, with serrae on ventral and posterior margins and several small antrorse spines on ventral margin near angle. Gill rakers on first arch are moderately long and slender and number 6 or 7 on upper limb and 12 or 13 on lower limb. Measurements are expressed as percent of SL: head length 37%– 40%, snout length 12%–14%, eye length 8%–9%, upper jaw length 18%– 19%, pectoral fin length 29%–30%, pelvic fin length 24%–26%, body depth 41%– 43%. Pectoral fin is long and rounded, with 13 or 14 rays. Dorsal fin is unnotched between spin-

ous and rayed sections, originates anterior to pectoral fin base, and has 10 spines and 14 to 16 rays. Anal fin has 7 rays. Caudal fin is strongly concave. Scales are relatively small and ctenoid. Lateral line is gently arched anteriorly and has 48 to 53 pored scales. Color in life is iridescent blue to black, with upper and lower margins of caudal fin dark blue to black and remainder of fin light. This species occurs in shallow water in the western Atlantic from southern Florida, the Dry Tortugas, and the Bahamas. It is considered to be a simultaneous hermaphrodite and to mimic the pomacentrid Chromis cyanea. Maximum known size is 130 mm TL. references: Longley and Hildebrand 1941; Thresher 1978; Domeier 1994; Lobel 2002.

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SERRANIDAE 159

Hypoplectrus nigricans

(Poey, 1852)

Hypoplectrus nigricans has a uniformly dark body and fins. Head is compressed and of moderate length. Small canine teeth occur in jaws. Vomer and palatine teeth are villiform and arranged in narrow bands. Preoperculum is angular, with serrae on ventral and posterior margins and several small antrorse spines on ventral margin near angle. Gill rakers on first arch are moderately long and slender and number 7 on upper limb and 13 or 14 on lower limb. Measurements are expressed as percent of SL: head length 38%–39%, snout length 13%–14%, eye diameter 8%–9%, upper jaw length 17%–18%, pectoral fin length 29%– 31%, pelvic fin length 25%–26%, body depth 44%– 46%. Pectoral fin is long and rounded, with 13 or 14 rays. Dorsal fin is unnotched between spinous and rayed sections, originates anterior to pectoral fin base, and has 10 spines and 14 or 15 rays. Anal fin has 7 rays. Caudal fin is moderately concave. Scales are rela-

tively small and ctenoid. Lateral line is gently arched anteriorly and has 48 to 53 pored scales. Color in life is dark bluish, purple, or brownish black, with head and lower part of body more bluish than brown. Median fins are dark bluish brown. This species occurs in the western Atlantic from southern Florida and the Bahamas to Panama, including the eastern Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from the Dry Tortugas in shallow water. This species is considered a simultaneous hermaphrodite and a mimic of the dark-colored pomacentrids of the genus Stegastes. Maximum known size is about 150 mm TL.

references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Thresher 1978; Domeier 1994; Smith 1997; Lobel 2002.

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160 PERCIFORMES

Hypoplectrus puella Barred hamlet

(Cuvier, 1828)

Hypoplectrus puella has a broad dark brown bar on front half of body and three narrow brown bars on posterior half of body. Head is compressed and of moderate length. Small canine teeth occur in jaws. Vomer and palatine teeth are villiform and arranged in narrow bands. Preoperculum is angular, with serrae on ventral and posterior margins and several small antrorse spines on ventral margin near angle. Gill rakers on first arch are moderately long and slender and number 6 to 8 on upper limb and 11 to 15 on lower limb. Measurements are expressed as percent of SL: head length 38%– 44%, snout length 13%–16%, eye length 8%–9%, upper jaw length 18%– 19%, pectoral fin length 25%–31%, pelvic fin length 23%–26%, body depth 36%– 46%. Pectoral fin is long and rounded, with 13 or 14 rays. Dorsal fin is unnotched between spinous and rayed sections, originates anterior to pectoral fin base, and has 10 spines and 14 to 16 rays. Anal fin has 7 rays. Caudal fin is slightly forked. Scales are relatively small and ctenoid. Lateral line is gently arched anteriorly and has 48 to 53 pored scales. Color in life is creamy white to pale yellow, with six bars on side of head and body. First bar runs through

eye; second extends from nape to pelvic fin insertion; third extends from base of spinous section of dorsal fin to belly; and fourth, fifth, and sixth are located on posterior half of body. First and third bars may narrow ventrally, and last bar is located on caudal peduncle. Blue lines are located on head and body between bars. Pectoral fin is transparent, and pelvic fin is usually white to yellowish but can be darkly pigmented. There are four color morphs that vary in darkness of bars on body. Small juveniles have two black spots on caudal peduncle, with squarish white blotch above each. Large juveniles have dark blotch anterior to upper black spot. Color in preservative is light with six dark bars, and light lines between bars on head and body represent blue lines in life. This species occurs in shallow water around reefs in the western Atlantic from southern Florida and Bermuda to Venezuela, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is apparently the most common species of the genus in the Gulf of Mexico, and unlike the other congeners, it occurs in shallow, murky waters. Food consists mainly of crustaceans. This spe-

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SERRANIDAE 161

cies is considered to be a simultaneous hermaphrodite. Maximum known size is about 140 mm TL. references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Barlow 1975; Smith

et al. 1975; Thresher 1978; Bullock and Smith 1991; Domeier 1994; Randall 1996; Smith 1997; Hoese and Moore 1998; Smith-Vaniz et al. 1999; Lobel 2002.

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162 PERCIFORMES

Hypoplectrus unicolor Butter hamlet

(Walbaum, 1792)

Hypoplectrus unicolor has a black blotch or saddle on the caudal peduncle. Head is compressed and of moderate length. Small canine teeth occur in jaws. Vomer and palatine teeth are villiform and arranged in narrow bands. Preoperculum is angular, with serrae on ventral and posterior margins and several small antrorse spines on ventral margin near angle. Gill rakers on first arch are moderately long and slender and number 6 or 7 on upper limb and 12 to 15 on lower limb. Measurements are expressed as percent of SL: head length 37%– 39%, snout length 12%–13%, eye length 8%– 9%, upper jaw length 18%–19%, pectoral fin length 29%–32%, pelvic fin length 24%– 25%, body depth 41%–51%. Pectoral fin is long and rounded, with 13 to 15 rays. Dorsal fin is unnotched between spinous and rayed sections, originates anterior to pectoral fin base, and has 10 spines and 13 to 15 rays. Anal fin has 7 rays. Caudal fin is slightly concave. Scales are relatively small and ctenoid. Lateral line is gently arched anteriorly and has 77

to 82 pored scales. Color in life is pale yellow, with dorsal section of body generally darker than ventral section and iridescent blue lines running vertically on head. Blue line encircles orbit and extends ventrally on side. Blue-edged black spot is occasionally located on snout. Pectoral fin is transparent, and pelvic fin is yellowish. This species occurs in shallow water in the western Atlantic from southern Florida and the Bahamas to Venezuela, including the Antilles, the northern Gulf of Mexico off Port Aransas, and the southern Gulf of Mexico off Veracruz. Food consists of penaeid shrimps and rayfinned fishes. Maximum known size is about 120 mm TL. references: Gunter and Knapp 1951; Böhlke and Chaplin 1968; Hoese and Moore 1977, 1998; Thresher 1978; Robins and Ray 1986 (in part); Cervigón 1991 (in part as H. puella); Domeier 1994; Randall 1996; Smith 1997; Lobel 2002.

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SERRANIDAE 163

Liopropoma eukrines Wrasse bass

(Stark and Courtenay, 1962)

Liopropoma eukrines has spinous dorsal fin interrupted by a scaled area. Head is conical and of moderate length. Snout is acute and slightly longer than eye diameter. Anterior and posterior nares are widely separated, with anterior naris tubular and adjacent to lip, and posterior naris just anterior to eye. Maxilla extends to posterior margin of pupil. Supramaxilla is present. Jaw, vomerine, and palatine teeth are villiform. Free margin of preoperculum is smooth. Operculum has weak upper and strong lower spine. Gill rakers on first arch number 16 to 18. Measurements are expressed as percent of SL: head length 35%– 41%, snout length 9%–11%, eye diameter about 8%, interorbital width about 5%, body depth 22%– 29%. Pectoral fin has 13 or 14 rays. Dorsal fin has eight spines and 12 rays, with seventh and eighth spines lacking membranes and surrounded by scales at base. Anal fin has 8 rays. Caudal fin is truncate. Scales are ctenoid. Head is scaled except for lips. Lateral line is arched anteriorly and has 44 pored scales. Vertebrae number 24: 10 precaudal and 14 caudal. Body

is bright red in life, with dark brown to black stripe extending from tip of snout through eye to caudal fin base, white to yellowish stripe on either side of dark stripe, and dark reddish brown mid-dorsal stripe. Rayed sections of dorsal and anal fins are pale red. Caudal fin has submarginal black blotch merging with midbody stripe and white posterior margin. Juveniles are yellow, with black stripe. This species occurs between 20 and 116 m in the western Atlantic from North Carolina to the Florida Keys and the northern Gulf of Mexico. In the Gulf of Mexico it is known from the Florida Middle Grounds and West Flower Garden Reef. This species is usually associated with rocky or coral reefs. Maximum known size is 100 mm SL.

references: Stark and Courtenay 1962 (as Chorististium eukrines); Bright and Cashman 1974; Hoese and Moore 1977, 1998; Shipp and Hopkins 1978; Clarke 1986; Robins and Ray 1986; Bullock and Smith 1991; Boschung 1992; Smith 1997; Heemstra 2002c.

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164 PERCIFORMES

Liopropoma rubre Poey, 1861 Peppermint bass

Liopropoma rubre has a spinous dorsal fin interrupted by a scaled area. Head is conical and of moderate length. Snout is acute and slightly longer than eye diameter. Anterior and posterior nares are widely separated, with anterior one tubular and adjacent to lip, and posterior one just anterior to eye. Maxilla extends to posterior margin of pupil. Supramaxilla is present. Jaw, vomerine, and palatine teeth are villiform. Preopercular margin is smooth. Operculum has weak upper and strong lower spine. Gill rakers on first arch number 5 on upper limb and 11 on lower limb. Measurements are expressed as percent of SL: head length 37%– 43%, snout length 9%–10%, eye diameter 8%–10%, interorbital width about 9%, body depth 27%–33%. Pectoral fin has 12 or 13 rays. Dorsal fin originates slightly posterior to pectoral fin base and has eight spines and 12 rays, with seventh and eighth spines short, free of membranes, and surrounded by scales. Anal fin has 8 rays. Caudal fin is truncate. Scales are ctenoid. Head is scaled except for

lips. Lateral line is arched anteriorly and has 45 to 47 pored scales. Color is pale yellow, with five deep reddish brown stripes alternating with narrow pink stripes. Fins are pinkish red. Outer aspects of rayed section of dorsal fin, anal fin, and caudal fin have oval black blotches. This species occurs between 2 and 43 m in the western Atlantic from southern Florida and Bermuda to Venezuela, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it occurs on the Flower Garden Reefs and in Campeche Bay. Maximum known length is about 80 mm SL.

references: Böhlke 1956 (as Chorististium rubrum); Stark and Courtenay 1962 (as C. rubrum); Böhlke and Chaplin 1968 (as C. rubrum); Randall 1968a, 1996; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Robins and Ray 1986; Cervigón 1991; Smith 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 165

Mycteroperca acutirostris Comb grouper

(Valenciennes, 1828)

Mycteroperca acutirostris has a relatively broad maxilla and a body patterned with irregular white blotches and spots. Profile of head is evenly convex. Interorbital region is convex. Maxilla extends almost to posterior margin of orbit, with distal part of ventral margin straight and lacking step or bony knob. Supramaxilla is well developed. Jaws bear welldeveloped canine teeth on side of symphysis. Preoperculum is angular and posterior margin is serrated, with serrae at angle enlarged. Dorsal margin of operculum is convex. Gill rakers number 48 to 55, with 16 to 20 on upper limb and 32 to 36 on lower limb. Measurements are expressed as percent of SL: head length 37%– 40%, maxilla width 4%– 6%, body depth 31%–37%. Pectoral fin is broadly rounded and has 15 to 17 rays. Dorsal fin has 11 spines and 15 to 17 rays, with membrane between spines incised and posterior margin rounded. Anal fin has 10 to 12 rays, with outer margin pointed in adults. Caudal fin is rounded in specimens less than 10 cm SL, truncate in specimens between 10 and 12 cm SL, and concave in specimens greater than 25 cm SL. Lateral body scales are ctenoid. Lateral line scales number 67 to 77, and scales in horizontal se-

ries number 85 to 106. Head and body are grayish brown and covered with irregular blotches and spots, with three or four brown stripes radiating posteriorly from eye and extending along ventral section of body, and a single brown stripe extending from maxilla to preoperculum. Median fins are darker than body and covered with white spots and streaks. Juveniles less than 15 cm SL have black saddle on caudal peduncle, and the largest individuals are uniformly gray. This species occurs between the shoreline and 25 m in the western Atlantic from the northwestern Gulf of Mexico and the Greater and Lesser Antilles to southern Brazil. In the western Gulf of Mexico it is common near jetties at Port Aransas and 7 Fathom Reef. Juveniles are associated with turtle-grass beds, mangroves, soft corals, and coral reefs. Adults are associated with hard bottoms with high relief. Maximum known size is 80 cm TL. references: Randall 1968a, 1996; Smith 1971; Hoese and Moore 1977, 1998; Cervigón 1991 (all previous as M. rubra); Heemstra and Randall 1993; Smith 1997; Hoese and Moore 1998; Heemstra 2002c.

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166 PERCIFORMES

Mycteroperca bonaci Black grouper

(Poey, 1860)

Mycteroperca bonaci has a rounded preoperculum, yellowish brown spots on ventral aspect of body, and pectoral fin outlined with orange pigment. Maxilla extends beyond middle of eye, with distal part of ventral margin straight and lacking step or bony knob. Supramaxilla is well developed. Jaws bear welldeveloped canines on side of symphysis. Preoperculum is evenly rounded, and posterior margin is serrated, without notch or lobe at angle. Dorsal margin of operculum is convex. Gill rakers on first arch number 19 to 27, with 2 to 5 on upper limb and 8 to 12 on lower limb. Measurements are expressed as percent of SL: head length 36%– 40%, body depth 29%– 30%. Pectoral fin is broadly rounded and has 16 or 17 rays. Dorsal fin has 11 spines and 15 to 17 rays, with membrane between spines distinctly incised. Anal fin has 11 to 13 rays. Posterior margins of dorsal and anal fins are rounded. Caudal fin is truncate to slightly emarginate. Lateral line scales number 78 to 83, and scales in horizontal series number 119 to 126. Color is grayish to dark brown and densely covered with irregular bronze or brassy spots separated by bluish reticulum. Brassy spots may coalesce into chainlike horizontal streaks. Dorsolateral aspect of body occasion-

ally has seven or eight columns of rectangular dark blotches; first is above operculum, and last is on caudal peduncle. Pectoral fins are dusky brown with orange margin. This species occurs between 10 and 30 m in the western Atlantic from Bermuda and the Gulf of Mexico to southern Brazil, including the Bahamas and the Greater and Lesser Antilles. It is known from the northern and southern Gulf of Mexico. Adults are associated with rocky bottoms and coral reefs, and juveniles are associated with mangrove habitats. Food consists of crustaceans (juveniles) and ray-finned fishes (adults). This species is a protogynous hermaphrodite, with individuals maturing as females between 50 and 100 cm TL and becoming functional males at 96 to 116 cm TL. Maximum age is 17 to 19 years, and maximum known size is 133 cm TL.

references: Springer and McErlean 1962; Böhlke and Chaplin 1968; Randall 1968a, 1996; Smith 1971, 1997; Smith et al. 1975; Hoese and Moore 1977; Robins and Ray 1986; Bullock and Smith 1991; Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 167

Mycteroperca interstitialis Yellowmouth grouper

(Poey, 1860)

Mycteroperca interstitialis has preoperculum notched above angle and rays of median fins produced beyond membrane in adults. Profile of head is evenly convex. Posterior naris is greatly enlarged in adults. Maxilla extends to about middle of eye, with ventral margin straight and lacking step or bony knob. Supramaxilla is well developed. Large canines occur in anterior section of jaws. Preoperculum is angular, with posterior margin serrated and angle extended as lobe bearing enlarged serrae. Dorsal margin of operculum is convex. Gill rakers on first arch number 23 to 27. Measurements are expressed as percent of SL: head length 33%–36%, body depth 29%–33%. Pectoral fin is broadly rounded and has 16 or 17 rays. Dorsal fin has 11 spines and 16 to 18 rays, with twelfth and thirteenth rays elongate in adults. Anal fin has 11 or 12 rays, with sixth to eighth rays elongated in adults. Caudal fin is emarginate. Lateral body scales are ctenoid. Lateral line scales number 70 to 74, and scales in horizontal series number 112 to 119. Color is light brownish gray, with dense covering of small brown spots on dorsal and upper lateral sections of body. Mouth and margin of spinous

dorsal fin are yellow, and pectoral fin rays are black, with membranes unpigmented and fin margin white. Occasionally specimens are uniformly brown dorsally or have faint irregular bars on dorsal half of body. Juveniles are brown dorsally and white ventrally, with white middorsal stripe from tip of lower jaw to origin of dorsal fin. Occasionally dorsal surface is patterned with dark bars or saddlelike blotches. This species occurs between 20 and 150 m in the western Atlantic from North Carolina and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with coral reefs and rocky areas. In the Gulf Mexico it is found off western Florida and the Flower Garden Banks and in Campeche Bay. Food consists of ray-finned fishes. Maximum known size is 74 cm TL.

references: Randall 1968a; 1996; Böhlke and Chaplin 1968; Smith 1971, 1997; Robins and Ray 1986; Dennis and Bright 1988b; Cervigón 1991; Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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168 PERCIFORMES

Mycteroperca microlepis Gag

(Goode and Bean, 1879)

Mycteroperca microlepis has preoperculum notched above angle and rays of median fins not produced beyond membrane in adults. Profile of head is evenly convex. Anterior and posterior nares are close set, and posterior naris is considerably larger than anterior naris in adults. Maxilla extends beyond posterior margin of orbit, with ventral margin straight and lacking step or bony notch. Supramaxilla is well developed. Canine teeth occur in anterior part of jaws. Preoperculum is angular, with serrae on posterior margin and angle extended as lobe bearing enlarged serrae. Dorsal margin of operculum is convex. Gill rakers on first arch number 21 to 29. Measurements are expressed as percent of SL: head length 37%– 40%, body depth 29%–33%. Pectoral fin is broadly rounded and has 16 to 18 rays. Dorsal fin has 11 spines and 16 to 18 rays, with third or fourth spine longest and membrane incised between spines. Anal fin has 10 to 12 rays. Posterior margins of dorsal and anal fins are rounded. Caudal fin is truncate in juveniles and emarginate in large adults. Scales on lateral aspect of body, except those under pectoral fin, are smooth. Lateral line scales number 88 to 96, and scales in horizontal series number 128 to 146. Females and juveniles are brownish gray, with darker vermiculations and five brown saddles separated by short white bars under dorsal fin. Males have two color patterns: Some are pale gray, with slightly dark

vermiculations below rayed section of dorsal fin, on belly, on ventral part of body above anal fin, and on posterior margins of pectoral fin, pelvic fin, and rayed part of dorsal fin. Other males are more darkly pigmented on posterior section of body, with snout and front of jaws, upper part of caudal peduncle, and rayed sections of dorsal and anal fins black. Posterior margin of caudal fin is black. This species occurs, generally between 40 and 152 m, in the western Atlantic from Massachusetts and Bermuda to southern Brazil, including the Gulf of Mexico and Cuba. It is found throughout the Gulf of Mexico. Adults are limited to continental waters and are associated with rocky areas of high relief, although they occasionally occur in shallow rocky or grassy areas. Juveniles occur in estuaries and sea-grass beds. Food consists of cephalopods, shrimps, crabs, and ray-finned fishes. Females are sexually mature by age five or six years, at 67 to 75 cm TL, and sex change occurs by age 10 or 11 years, at 95 to 100 cm TL. Maximum known size is 120 cm TL.

references: Longley and Hildebrand 1941; Smith 1971, 1997; Smith et al. 1975; Hoese and Moore 1977, 1998; Robins and Ray 1986; Collins et al. 1987; Bullock and Smith 1991; Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 169

Mycteroperca phenax Scamp

Jordan and Swain, 1884

Mycteroperca phenax has notch in posterior margin of preoperculum above angle, and body is covered with small reddish brown spots. Posterior naris is 2 to 4 times larger than anterior naris. Maxilla extends to posterior margin of orbit, with ventral margin straight and lacking step and bony knob. Supramaxilla is well developed. Canine teeth are present in anterior section of jaws. Preoperculum is angular, with posterior margin serrate and angle extended as lobe bearing serrae. Gill rakers on first arch number 26 to 31, with 8 to 10 on upper limb and 17 to 21 on lower limb. Measurements are expressed as percent of SL: head length 33%– 38%, body depth 29%–33%. Pectoral fin is broadly rounded and has 15 to 17 rays. Dorsal fin has 11 spines and 16 to 18 rays. Anal fin has 10 to 12 rays. Caudal fin is concave. Fin rays of dorsal, anal, and caudal fins are unevenly exserted (extended beyond membranes). Lateral body scales are ctenoid. Lateral line scales number 76 to 82, and scales in horizontal series number about 124. Color is variable. Generally head and body are pale brown, with a single small reddish brown spot on each scale

of dorsal and lateral surfaces and on medial fins. Occasionally body is pale brown, with several clusters of dark brown spots resembling cat paw prints on dorsolateral parts of body. In large specimens, head and anterior one-third of body are silvery gray with dark reticulations, and posterior two-thirds of body is dark. This species occurs between 30 and 100 m in the western Atlantic from Massachusetts to Venezuela, including the Gulf of Mexico and the islands of Margarita and Cubagua off eastern Venezuela. It is found throughout the Gulf of Mexico. This species is associated with rocky and live bottom areas. Juveniles are found in shallow waters around jetties and in mangrove habitats. Food consists of octopods, crustaceans, and ray-finned fishes. Spawning occurs from March through May in the Gulf of Mexico. Maximum known age is 21 years, and maximum known size is 900 mm TL. references: Smith 1971, 1997; Hoese and Moore 1977, 1998; Robins and Ray 1986; Cervigón 1991; Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; Heemstra 2002c.

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170 PERCIFORMES

Mycteroperca tigris Tiger grouper

(Valenciennes, 1833)

Mycteroperca tigris has a rounded preoperculum and exserted median fin rays. Profile of head is evenly convex. Posterior naris is 3 to 5 times greater than anterior naris. Maxilla extends distinctly beyond posterior margin of orbit in adults, and ventral margin is straight and lacks step or bony knob. Supramaxilla is well developed. Canines occur in anterior section of jaws. Preoperculum is rounded, with posterior margin serrated. Dorsal margin of operculum is convex. Gill rakers on first arch are short and number 23 to 25, with 8 on upper limb and 15 to 17 on lower limb. Measurements are expressed as percent of SL: head length 36%– 40%, body depth 28%–32%. Pectoral fin is broadly rounded and has 17 rays. Dorsal fin has 11 spines and 15 to 17 rays. Anal fin has 11 rays. Large specimens have pointed dorsal and anal fins. Caudal fin is rounded in juveniles and truncate or emarginate in adults. Midlateral body scales are ctenoid in juveniles and cycloid in adults. Lateral line scales number 82 or 83, and scales in horizontal series number about 120. Color in adults is greenish brown to brownish gray, darker dorsally

than ventrally, and with 9 to 11 oblique pale stripes. Median fins have irregular pale spots and stripes, and pectoral fin is pale yellow near tips. Small juveniles are yellow, with dark brown midlateral stripe in addition to oblique pale stripes. This species occurs between 10 and 40 m in the western Atlantic from southern Florida and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it is found along southwestern Florida and from Texas to Yucatán, and it is abundant on the Flower Garden Banks. It is associated with coral reefs and rocky substrates. Food consists of a variety of ray-finned fishes. Specimens less than 37 cm TL are females, and those greater than 45 cm TL are males. Maximum known size is 100 cm TL. references: Longley and Hildebrand 1941; Smith 1971, 1997; Dennis and Bright 1988a; Cervigón 1991; Heemstra and Smith 1993; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 171

Mycteroperca venenosa Yellowfin grouper

(Linnaeus, 1758)

Mycteroperca venenosa has a rounded preoperculum, and distal third of pectoral fin is bright yellow. Posterior naris is slightly larger to twice as large as anterior naris. Maxilla extends to posterior margin of orbit, with ventral margin straight. Supramaxilla is well developed. Well-developed canines occur in front of jaws. Preoperculum is evenly rounded or has slight notch above angle, with posterior margin serrated. Dorsal margin of operculum is convex. Gill rakers on first arch number 24 to 27, with 8 to 10 on upper limb and 17 or 18 on lower limb. Measurements are expressed as percent of SL: head length 34%–38%, body depth 31%–34%. Pectoral fin is broadly rounded and has 16 to 18 rays. Dorsal fin has 11 spines and 15 or 16 rays, with membrane incised between spines. Anal fin has 10 to 12 rays. Caudal fin is truncate in juveniles and concave in adults. Midbody scales are ctenoid in juveniles and cycloid in adults. Lateral line scales number 72 to 81, and scales in horizontal series number 111 to 125. Color is reddish in deep water and greenish in shallow water, with oblong dark blotches on head and body

overlying small black spots. Ventral sections of head and body are covered with small dark red spots, and caudal fin and rayed sections of dorsal and anal fins have dark margin and white edge. This species occurs between 2 and 137 m in the western Atlantic from North Carolina and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is uncommon in the northern Gulf of Mexico. Juveniles are associated with turtle-grass beds in shallow water, and adults are associated with rocky substrates and coral reefs. Specimens in the Gulf of Mexico are also associated with muddy bottoms. Food consists of squids and ray-finned fishes, though squids are a minor resource. Maximum known size is 100 cm TL.

references: Longley and Hildebrand 1941; Smith 1971, 1997; Moore 1975a; Smith et al. 1975; Hoese and Moore 1977, 1998; Bullock and Smith 1991; Cervigón 1991; Boschung 1992; Heemstra and Randall 1993; Schaldach et al. 1997; SmithVaniz et al. 1999; Heemstra 2002c.

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172 PERCIFORMES

Paranthias furcifer (Valenciennes, 1828) Creolefish

Paranthias furcifer has a small mouth and a deeply forked caudal fin. Profile of head is convex. Maxilla extends to, but not beyond, center of eye, with ventral margin smoothly curved and lacking step or bony knob. Supramaxilla is rudimentary or absent. Canine teeth in jaws are poorly developed, and teeth are arranged in oval patch in vomer. Preoperculum is subangular, with vertical margin and posterior section of ventral margin finely serrate. Dorsal margin of operculum is slightly convex. Gill rakers on first arch number 38, with 12 to 14 on upper limb and 24 to 26 on lower limb. Measurements are expressed as percent of SL: head length 26%–29%, body depth 29%– 34%. Pectoral fin is bluntly pointed and has 19 or 20 rays. Dorsal fin has nine spines and 17 to 19 rays, with membrane slightly incised between spines. Anal fin has 8 or 9 (rarely 10) rays. Caudal fin has 8 branched rays and 12 or 13 procurrent rays in upper section, and 7 branched rays and 11 or 12 procurrent rays in lower section. Midlateral body scales are ctenoid. Lateral line scales number 69 to 77, and scales in horizontal series number 124 to 129. Color is reddish brown and paler ventrally than dorsally, with bright orangish red spot on upper end of pectoral fin base, three widely

separated white spots between lateral line and dorsal fin base, two blue stripes on cheek, yellowish green spot on each interspinous dorsal fin membrane, and dark green submarginal line on posterior section of dorsal fin. This species occurs in the tropical and subtropical Atlantic. In the western Atlantic it occurs between 10 and 128 m, from North Carolina and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is known throughout the Gulf of Mexico and is associated with coral reefs and hard substrates. Food consists of zooplankton (copepods, pelagic tunicates, and shrimps and shrimp larvae). Feeding takes place in the water column above reefs. This species is thought to be a protogynous hermaphrodite, but no specimens in transition from females to males have been observed. Maximum known size is 380 mm TL.

references: Smith 1971; Bright and Cashman 1974; Moore 1975a; Smith et al. 1975; Hoese and Moore 1977, 1998; Robins and Ray 1986; Bullock and Smith 1991; Cervigón 1991; Boschung 1992; Heemstra and Randall 1993; Randall 1996; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 173

Plectranthias garrupellus Apricot bass

Robins and Stark, 1961

Plectranthias garrupellus has spinous section of dorsal fin angular and lateral line running close to dorsal fin base. Anterior and posterior nares are close set, with anterior naris at end of short tube. Orbital diameter is greater than interorbital width. Supramaxilla is rudimentary or absent. Teeth in jaws are sharp, narrow, conical, and arranged in bands, with anterior teeth on inner and outer rows caninelike. Teeth in vomer and palatine are small and arranged in chevron-shaped patch without posterior prolongation on vomer. Preoperculum is angular and has coarse serrations along posterior margin and two antrorse spines on lower margin. Margin of interoperculum is smooth and covered with scales. Three opercular spines are of about equal size. Gill rakers on first arch are rather short and slender and number 14 to 18, with 4 to 6 on upper limb and 10 to 13 on lower limb. Measurements are expressed as percent of SL: head length 40%– 44%, snout length 8%–11%, orbit diameter 10%–12% (for specimens 40 mm SL), and body depth 29%– 42%. Pectoral fin has 12 to 14 rays. Dorsal fin is notched between spinous and rayed sections and has 10 spines and 15 to 17 rays, with fleshy tabs extending from interspinal membranes of anterior spines. Anal fin has

7 rays. Caudal fin is nearly truncate to truncate. Scales are large and ctenoid and cover body and head, with exception of snout, maxilla, and most of lower jaw. Scales encircling caudal peduncle number 12 to 14. Lateral line is continuous and has 28 or 29 tubed scales. Vertebrae number 26. Color is red to orange dorsally and pale laterally and ventrally, with dark red spot at base of each scale. Cheek is largely dull yellowish orange, and lateral section of body has numerous dull yellow flecks. Iris of eye is mostly reddish yellow. Spinous section of dorsal fin is reddish orange; rayed sections of dorsal, anal, and caudal fins are yellow; and pelvic fin is largely pallid. This species occurs between 13 and 375 m in the western Atlantic from North Carolina and the Bahamas to the eastern and western Caribbean Sea, including the eastern Gulf of Mexico and Cuba. Food consists of shrimps, crabs, and possibly polychaetes. This species is probably a protogynous hermaphrodite. In the Gulf of Mexico spawning is thought to take place in the late summer to fall. Maximum known size is 70 mm SL. references: Robins and Stark 1961; Randall 1980; Robins and Ray 1986; Heemstra 2002c.

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174 PERCIFORMES

Pronotogrammus martinicensis Roughtongue bass

(Guichenot, 1868)

Pronotogrammus martinicensis has scales on head and maxilla and an oval tooth patch on tongue. Anterior and posterior nares are close set, and posterior margin of anterior naris is produced into slender filament. Orbit diameter is greater than interorbital width. Upper jaw is indented at symphysis. Upper jaw has outer row of small conical teeth and narrow inner band of villiform teeth, with two large posteriorly or outwardly directed canine teeth near symphysis. Lower jaw has row of small conical teeth, narrow band of villiform teeth, and three large canine teeth directed either posteriorly or outwardly near symphysis. Vomerine and palatine teeth are granular, and vomerine tooth patch has well-developed posterior prolongation. Preoperculum is angular, with serrae on ventral and posterior margins. Suboperculum and interoperculum are serrated along free margins near their articulation. Gill rakers on first arch are long and slender and number 34 to 41, with 9 to 13 on upper limb and 24 to 29 on lower limb. Measurements are expressed as percent of SL: head length 31%– 37%, snout length 5%–7%, orbit length 9%– 13%, interorbital width 6%– 8%, body depth 33%– 42%. Pectoral fin is large and has 16 to 18 rays. Dorsal fin is not incised between

spinous and rayed sections and has 10 spines and 13 to 16 (usually 15) rays, with third spine longest. Anal fin has 7 or 8 rays. Caudal fin is crescent shaped and has filaments extending from upper and lower lobes. Scales are large and ctenoid and cover body and head. Scales encircling caudal peduncle number 18 to 22. Lateral line runs high along flank, is continuous, and has 35 to 41 (usually 38 to 40) tubed scales. Vertebrae number 25 or 26. Color is red orange to pink and slightly lighter ventrally than dorsally, with yellow or greenish yellow stripe running from snout below eye to operculum. Iris of eye is red orange, orange, yellow, or grayish green. Fins are yellow to orange. Some specimens may have two yellow stripes radiating posteriorly from orbit and/or golden or yellow stripes and yellow green bars on body. This species occurs between 55 and 230 m in the western Atlantic from North Carolina and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico. Food consists of ostracods, pteropods, and copepods. This species is a protogynous hermaphrodite. Females range from 47 to 112 mm SL, and males range from 66

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SERRANIDAE 175

to 132 mm SL. Maximum known size is about 150 mm SL. references: Longley and Hildebrand 1941 (as Ocyanthias martinicensis); Anderson and Heem-

stra 1980 (as Holanthias martinicensis); Coleman 1981 (as H. martinicensis); Matsuura 1983a (as O. martinicensis); Robins and Ray 1986; Dennis and Bright 1988a; Bullock and Smith 1991; Cervigón 1991; Boschung 1992 (all as H. martinicensis); Anderson 2002b.

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176 PERCIFORMES

Rypticus bistrispinus Freckled soapfish

(Mitchill, 1818)

Rypticus bistrispinus has rayed sections of dorsal and anal fins fleshy, inner pelvic rays connected to body by membrane, and closely spaced ocher or dark brown spots on body. Head is compressed and relatively long. Anterior naris is tubular and moderately separated from posterior naris. Maxilla extends to posterior margin of orbit. Jaw teeth are villiform, with those in inner row of lower jaw slightly enlarged and directed posteriorly. Vomer and palatine have bands of villiform teeth, with those of inner row of palatine slightly enlarged and directed posteriorly. Margin of preoperculum is semicircular, with three (occasionally two) spines. Gill rakers on first arch are short and number seven to nine, excluding rudiments. Measurements are expressed as percent of SL: head length 32%– 42%, snout length 7%–10%, eye diameter 7%–9%, upper jaw length 15%–20%, pectoral fin length 21%–25%, pre-dorsal-fin length 38%–50%, body depth 29%–34%. Pectoral fin is rounded and has 13 to 16 rays. Dorsal fin has rounded posterior margin and consists of two spines and 24 or 26 rays. Anal fin has rounded posterior margin and 15 to 17 (usually 15 or 16) rays; spines are lacking. Caudal fin is rounded. Scales are small and embedded. Head, with

exception of lips, is covered with scales. Skin is covered with thick mucus that contains toxin (grammistin). Vertebrae number 25: 10 precaudal and 15 caudal. Pyloric caecae number three. Color is yellow to light brown, with ocher to reddish brown spots on upper surface and cream to tan belly. Cheeks, upper and lower jaws, and belly are mottled with ocher to reddish brown spots. Juveniles have broad brown stripe on flank. This species occurs from near shore to at least 80 m in the western Atlantic from Cape Canaveral, FL, and the Bahamas to Venezuela, including the eastern and southern Gulf of Mexico and the Antilles. It is secretive and apparently remains in shells and burrows during the day and forages at night. Food consists of small amphipods, crabs, and shrimps. In the eastern Gulf of Mexico spawning takes place between July and September. Maximum known size is 115 mm SL.

references: Longley and Hildebrand 1941 (in part as R. arenatus); Courtenay 1967; Böhlke and Chaplin 1968; Darcy and Gutherz 1984; Williams and Shipp 1980; Robins and Ray 1986; Boschung 1992; Cervigón 1991; Smith 1997; Heemstra 2002c.

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SERRANIDAE 177

Rypticus maculatus (Holbrook, 1855) Whitespotted soapfish Rypticus maculatus has rayed sections of dorsal and anal fins fleshy, inner pelvic rays connected to body by membrane, and pale spots on body. Anterior naris is tubular and moderately separated from posterior naris. Maxilla extends beyond posterior margin of orbit. Jaws, vomerine, and palatine teeth are villiform. Margin of preoperculum is semicircular, with two (occasionally one) spines. Dorsal margin of operculum is connected to head by membrane. Gill rakers on first arch are short and number seven to nine, excluding rudiments. Measurements are expressed as percent of SL: head length 32%–38%, snout length 6%–9%, eye diameter 6%–7%, upper jaw length 13%–18%, pectoral fin length 17%– 22%, pre-dorsal-fin length 39%– 44%, body depth 29%– 40%. Pectoral fin is rounded and has 13 to 16 rays. Dorsal fin has two (rarely three) spines and 22 to 27 (usually 24 to 26) rays. Anal fin has rounded posterior margin and 13 to 17 (usually 15 or 16) rays; spines are lacking. Caudal fin is rounded. Scales are small and embedded. Head, with exception of lips, is

covered with scales. Skin is covered with thick mucus that contains toxin (grammistin). Vertebrae number 24 or 25: 10 precaudal and 14 or (rarely) 15 caudal. Color is brown, with scattered to coalesced white spots on upper half of head trunk, and cream on jaws and belly. This species occurs from near shore to 91 m in the western Atlantic from Cape Hatteras, NC, to the southern Gulf of Mexico. In the Gulf of Mexico it is known from Florida to Texas and from Campeche Bay, and it is associated with rocky or coral substrata. Food consists of crabs, shrimps, and small ray-finned fishes. In the Gulf of Mexico spawning takes place during the summer. Maximum known size is 198 mm SL.

references: Courtenay 1967; Springer and Woodburn 1960 (as R. saponaceus); Smith et al. 1975; Hastings et al. 1976; Hoese and Moore 1977, 1998; Hastings 1979; Robins and Ray 1986; Retzer 1990; Boschung 1992; Smith 1997; Heemstra 2002c.

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178 PERCIFORMES

Rypticus randalli

Courtenay, 1967

Rypticus randalli has rayed sections of dorsal and anal fins fleshy, inner pelvic rays connected to body by a membrane, and a uniformly dark-colored body. Anterior naris is tubular and moderately separated from posterior naris. Maxilla extends beyond posterior margin of orbit. Jaw, vomer, and palatine teeth are villiform and about same size. Preoperculum is semicircular, with two (occasionally one) spine. Dorsal margin of operculum is joined to head by membrane. Gill rakers on first arch are short and number 7 to 11, excluding rudiments. Measurements are expressed as percent of SL: head length 30%–34%, snout length 5%– 6%, upper jaw length 13%–16%, predorsal-fin length 36%– 42%. Pectoral fin is rounded and has 14 to 16 rays. Dorsal fin has rounded posterior margin, three spines, and 23 to 25 rays. Anal fin has rounded posterior

margin and 15 or 16 (rarely 14) rays; spines are lacking. Caudal fin is rounded. Scales are small and embedded. Head, with exception of lips, is scaled. Skin is covered with thick mucus that contains toxin (grammistin). Vertebrae number 24: 10 precaudal and 14 caudal. Pyloric caecae number three. Color is dark brown. This species occurs in shallow water on muddy bottoms in the western Atlantic from the southern Gulf of Mexico and from Honduras to Brazil. It has been recorded only once from the southern Gulf of Mexico, at Isla de Lobos, Veracruz. Maximum known size is 138 mm SL.

references: Courtenay 1967; Castro-Aguirre and Márquez-Espinoza 1981; Heemstra 2002c.

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SERRANIDAE 179

Rypticus saponaceus Greater soapfish

(Bloch and Schneider, 1801)

Rypticus saponaceus has rayed sections of dorsal and anal fins fleshy, inner pelvic fin rays connected to body by membrane, and pale blotches on body. Anterior naris is tubular and moderately separated from posterior naris. Maxilla extends beyond posterior margin of orbit. Jaws, vomerine, and palatine teeth are villiform, with those in posterior aspect of upper jaw, lower jaw, and vomer enlarged and directed posteriorly. Margin of preoperculum is semicircular, with two (rarely one or three) spines. Gill rakers on first arch are short and number 5 to 11 (usually 7 to 9), excluding rudiments. Measurements are expressed as percent of SL: head length 32%–39%, snout length 6%–9%, upper jaw length 13%–17%, pre-dorsal-fin length 34%– 46%. Pectoral fin is rounded and has 14 to 17 (usually 15 or 16) rays. Dorsal fin has rounded posterior margin and consists of three spines and 21 to 25 rays (usually 23 or 24). Anal fin has rounded posterior margin and 14 to 17 (usually 16 or 17) rays; spines are lacking. Caudal fin is rounded. Scales are small and embedded. Head, with exception of lips, is covered with scales. Skin is

covered with thick mucus that contains toxin (grammistin). Vertebrae number 24: 10 precaudal and 14 caudal. Pyloric caecae number four. Color is dark brown to blue black, with blotches of gray to brown on sides. Lower jaw and belly are cream, with dark mottling. Juveniles are pale gray, with dark markings that often form reticulations. This species occurs in the tropical Atlantic. In the western Atlantic it occurs from near shore to 82 m, from Miami, FL, and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it is known from banks off Texas and at Isla de Lobos, Veracruz. It is associated with muddy bottoms and coral reefs and is generally secretive during daylight. Maximum known size is 245 mm SL.

references: Courtenay 1967; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1998; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Dennis and Bright 1988b; Retzer 1990; Cervigón 1991; Smith 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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180 PERCIFORMES

Rypticus subbifrenatus Spotted soapfish

Gill, 1861

Rypticus subbifrenatus has rayed sections of dorsal and anal fins fleshy, inner pelvic rays connected to body by membrane, and small dark spots on body. Anterior naris is tubular and moderately separated from posterior naris. Maxilla extends beyond posterior margin of orbit. Jaw, vomerine, and palatine teeth are villiform, with posterior teeth in jaws and vomer slightly enlarged. Margin of preoperculum is semicircular, with one or two spines. Gill rakers on first arch are short and number 7 to 10, excluding rudiments. Measurements are expressed as percent of SL: head length 36%– 42%, snout length 6%–10%, upper jaw length 16%–20%, pre-dorsal-fin length 40%–52%. Pectoral fin is rounded and has 14 to 16 (usually 14 or 15) rays. Dorsal fin has rounded posterior margin and consists of three or four spines and 20 to 24 rays. Anal fin has rounded posterior margin and 13 to 16 (usually 14 or 15) rays; spines are lacking. Caudal fin is rounded. Scales are small and embedded. Head, with exception of lips, is covered with

scales. Skin is covered with thick mucus that contains toxin (grammistin). Vertebrae number 24: 10 precaudal and 14 caudal. Pyloric caecae number three. Color is olive green to reddish brown, with widely spaced dark brown to black spots. Spots are limited to head and anterior part of body in adults. This species occurs in shallow water in the western Atlantic from southern Florida and Bermuda to Venezuela, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it is known from Texas and Veracruz and is associated with coral reefs and other areas with clear water. Maximum known size is 105 mm SL.

references: Courtenay 1967; Böhlke and Chaplin 1968; Hoese and Moore 1977, 1998; CastroAguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Retzer 1990; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 181

Schultzea beta School bass

(Hildebrand, 1940)

Schultzea beta is elongate, with a very protrusible mouth that lacks teeth. Anterior and posterior nares are moderately separated, with posterior margin of anterior naris bearing small flap. Maxilla extends to middle of eye. Preoperculum is angular and serrated along ventral and posterior margins, with serrae at corner slightly enlarged. Margins of suboperculum and interoperculum are serrate. Opercular spines are closely spaced. Branchiostegal rays number 6. Gill rakers on first arch are long and number 29 to 39, with 9 to 11 on upper limb and 20 to 26 on lower limb. Measurements are expressed as percent of SL: head length 30%–33%, snout length 7%– 8%, eye diameter 9%–11%, body depth 21%–24%. Pectoral fin is truncate and has 15 to 17 rays. Dorsal fin is distinctly notched and has 10 spines and 11 to 13 rays. Anal fin has 6 to 8 rays. Caudal fin is shallowly forked, with upper lobe slightly longer than lower lobe. Scales are ctenoid and extend anteriorly to posterior margin of head. Scales encircling caudal peduncle number 25 to 30. Pored lateral line scales number 48 to 56. Vertebrae number 24:

10 precaudal and 14 caudal. Color is reddish brown dorsally with mottling of dark reddish brown, and irregular white blotches with reddish brown mottling ventrally. Rusty bar runs under eye, and dusky crescent marking is present along midlength of caudal fin lobes. This species occurs between 21 and 170 m in the western Atlantic from the Carolinas and the Bahamas to northern South America, including the eastern and southern Gulf of Mexico and the Antilles. It is often observed in small schools within a meter of the bottom near coral patches. Food consists of zooplankton, such as copepods, ostracods, crab megalops, and bivalve and gastropod veligers. It is thought to be a simultaneous hermaphrodite. Maximum known size is about 100 mm SL.

references: Longley and Hildebrand 1940, 1941 (as Serranus beta); Woods 1958 (as Schultzea campechanus); Robins and Stark 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Davis and Birdsong 1973; Darcy and Gutherz 1984; Robins and Ray 1986; Johnson and Smith-Vaniz 1987; Heemstra 2002c.

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182 PERCIFORMES

Serraniculus pumilio Ginsburg, 1952 Pygmy seabass

Serraniculus pumilio is spindle shaped, with head and body mottled. Anterior and posterior nares are close set, with large flap extending from posterior margin of anterior naris. Maxilla extends to near center of eye. Jaw teeth are villiform and arranged in broad bands, with those in outer row enlarged in both jaws and those of inner row enlarged in lower jaw. Vomer and palatine have villiform teeth arranged in bands. Preoperculum is serrated along posterior margin and smooth along ventral margin. Branchiostegal rays number 6. Gill rakers on first arch are short and number 9 to 13. Measurements are expressed as percent of SL: head length 31%–36%, snout length 7%– 8%, eye diameter 7%–9%, body depth 24%–28%. Pectoral fins have 14 or 15 rays. Dorsal fin is slightly notched, with 10 spines and 10 or 11 rays. Anal fin has 6 or 7 rays. Posterior margin of caudal fin is slightly convex. Scales are ctenoid and extend onto head to near posterior margin of orbit. Lateral line follows dorsal contour of body but is widely separated from dorsal fin base and has 40 to 46 pored scales.

Color is mottled with buff and dark brown dorsally, and is abruptly white ventrally, with poorly defined dark bars below spinous section of dorsal fin, near end of rayed section of dorsal fin, and at base of caudal peduncle. Narrow dark bar extends between eyes, and a row of brown dots is located along lateral line. This species occurs from near shore to 165 m in the western Atlantic from North Carolina to Venezuela, including the Gulf of Mexico and the Greater Antilles. In the Gulf of Mexico it is most abundant between 19 and 70 m over sandy to shelly bottoms. Food consists of crabs and shrimps. Spawning in the Gulf of Mexico takes place in August and September. Maturity is reached at about 25 mm SL, and maximum known size is 63 mm SL.

references: Ginsburg 1952a; Reid 1954; Springer and Woodburn 1960; Hastings 1973; Hoese and Moore 1977, 1998; Ogren and Brusher 1977; Darcy and Gutherz 1984; Robins and Ray 1986; Bullock and Smith 1991; Boschung 1992; Cervigón 1991; Heemstra 2002c.

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SERRANIDAE 183

Serranus annularis (Günther, 1880) Orangeback bass

Serranus annularis has two yellow squares outlined in black behind eye and a series of black spots on upper flank. Anterior and posterior nares are close set, with flap extending from posterior margin of anterior naris. Maxilla extends to about middle of pupil. Jaw teeth are villiform and in bands that decrease in width posteriorly, with several large teeth in inner and outer rows of anterior section of upper jaw and midsection of lower jaw. Vomerine teeth are small and in V-shaped patch, and palatine teeth are small and in short patch. Margin of preoperculum is finely serrate. Margin of interoperculum is smooth. Middle opercular spine is straight. Gill rakers on first arch are moderately short and number 15 to 18. Measurements are expressed as percent of SL: head length 38%– 40%, snout length 10%–12%, eye diameter 9%–11%, body depth 30%– 33%. Pectoral fin has 13 or 14 rays. Dorsal fin is not notched and has 10 spines and 10 to 12 rays. Anal fin has 6 or 7 rays. Caudal fin has concave margin. Scales are ctenoid and occur

on body, nape, operculum, and cheek but not on top of head. Cheek scale rows number 10 to 12. Scales encircling caudal peduncle number 31 to 36. Pored lateral line scales number 46 to 50. Color is orange or salmon dorsally, shading to white on side, with two squarish orange or yellow spots outlined in black and located on head behind eye, row of black spots along base of dorsal fin, and seven pale orangish yellow bars on lower side. Dorsal fin spines have blue tips. This species occurs in the western Atlantic from southern Florida and Bermuda to Brazil, including the eastern and northern Gulf of Mexico and the Antilles. It is common in rocky and coral reef habitats between 30 and 60 m. Maximum known size is 63 mm SL. references: Robins and Stark 1961; Hoese and Moore 1977, 1998; Robins and Ray 1986; Dennis and Bright 1988a; Bullock and Smith 1991; Randall 1996; Smith-Vaniz et al. 1999; Heemstra 2002c.

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184 PERCIFORMES

Serranus atrobranchus Blackear bass

(Cuvier, 1829)

Serranus atrobranchus has weak spines on operculum, scales on head, and black lanceolate mark on inner side of operculum. Maxilla extends to about anterior margin of pupil. Nares are close set, with moderate-sized flap extending from posterior margin of anterior naris. Jaw teeth are villiform and arranged in bands narrowing posteriorly (to a single row in lower jaw), with enlarged conical and recurved teeth in outer row of both jaws. Vomerine and palatine teeth are granular and arranged in Vshaped patch and narrow band, respectively. Preoperculum forms obtuse angle and has serrated margin, with serrae largest at corner. Suboperculum has smooth margin. Operculum has three weak spines, with upper and lower ones hidden by scales. Gill rakers on first arch number 15 to 20. Measurements are expressed as percent of SL: head length 35%–38%, snout length 8%–10%, eye diameter 11%–13%, body depth 33%–35%. Pectoral fin is long and has 15 to 17 rays. Dorsal fin is unnotched and has 10 spines and 12 rays. Anal fin has 7 rays. Caudal fin is emarginate to slightly concave.

Scales are weakly ctenoid, rather deciduous, and occur on nape, occiput, posterior aspect of interorbital region, operculum, and cheek. Scales encircling caudal peduncle number 22 to 25. Pored lateral line scales number 44 to 47, and rows of cheek scales number five. Color is brown dorsally and light tan ventrally, with black lanceolate mark on inner surface of operculum, visible externally, and faint dusky bar from base of eighth or ninth dorsal fin spine to belly. This species occurs between 18 and 276 m in the western Atlantic from the northern Gulf of Mexico to northern Brazil, including the Antilles. It is found throughout the Gulf of Mexico. Maximum known size is 90 mm SL.

references: Longley and Hildebrand 1941 (as Prionodes atrobranchus); Ginsburg 1952a (as Paracentropristis pomospilus and P. atrobranchus); Hildebrand 1954; Robins and Stark 1961; Hoese and Moore, 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; CastroAguirre et al. 1999; Heemstra 2002c.

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SERRANIDAE 185

Serranus baldwini Lantern bass

(Evermann and Marsh, 1899)

Serranus baldwini has well-developed opercular spines, top of head naked, and caudal fin bearing four dark blotches at base. Nares are close set, with flap extending from posterior margin of anterior naris. Maxilla extends to center of eye. Jaw teeth are villiform and in bands that narrow posteriorly, with additional canine teeth occurring on either side of symphysis of upper jaw and in lateral aspect of lower jaw. Vomerine and palatine villiform teeth are arranged in V-shaped patch and short band, respectively. Preoperculum forms obtuse angle and has serrations along posterior margin. Middle opercular spine is relatively large and straight. Gill rakers on first arch are short and number six or seven. Measurements are expressed as percent of SL: head length 38%– 41%, snout length 8%–10%, eye diameter 9%–11%, body depth 28%–32%. Pectoral fin is large and has 13 to 15 rays. Dorsal fin is unnotched and has 10 (rarely 9 or 11) spines and 11 or 12 rays. Anal fin has 7 rays. Caudal fin is truncate. Scales are ctenoid and cover body, nape, opercula, and cheek. Pored lateral

line scales number 42 to 48. Rows of cheek scales number six. Scales encircling caudal peduncle number 23 to 26. Color is white with three stripes or series of blotches on upper side and series of four black blotches, each with red or yellow bar below it, along lower side of body. Bars along lower flank are red in shallow-water specimens and yellow in deepwater ones. Caudal fin base has four black spots in curved vertical series. This species occurs between the shore and 75 m in the western Atlantic from southeastern Florida and the Bahamas to Suriname, including the eastern Gulf of Mexico and the Antilles. It is associated with Thalassia beds and areas with vertical relief. Juveniles have been found in queen conchs. Food consists of caridean shrimps and small ray-finned fishes such as labrids. Maximum known size is 51 mm SL.

references: Robins and Stark 1961; Longley and Hildebrand 1940, 1941; Robins and Ray 1986; Randall 1996; Smith 1997; Heemstra 2002c.

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186 PERCIFORMES

Serranus chionaraia Robins and Stark, 1961 Snow bass Serranus chionaraia has well-developed opercular spines, top of head naked, and caudal fin bearing dark hatch marks along upper and lower rays. Nares are close set, with flap extending from posterior margin of anterior naris. Maxilla extends to center of eye. Jaw teeth are villiform and in bands that narrow posteriorly, with additional canine teeth occurring on either side of symphysis of upper jaw and in lateral aspect of lower jaw. Villiform vomerine and palatine teeth are arranged in V-shaped patch and short band, respectively. Preoperculum forms obtuse angle and has minute serrations along posterior margin. Middle opercular spine is relatively large and straight. Gill rakers on first arch are short and number 17 to 20. Measurements are expressed as percent of SL: head 40%– 41%, snout length 8%– 11%, eye diameter 11%–13%, body depth 31%–36%. Pectoral fin is large and has 13 or 14 rays. Dorsal fin is unnotched and has

10 spines and 12 rays. Anal fin has 7 rays. Caudal fin is very shallowly forked. Scales are ctenoid and cover body, nape, opercula, and cheek. Scales encircling caudal peduncle number 26 to 30. Pored lateral line scales number 45 to 47. Color is brown, with belly white from pectoral fin base to anus and caudal peduncle cream colored or pale tan. Rayed sections of dorsal and anal fins have brown spots. Head has three parallel blue stripes, and caudal fin has brown to blackish crossbars or series of spots on upper and lower margins. This species occurs between 45 and 90 m in the western Atlantic from southeastern Florida, the Florida Keys, Puerto Rico, and Honduras. It has not been reported from the Gulf of Mexico, but it may occur there. Maximum known size is 49 mm SL. references: Robins and Stark 1961; Robins and Ray 1986; Smith 1997; Heemstra 2002c.

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SERRANIDAE 187

Serranus luciopercanus

Poey, 1852

Serranus luciopercanus has a notch between spinous and rayed sections of dorsal fin, ctenoid scales, and middle opercular spine turned upward. Anterior profile of head is slightly convex. Nares are close set, with flap extending from posterior margin of anterior naris. Maxilla extends to anterior margin of pupil. Jaw teeth are villiform and in bands narrowing posteriorly, with outer row enlarged, conical, and recurved. Vomerine tooth patch is chevron shaped, with teeth small except for canine teeth at each end of V. Palatine teeth are in irregular rows. Premaxilla is obtusely angled and has serrae along posterior and ventral margins, with serrae best developed at corner. Gill rakers on first arch number 20 to 24. Measurements are expressed as percent of SL: head length 40%– 41%, snout length 12%–14%, eye diameter 10%–11%, body depth 29%– 32%. Pectoral fin is large and fanlike and has 14 rays. Dorsal fin has 10 spines and 11 or 12 rays. Anal fin is slightly falcate and has 7 rays. Caudal fin is deeply forked, and upper lobe is

longer than lower lobe. Scales are ctenoid and occur on nape, operculum, cheek, pectoral fin base, and bases of dorsal and anal fins but not on top of head. Scales encircling caudal peduncle number 38 to 42. Pored lateral line scales number 50 to 55, and rows of cheek scales number 8 to 10. Color is brown dorsally and tan ventrally, with broad dark band extending diagonally from base of spinous section of dorsal fin to level of pectoral fin. Caudal fin has dark upper and lower bands that meet at base of fin and form horizontal V-shaped pattern. Upper and lower margins of caudal fin are dark. This species occurs at about 110 m in the western Atlantic from the southern Gulf of Mexico (off Veracruz), Cuba, Grand Cayman, the Lesser Antilles, and Honduras. Maximum known size is 119 mm SL. references: Jordan and Evermann 1898; Robins and Stark 1961; Castro-Aguirre and MárquezEspinoza 1981; Heemstra 2002c.

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188 PERCIFORMES

Serranus notospilus Saddle bass

Longley, 1935

Serranus notospilus is spindle shaped in profile and moderately compressed, with no notch between spinous and rayed sections of dorsal fin, deciduous scales, and black blotch on rayed section of dorsal fin. Nares are close set, with long flap extending from posterior margin of anterior naris. Maxilla extends to about middle of eye. Jaw teeth are villiform and in bands narrowing posteriorly, with outer row enlarged, conical, and recurved. Granular vomerine and palatine teeth are arranged in chevron-shaped patch and narrow band, respectively. Premaxilla is obtusely angled and has serrae along posterior and ventral margins. Middle opercular spine is relatively large and straight. Gill rakers on first arch number 17 to 23. Measurements are expressed as percent of SL: head length 36%– 40%, snout length 7%–11%, eye diameter 11%–15%, body depth 26%–33%. Pectoral fin is large and fanlike and has 14 to 17 (usually 15 or 16) rays. Dorsal fin has 10 spines and 12 or 13 (usually 12) rays. Anal fin has 7 or 8 rays. Caudal fin is emarginate. Scales are weakly ctenoid and occur on nape, occiput, interorbital region, operculum, and

cheek. Pored lateral line scales number 45 to 48. Rows of cheek scales number five to seven, and scales encircling caudal peduncle number 25 to 29. Color is brown to tan dorsally and tan ventrally, with faint dark bands below dorsal fin and on caudal peduncle. Head is silvery, with broad dark brown band running from eye to interoperculum. Rayed section of dorsal fin has black blotch above middle band. Specimens from the southern section of the range (Caribbean Sea) have broad white bar extending from lateral line to belly, just anterior to anus. This species occurs between 44 and 229 m in the western Atlantic from the Florida Keys to Suriname, including the Florida Keys and the Gulf of Mexico. It occurs throughout the Gulf of Mexico. Maximum known size is about 100 mm SL. references: Longley 1935; Longley and Hildebrand 1941 (as Prionodes notospilus); Ginsburg 1952a (as Paracentropristis notospilus); Robins and Stark 1961; Bullock and Smith 1991; Boschung 1992; Smith 1997; Heemstra 2002c.

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SERRANIDAE 189

Serranus phoebe Tattler

Poey, 1851

Serranus phoebe has well-developed opercular spines and vertical white band on body just anterior to anus. Nares are close set, with moderate-sized flap extending from posterior margin of anterior naris. Maxilla extends to about center of eye. Jaw teeth are villiform and arranged in rows that narrow posteriorly, with teeth in outer rows enlarged, conical, slightly recurved, and widely spaced. Granular vomerine and palatine teeth are arranged in chevronshaped patch and narrow row, respectively. Preoperculum forms obtuse angle and has serrae on posterior and ventral margins, with serrae largest on posterior margin near angle. Suboperculum has smooth margin. Middle opercular spine is relatively large and straight. Gill rakers on first arch number 17 to 20. Measurements are expressed as percent of SL: head length 37%– 41%, snout length 10%–12%, eye diameter 10%–12%, body depth 30%– 34%. Pectoral fin is large and usually has 15 or 16 rays. Dorsal fin is not notched between spinous and rayed sections and has 10 spines and 12 rays. Anal fin has 7 rays. Caudal fin is shallowly forked. Scales are ctenoid and cover trunk, nape, operculum, interoperculum, and cheek but not occiput or interorbital region. Scales encircling caudal peduncle number 29 to 31. Pored lateral line scales number 47 to 51, and rows on cheek number 9 to 12. Basal third

of interspinous membranes of dorsal and anal fins are covered with scales. Color is buff to brown dorsally and white to light tan ventrally. Broad dark brown bars occur on head below orbit, below anterior section of spinous dorsal fin to belly just in front of anus, and under center of rayed section of dorsal fin. Bar under spinous dorsal fin is immediately followed by silvery white bar extending from midflank to anus. Two poorly defined dark areas occur on rear half of body, one near dorsal fin and one at midline of body. Spinous section of dorsal fin has dusky blotch above band. Pigment patterns are best developed in juvenile specimens and may be indistinct, with exception of silvery white band in adults. This species occurs between 146 and 402 m in the western Atlantic from South Carolina and Bermuda to Guiana, including the Gulf of Mexico and the Greater Antilles. It is more common in the northeastern than in the northwestern Gulf of Mexico. Maximum known size is 154 mm SL. references: Robins and Stark 1961; Moore 1975a; Hoese and Moore 1977, 1998; Matsuura 1983a; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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190 PERCIFORMES

Serranus subligarius Belted sandfish

(Cope, 1870)

Serranus subligarius has well-developed opercular spines and a sharply defined white belly. Dorsal profile of head is strongly sloping and straight to slightly concave. Nares are close set, with large flap extending from posterior margin of anterior naris. Maxilla extends to about center of eye. Jaw teeth are villiform and arranged in bands that narrow posteriorly, with those in outer row enlarged and slightly recurved. Vomerine teeth are arranged in triangular patch, and palatine teeth form narrow band. Preoperculum is rounded at corner and has serrae on margin. Margin of interoperculum is smooth. Middle opercular spine is straight. Gill rakers on first arch are short and number 15 to 19. Measurements are expressed as percent of SL: head length 40%– 44%, snout length 10%–11%, eye diameter 9%–10%, body depth 34%–39%. Pectoral fin is large and has 15 to 17 rays. Dorsal fin has 10 spines and 12 to 14 rays. Anal fin has 7 rays, and second spine is longer and thicker than third spine. Caudal fin is truncate, with rounded corners. Scales are ctenoid and adherent and occur on trunk, nape, operculum, and cheek but not on occiput or interorbital region. Interradial membrane of dorsal and anal fins

is scaled along basal section. Scales encircling caudal peduncle number 24 to 28. Pored lateral line scales number 42 to 46, and scale rows on cheek number 10 to 12. Color is reddish brown, with series of broad dark bands extending to ventral midline except on sharply defined white belly. Center of each scale is slightly lighter than margin. Large black blotch is located on anterior portion of rayed section of dorsal fin and flank below. Rayed sections of dorsal and anal fins and pectoral and caudal fins are finely banded. This species occurs between 2 and 80 m in the western Atlantic from North Carolina to the northern (and western) Gulf of Mexico. In the Gulf of Mexico it occurs from southern Florida to Vera Cruz, but it is apparently rare in the Florida Keys and clear waters of southern Florida. Maximum known size is 100 mm TL.

references: Ginsburg 1952a (as Serranellus subligarius); Robins and Stark 1961; Hoese and Moore 1977, 1998; Hastings and Bortone 1980; Robins and Ray 1986; Bullock and Smith 1991; Boschung 1992; Schaldach et al. 1997; Smith 1997; Heemstra 2002c.

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SERRANIDAE 191

Serranus tabacarius (Cuvier, 1829) Tobaccofish

Serranus tabacarius has well-developed opercular spines, top of head naked, and upper and lower sections of caudal fin darkly pigmented. Dorsal profile of head is slightly convex. Nares are close set, with flap extending from posterior margin of anterior naris. Maxilla extends to about center of eye. Jaw teeth are villiform and in bands that narrow posteriorly. Granular vomerine and palatine teeth are arranged in chevron-shaped patch and narrow band, respectively. Preoperculum forms obtuse angle and is serrated along posterior and ventral margins, with serrae enlarged along lower posterior margin. Interoperculum has serrated margin. Gill rakers on first arch number 21 to 25. Measurements are expressed as percent of SL: head length 36%–39%, snout length 9%–11%, eye diameter 9%–12%, body depth 27%–30%. Pectoral fin is large and has 14 or 15 rays. Dorsal fin is shallowly notched and has 10 spines and 11 or 12 rays. Anal fin has 7 rays. Scales are small and ctenoid and cover body, nape, cheek, and operculum. Scales encircling caudal peduncle number 37 to 39.

Pored lateral line scales number 50 to 52, and rows of cheek scales number 10 to 12. Color is pale brownish orange, with pale blotches and black saddles or bars on dorsal part of body. Caudal fin is marked with two dark submarginal black stripes that converge at base to form horizontal V-shaped pattern. This species occurs between the shore and 69 m in the western Atlantic from Georgia and Bermuda to northern Brazil, including the eastern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with sandy bottoms and reefs. Food consists of shrimps and atherinid fishes. This species is a synchronous hermaphrodite. Maximum known size is 140 mm SL.

references: Kanazawa 1952 (as Prionodes tabacarius); Robins and Stark 1961; Stark and Davis 1966; Böhlke and Chaplin 1968; Randall 1968a, 1996; Smith et al. 1975; Robins and Ray 1986; Bullock and Smith 1991; Smith 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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192 PERCIFORMES

Serranus tigrinus Harlequin bass

(Bloch, 1790)

Serranus tigrinus has well-developed opercular spines, top of head naked, and irregular black spots and bars on side of body. Anterior dorsal profile is slightly convex. Maxilla extends to center of eye. Nares are close set, with flap extending from posterior margin of anterior naris. Jaw teeth are villiform and in bands that narrow posteriorly, with additional canines on either side of symphysis of upper and lower jaws, and several canines near midlength of lower jaw. Vomer has V-shaped patch of villiform teeth. Palatine teeth are granular and in narrow band. Preoperculum is finely serrate on posterior margin and smooth on ventral margin. Interoperculum has smooth margin. Gill rakers on first arch number 15 to 19. Measurements are expressed as percent of SL: head length 38%– 42%, snout length 9%–12%, eye diameter 8%–12%, body depth 26%–31%. Pectoral fin is large and has 14 rays. Dorsal fin is unnotched and has 10 spines and 12 rays. Anal fin has 7 rays. Caudal fin is concave, with upper rays longer than lower rays. Scales are ctenoid and cover body, nape, operculum, and cheek. Scales encircling caudal peduncle number 36 to 41. Lateral line scales number 48 to 51, and cheek scale rows number 11 to 13.

Body is white, with irregular dark brown to black spots dorsally, and six or seven dark brown to black bars extending to ventral midline. Middle four bars run onto dorsal fin. Head is white, with irregular black spots dorsally and ventrally. Dorsal, anal, and caudal fins are densely covered with small dark brown spots. This species occurs between the shore and 37 m in the western Atlantic from southern Florida and Bermuda to Venezuela, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from the Florida Middle Grounds and the Yucatán Peninsula and is associated with reefs and rubble around reefs. Food consists of stomatopods, caridean shrimps, crabs, and ray-finned fishes. This species is a synchronous hermaphrodite. Pairs of adults spawn in a shared territory, with sexual roles often reversed. Maximum known size is about 75 mm SL. references: Robins and Stark 1961; Smith 1965; Böhlke and Chaplin 1968; Randall 1968a, 1996; Smith et al. 1975; Pressley 1981; Bullock and Smith 1991; Smith 1997; Smith-Vaniz et al. 1999; Heemstra 2002c.

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SERRANIDAE 193

Serranus tortugarum Longley, 1935 Chalk bass

Serranus tortugarum has well-developed opercular spines, top of head naked, and a number of pale bands on upper part of body. Dorsal profile of head is slightly convex. Nares are close set, with flap extending from posterior margin of anterior naris. Maxilla extends to about anterior margin of pupil. Jaw teeth are villiform and in bands that narrow posteriorly. Granular vomerine and palatine teeth are arranged in V-shaped patch and narrow band, respectively. Preoperculum forms obtuse angle and has serrae along posterior and ventral margins, with serrae largest on posterior margin near angle. Interoperculum has smooth margin. Gill rakers on first arch number 27 to 31. Measurements are expressed as percent of SL: head length 33%–36%, snout length 7%–9%, eye diameter 10%–13%, body depth 29%– 31%. Pectoral fin is large and has 14 or 15 rays. Dorsal fin is shallowly indented and has 10 spines and 10 to 12 rays. Anal fin has 7 rays. Scales are ctenoid and cover body, nape, operculum, interoperculum, and cheek. Scales encircling caudal peduncle number 30 to 34. Pored lateral line scales number 46 to 50, and rows of cheek scales number five or six. Color is orangish brown, shading to white ventrally, with seven white or pale blue bars above lat-

eral line (first on nape, last on caudal peduncle, and remainder extending onto dorsal fin) and three or four chalky bars below lateral line. Head has four dusky blue transverse bands: one on snout, two across interorbital region, and one on nape. Caudal fin is mottled with dull orange and pale blue. Pectoral fin is pale pink. This species occurs between 18 and 402 m in the western Atlantic from southern Florida and the Bahamas to Venezuela, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from the Dry Tortugas, southwestern Florida, and Campeche Bay. This species is a simultaneous hermaphrodite that may exchange both sperm and eggs with its partner during spawning. Spawning occurs at dusk among large aggregations of individuals. Maximum known size is 78 mm SL.

references: Longley 1935; Longley and Hildebrand 1940, 1941; Robins and Stark 1961; Stark and Courtenay 1962; Stark and Davis 1966; Randall 1968a; Böhlke and Chaplin 1968; Darcy and Gutherz 1984; Fischer 1984; Bullock and Smith 1991; Boschung 1992; Smith 1997; Heemstra 2002c.

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194

GRAMMATIDAE

Basslets

Basslets are moderately elongate and compressed, with large eyes, a continuous dorsal fin, and lateral line interrupted or absent on body. Dorsal profile of head is moderately convex. Snout is very short. Naris is paired, with anterior naris tubular and posterior naris elliptical. Mouth is terminal, oblique, and usually rather small to moderate in size. Upper jaw is moderately protrusible. Teeth in upper jaw are arranged in a villiform patch on each side of symphysis, are enlarged and conical lateral to patch, and arranged in one to several rows. Teeth in lower jaw are arranged in a villiform patch on either side of symphysis, and are enlarged, recurved, and caninelike lateral to patch. Vomerine and palatine teeth are slightly enlarged and conical. Supramaxilla is present or absent. Posterior and ventral margins of preoperculum are serrated or smooth. Operculum either lacks or has one or more small, flat spine(s) on upper posterior margin. Gill membranes are joined at level of orbit. Gill rakers on first arch are usually long and slender. Pectoral fin is moderate to large. Dorsal fin is unnotched or shallowly notched and has 11 to 13 spines. Pelvic fin inserts below or slightly anterior to pectoral fin base, is moderately long to filamentous, and has 1 spine and five rays. Anal fin has 3 spines. Caudal fin is strongly emarginate to rounded. Scales are ctenoid and cover body and head, including operculum and cheek. Maximum known size is about 100 mm TL. Basslets occur in the tropical western Atlantic in shallow water to a depth of 365 m and are generally associated with clear water around coral reefs. They are often found near ledges and dropoffs on deep reefs. There are 11 species in two genera in the western Central Atlantic, and 2 or 3 species in one or two genera occur in the Gulf of Mexico. references: Jordan and Evermann 1898; Böhlke and Chaplin 1968; Fritzsche and Fuiman 1982; Johnson 1984; Gill and Mooi 1993; Nelson 1994; Smith 1997; Smith-Vaniz et al. 1999; Mooi 2002a.

Key to the Species of the Gulf of Mexico (Adapted from Böhlke and Chaplin 1968) 1a. Lateral line on body divided into two sections: anterior section high on body and posterior section at midline; procurrent caudal fin rays thin and weak; body bicolored, purple anteriorly shading to yellow posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gramma loreto p. 196 1b. Lateral line on body absent; procurrent caudal fin rays consisting of three strong spines above and below principal caudal fin rays; body uniformly colored or bicolored, yellow anteriorly shading to blue posteriorly . . . . . . . . . . . . 2

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GRAMMATIDAE 195

2a. Caudal fin margin rounded; pectoral fin rays 17; lateral scale rows about 26; body uniformly bronze and lacking blue lines on head and anterior body . . . . . . . . . . . . . . . . . . . Lipogramma anabantoides p. 197 2b. Caudal fin margin emarginate; pectoral fin rays 15; lateral scale rows about 29 or 30; body yellow anteriorly and bluish posteriorly, with three blue lines on head and anterior body. . . . . . . . . . . . . . Lipogramma trilineatum p. 198

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196 PERCIFORMES

Gramma loreto Poey, 1868 Royal gramma

Gramma loreto has a disjunct lateral line and poorly developed procurrent caudal fin rays. Snout is blunt. Anterior naris is at end of welldeveloped tube and is widely separated from posterior naris. Lower jaw is even with upper jaw, and upper jaw extends to posterior margin of orbit. Upper jaw has an outer row of enlarged, slightly recurved teeth and an inner patch of villiform teeth on either side on symphysis. Lower jaw has one or two large recurved canine(s) and an inner patch of villiform teeth on either side of symphysis, and a row of enlarged canines running to corner of mouth. Vomer has small patch of slightly enlarged conical teeth, and palatine has narrow band of similar teeth. Preoperculum has broadly rounded and serrated posterior margin. Operculum has 2 flat spines on upper posterior margin, with lower spine larger than upper. Gill rakers on first arch number 12 or 13 on upper limb and 20 to 22 on lower limb. Measurements are expressed as percent of SL: head length 32%–35%, snout length 6%– 8%, upper jaw length 16%–18%, eye diameter 9%– 13%, pelvic fin length 31%–53%, body depth 29%–36%. Pectoral fin extends to near anus and has 15 or 16 rays. Dorsal fin is shallowly notched and has 12 spines and 9 or 10 rays. Anal fin has 9 or 10 rays. Caudal fin is slightly

emarginate to slightly rounded. Head lacks scales except for cheek and opercular flap. Anterior section of lateral line originates at upper end of operculum and runs parallel with base of dorsal fin, and posterior section of lateral line originates at about level of end of anterior section and runs along midlateral flank. Lateral line scales number 34 to 36 in anterior section and 10 to 12 in posterior section. Most scales on body are ctenoid. Vertebrae number 27 or 28: 10 precaudal and 17 or 18 caudal. Color is blue or purplish anteriorly and orangish yellow posteriorly, with two golden streaks on head and a black spot on anterior margin of dorsal fin. This species occurs between the shoreline and 65 m in the western Atlantic in the Bahamas, the Florida Keys, and the Greater and Lesser Antilles, and from Belize to Venezuela. It has not been reported from the Gulf of Mexico, but it may occur there. This species is associated with caves and crevices of reefs and is often seen upside down under ledges. Males incubate the eggs, but they are not mouth brooders. Maximum known size is 65 mm SL. references: Böhlke and Randall 1963; Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986; Smith 1997; Mooi 2002a.

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GRAMMATIDAE 197

Lipogramma anabantoides Dusky basslet

Böhlke, 1960

Lipogramma anabantoides has well-developed procurrent caudal fin rays and a rounded caudal fin and lacks a lateral line. Snout is acute. Anterior naris is at end of well-developed tube and is widely separated from posterior naris. Lower jaw is even with upper jaw, and upper jaw extends to about middle of orbit. Supramaxilla is absent. Upper jaw has a band of villiform teeth, with a few of outermost teeth near symphysis enlarged. Lower jaw has a band of villiform teeth, with four of outermost teeth enlarged, recurved, and progressively larger from front to back. Vomer and palatine have narrow patches of villiform teeth. Preoperculum has smooth posterior margin. Operculum lacks spines but has flat projection. Gill membranes are united beneath posterior section of eye to form free fold over isthmus. First gill arch has 11 small gill rakers on lower limb. Measurements are expressed as percent of SL: head length 40%– 42%, snout length 9%–10%, upper jaw length 13%–14%, eye diameter 13%– 14%, pelvic fin length 49%–52%, body depth 40%– 41%. Pectoral fin is short and rounded and has 17 rays. Dorsal fin is continuous and has 12 or 13 spines and 8 to 10 rays. Pelvic fin is long and filamentous. Anal fin has 7 to

9 rays. Head has cycloid scales on cheeks, on posterior section of interorbital region, and on opercular bones. Scales on body are finely ctenoid. Lateral scale series number about 26, and vertebrae number 25: 10 precaudal and 15 caudal. Lateral line pores on head are large. Color in life is brilliant rose red on head and burnt sienna on remainder of body, with light lines between darkly pigmented scale rows. Dorsal fin has black margin, three black spots between first 3 fin spines, and partial ocellus on posterior section of fin. Color in preservative is dark with a reddish cast, with 10 dark lines running length of body and small dark blotch on anterior section and large, dark oval blotch on posterior section of dorsal fin. Unpaired fins are dark. This species occurs between 10 and 79 m in the western Atlantic off southern Florida, the Bahamas, Quintana Roo, Belize, Honduras, and Haiti, and in the Gulf of Mexico off the southwest coast of Florida. Maximum known size is 26 mm TL. references: Böhlke 1960; Böhlke and Chaplin 1968; Gilmore and Jones 1988; Gilmore 1997; Smith 1997; Mooi 2002a.

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198 PERCIFORMES

Lipogramma trilineatum Threeline basslet

Randall, 1963

Lipogramma trilineatum has well-developed procurrent caudal fin rays and an emarginate caudal fin and lacks a lateral line. Snout is acute. Anterior naris is at end of welldeveloped tube and is well separated from posterior naris. Maxilla extends to about middle of orbit. Supramaxilla is absent. Upper jaw has a band of villiform teeth, with six outermost teeth near symphysis enlarged and slightly recurved. Lower jaw has a band of villiform teeth, with four outermost teeth enlarged and recurved. Vomer and palatine have narrow patches of villiform teeth. Preoperculum has a smooth margin. Operculum lacks spines but has flat process. Gill membranes are united beneath posterior section of eye to form free fold over isthmus. Measurements are expressed as percent of SL: head length 35%–37%, snout length 8%–9%, eye diameter 11%–14%, pelvic fin length 37%– 69%, body depth 33%–34. Pectoral fin is short and slightly pointed and has 15 rays. Dorsal fin is continuous and has 12 spines and 10 rays. Pelvic fin is long and

filamentous. Anal fin has 7 rays. Cycloid scales occur on head, nape, thorax, pectoral fin base, and dorsal fin base. Scales on remainder of body are finely ctenoid. Scales in horizontal series number about 29 or 30, and vertebrae number 25: 10 precaudal and 15 caudal. Color is bright yellow to orange anteriorly and dorsally, grading to bluish posteriorly and ventrally, with three bright blue stripes running from head to middle of back. Top of head is reddish, and cheeks are lavender. This species occurs between 22 and 100 m in the western Atlantic from southeast Florida, the Dry Tortugas, Quintana Roo, Belize, Honduras, the Grand Cayman Islands, Jamaica, Puerto Rico, and the Bahamas to Curaçao. In the Gulf of Mexico it is known only from the Dry Tortugas. Maximum known size is about 35 mm TL. references: Randall 1963c; Böhlke and Chaplin 1968; Robins and Ray 1986; Gilmore 1997; Smith 1997; Mooi 2002a.

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199

OPISTOGNATHIDAE

Jawfishes

Jawfishes are oblong to elongate, moderately compressed, and tapering posteriorly, with bulbous heads, large jaws, and an incomplete lateral line. Anterior profile of head is strongly convex. Eye is large and high on head. Mouth is terminal, horizontal, and very large. Upper jaw is protractile. Single supramaxilla is usually present. Jaw teeth are small, cardiform to conical, and arranged in a single row along sides of jaw, with several additional rows of small teeth on either side of symphyses. Small teeth usually occur in vomer, but palatine lacks teeth. Operculum lacks spines. Pseudobranch is present on inner surface of gill cover. Gill rakers on first arch are well developed, and slit is present behind fourth gill arch. Gill membranes are united but are free of isthmus. Branchiostegal rays number six. Pectoral fin is fan shaped. Dorsal fin is long, continuous, and unnotched to slightly notched between spinous and rayed sections, with first half consisting of 9 to 12 slender spines and second half consisting of 10 to 22 rays. Pelvic fin is jugular and consists of 1 spine and 5 rays, the outer 2 of which are unbranched and stout while the inner 3 are branched and weak. Anal fin is long, low, and consists of 2 or 3 slender spines and 10 to 20 rays. Caudal fin has a rounded or lanceolate posterior margin. Scales are small and cycloid and cover body but usually not head. Lateral line runs high on flank and terminates at about center of dorsal fin base; tubes or pores are usually embedded in skin rather than in scales. Maximum known size is about 500 mm TL, but species are usually less than 120 mm TL. Jawfishes occur worldwide in tropical to subtropical seas on sandy to muddy bottoms from less than a meter to 375 m. Most species occur in shallow water over sandy bottoms near coral reefs. All species dwell in vertical burrows or crevices and are active during daylight hours. Some live commensally with invertebrates. Food consists of benthic and free-swimming invertebrates. Males practice oral incubation. There are about 60 species in three genera worldwide, 17 species in two genera in the western Central Atlantic, and 7 to 9 species in two genera in the Gulf of Mexico. references: Böhlke and Chaplin 1968; Smith-Vaniz 1986b, 2001a, 2002b; Nelson 1994; Randall 1996; Smith 1997.

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200 PERCIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Smith-Vaniz 1997) 1a. Posterior end of maxilla straight or rounded; cheeks rarely scaled; caudal fin rounded . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Posterior end of maxilla weakly concave, strongly concave, or hooklike; cheeks scaled; caudal fin lanceolate . . . . . . . . . . . . . . . . . Lonchopisthus micrognathus p. 202 2a. Anterior naris raised into short tube and lacking cirrus .............................................3 2b. Anterior naris raised into short tube and possessing simple cirrus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 3a. Dorsal fin spines 10; vomerine teeth absent; inner aspect of upper jaw mostly black; gill rakers on first arch 26 to 32 .............................................4 3b. Dorsal fin spines usually 11; vomerine teeth usually present; inner aspect of upper jaw pale; gill rakers on first arch 34 to 62 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4a. Posterior end of maxilla extended as thin, flexible lamina; coranoid process of articular bone of lower jaw club shaped, with evenly convex dorsal margin . . . . . . . . . . . . . . . . . . . Opistognathus melachasme p. 207 4b. Posterior end of maxilla rigid, not extended as thin, flexible lamina; coranoid process of articular bone of lower jaw hatchet shaped, with straight dorsal margin . . . . . . . . . . . . . . . . . . . . . . . Opistognathus nothus p. 208 5a. Outermost pelvic fin ray very closely bound to adjacent ray, and membrane between rays not incised distally; dorsal fin with narrow dark margin; anal fin rays 14 to 17; caudal fin length 30% to 41% of SL . . . . . . . . . . . . . . . . . . . . . Opistognathus aurifrons p. 203 5b. Outermost pelvic fin ray not closely bound to adjacent ray, and membrane between rays incised distally; dorsal fin without narrow dark margin; anal fin rays 12 to 14 (usually 12 or 13); caudal fin length 19% to 30% of SL . . . . . . . . . . . . . . . . . . . . Opistognathus lonchurus p. 204 6a. Posterior end of maxilla in adults terminating as thin, flexible lamina (slightly elongated in females and greatly elongated in males); dorsal fin spines thin, flexible, and usually curved distally, tips without swollen fleshy tabs; dorsal fin rays 15 to 18; posterior mid-dorsal half of head, anterior to dorsal fin, with few sensory pores . . . . . . . . . . . . . . . . 7 6b. Posterior end of maxilla in adults rigid, not ending in thin, flexible lamina; dorsal fin spines stiff and straight, tips with swollen fleshy tabs; dorsal fin rays 13 to 16; posterior middorsal half of head, anterior to dorsal fin, densely covered with pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7a. Dorsal fin with five or six dusky bands, with some extend-

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OPISTOGNATHIDAE 201

ing above fin base; gill rakers on first arch 31 to 43 in specimens greater than 50 mm SL . . . . . . . . . . . . . . . . . Opistognathus macrognathus p. 205 7b. Dorsal fin generally without five or six dusky bands but, if present, bands limited to fin base; gill rakers on first arch 28 to 35 in specimens greater than 50 mm SL . . . . . . . . . . . . . . . . . . . . . . . Opistognathus robinsi p. 209 8a. Fleshy cirrus on anterior naris relatively slender; upper margin of suboperculum not forming broad, fanlike truncated flap; premaxilla with a single row of teeth anteriorly; supramaxilla present; 42 to 54 oblique scale rows on body; posteriormost premaxillary tooth of mature males stouter and more strongly hooked than adjacent teeth . . . . . . . . . . . . . . . . . . . Opistognathus whitehursti p. 210 8b. Fleshy cirrus on anterior naris rounded to palmate; upper margin of suboperculum forming broad, fanlike truncated flap; premaxilla with two or more rows of teeth anteriorly; supramaxilla absent; 69 to 85 oblique scales rows on body; posteriormost two to four premaxillary teeth similar in shape to adjacent teeth . . . . . . . . . . . . . . . . . . . . Opistognathus maxillosus p. 206

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202 PERCIFORMES

Lonchopisthus micrognathus Swordtail jawfish

(Poey, 1860)

Lonchopisthus micrognathus has very large eyes, a relatively small maxilla, and a lanceolate caudal fin. Maxilla extends to about half of eye diameter behind posterior margin of orbit. Posterior margin of maxilla is weakly concave. Jaw teeth are small and conical. Vomer lacks teeth. Gill rakers on lower limb of first arch number 60 to 63. Measurements are expressed as percent of SL: head length 25%– 28%, snout length 3%– 4%, eye diameter 8%– 9%, upper jaw length 14%–15%, pectoral fin length 16%–21%, body depth 22%–26%. Pectoral fin has 16 to 19 rays. Dorsal fin has 11 pungent spines and 16 to 18 rays. Anal fin has 3 spines and 15 to 17 rays. Middle 4 rays of caudal fin are longest. Body, cheek, and operculum are scaled. Scales in lateral series number about 63. Anterior section of lateral line is very near dorsal fin base. Vertebrae number 28: 10 precaudal and 18 caudal. Color is

tan, with narrow blue bands best developed anteriorly. This species occurs in the western Atlantic from southern Florida to northern South America, including the Gulf of Mexico and Cuba. In the Gulf of Mexico it occurs to a depth of 86 m and from the Dry Tortugas to southern Texas and the Gulf of Campeche. It lives commensally with the crab Chasmocarcinus cylindricus and occurs in dense colonies. Food consists of zooplankton. Maximum known size is about 100 mm TL. Lonchopisthus lindneri and L. vanderbilti are synonyms of this species. references: Jordan and Evermann 1898; Longley and Hildebrand 1941; Mead 1959; Hoese and Moore 1977 (as Lonchopisthus lindneri), 1998; Robins and Ray 1986; Boschung 1992; SmithVaniz 2002b.

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OPISTOGNATHIDAE 203

Opistognathus aurifrons (Jordan and Thompson, 1905) Yellowhead jawfish Opistognathus aurifrons has elongated pelvic fins and a dorsal fin with a dark margin. Anterior naris is at end of short tube and lacks cirrus. Maxilla extends about one eye diameter behind posterior margin or orbit. Jaw teeth are relatively large, conical, recurved, and, with exception of those in front of upper jaw, arranged in a single row. Vomer has zero to two (usually one) short, pointed teeth. Gill rakers on first arch number 40 to 62, and number increases with growth. Measurements are expressed as percent of SL: head length 24%– 27%, snout length 3%– 4%, eye diameter 5%– 6%, upper jaw length 13%–16%, pectoral fin length 16%–19%, body depth 19%–21%. Pectoral fin has 19 to 21 rays. Dorsal fin has 9 to 12 (usually 11) spines and 14 to 18 (usually 15 or 16) rays. Anal fin has 3 spines and 14 to 17 rays. Caudal fin is rounded to slightly pointed. Body and operculum are covered with cycloid scales. Scales in horizontal series, posterior to opercular flap, number 81 to 94. Anterior section of lateral line is very near dorsal

fin base. Vertebrae number 25 to 27: 10 or 11 precaudal and 15 to 17 caudal. Color is tan with bluish cast and pale blue dots on body. Head, anterior part of body, and dorsal fin are yellowish. Dorsal fin has narrow dark margin. Pair of black spots is often present on chin. Inside of upper jaw is pale. This species occurs to a depth of 59 m in the western Atlantic from southern Florida and the Bahamas to Central America and Barbados, including the eastern and northern Gulf of Mexico. In the Gulf of Mexico it is known only from the Dry Tortugas and the Flower Garden Reefs. During daylight hours, it generally hovers about 15 to 20 cm above its burrow, with tail down, revolving slowly and feeding on zooplankton. Maximum known size is 97 mm SL. references: Longley and Hildebrand 1941; Böhlke and Thomas 1961; Böhlke 1967; Böhlke and Chaplin 1968; Robins and Ray 1986; Dennis and Bright 1988a; Randall 1996; Smith 1997; Smith-Vaniz 2002b.

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204 PERCIFORMES

Opistognathus lonchurus Moustache jawfish

Jordan and Gilbert, 1882

Opistognathus lonchurus has outermost pelvic fin ray largely free of remainder of fin and dusky markings on snout and upper jaw. Anterior naris is at end of short tube and lacks cirrus. Maxilla extends more than one eye diameter behind posterior margin of orbit. Outer jaw teeth are moderately large, conical, recurved, and arranged in a single row. Inner jaw teeth are filiform and arranged in bands that narrow posteriorly. Vomer has two to eight pointed and recurved teeth, and number increases with size. Gill rakers on lower limb of first arch number 34 to 45, with number increasing with growth. Measurements are expressed as percent of SL: head length 30%– 31%, snout length 4%–5%, eye diameter about 8%, upper jaw length 21%–23%, pectoral fin length 14%–15%, body depth 20%– 22%. Pectoral fin has 18 to 20 rays. Dorsal fin has 11 spines and 12 or 13 rays. Anal fin has 3 (rarely 2) spines and 12 or 13 rays. Caudal

fin is rounded to slightly pointed. Body is covered with cycloid scales. Scales in lateral series number 67 to 86. Anterior section of lateral line is very near dorsal fin base. Vertebrae number 26: 10 precaudal and 16 caudal. Color is pale olive gray, with dusky pigment on snout and upper jaw forming moustachelike marking. Dorsal fin has yellow stripe that is occasionally divided distally. Inside of upper jaw is pale. This species occurs to a depth of 91 m in the western Atlantic from South Carolina to Guyana, including the northeastern and northwestern Gulf of Mexico and the Greater Antilles. It is associated with silty sand mixed with rubble. Maximum known size is 122 mm SL. references: Jordan and Evermann 1898; Longley and Hildebrand 1941; Böhlke and Thomas 1961; Böhlke 1967; Robins and Ray 1986; Dennis and Bright 1988a; Boschung 1992; Smith-Vaniz 2002b.

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OPISTOGNATHIDAE 205

Opistognathus macrognathus Banded jawfish

Poey, 1860

Opistognathus macrognathus has the maxilla extended as a lamina and has dark bands on dorsal fin. Anterior naris is at end of short tube, with a simple, flattened cirrus on posterior rim. Maxilla extends about one eye diameter behind posterior margin of orbit in juveniles and females, and is extended as a thin, flexible lamina beyond posterior margin of operculum in mature males. Outer jaw teeth are moderately large, conical, and recurved, and inner jaw teeth are strongly recurved and arranged in two or three irregular rows. Vomer has one or two large, blunt teeth. Gill rakers on first arch number 31 to 43, with 9 to 15 on upper limb and 21 to 28 on lower limb, and numbers increase with growth. Measurements are expressed as percent of SL: head length 32%– 44%, eye diameter 10%–14%, upper jaw length 19%–25% in females and 24%–39% in males, pelvic fin length 15%–22%, caudal fin length 17%–22%, body depth 17%–24%. Pectoral fin has 19 to 22 rays. Dorsal fin has 10 to 12 spines and 15 to 17 rays. Anal fin has 2 or 3 (usually 3) spines and 15 or 16 rays. Caudal fin is rounded to slightly pointed. Body is covered with cycloid scales. Scales in horizontal series behind opercular flap number 97 to

110. Lateral line is very near dorsal fin base and extends from second to seventh dorsal fin ray. Vertebrae number 28 to 30: 10 precaudal and 18 to 20 caudal. Color is tan, with four to six dark brown blotches on base of dorsal fin and flank, and upper and lower blotches are occasionally joined. Dark blotch is occasionally present on outer part of dorsal fin between sixth and ninth spines. Males have two dark bands, separated by a white band, on underside of maxilla. This species occurs between 0.5 and 44 m in the western Atlantic from southern Florida and the Bahamas to northern South America, including the Antilles. It has been reported once from the eastern Gulf of Mexico; however, this record is questionable (W. F. SmithVaniz, pers. comm., March 1998). This species burrows in mixed sand and gravelly bottoms. Maximum known size is 100 mm TL. references: Jordan and Evermann 1898; Longley and Hildebrand 1940, 1941 (both as Opistognathus fasciatus); Böhlke and Chaplin 1968 (as O. fasciatus); Robins and Ray 1986; Randall 1996; Smith-Vaniz 1997, 2002b.

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206 PERCIFORMES

Opistognathus maxillosus Mottled jawfish

Poey, 1860

Opistognathus maxillosus has a cirrus on anterior naris and a broad, fanlike flap on upper margin of suboperculum. Anterior naris is at end of short tube bearing broadly rounded or palmate cirrus on posterior margin. Maxilla extends about one eye diameter behind posterior margin of orbit and is not sexually dimorphic. Supramaxilla is absent. Outer jaw teeth are moderately large, conical, and recurved, and inner jaw teeth are villiform and posteriorly slanting. Vomer has zero to three large, blunt teeth. Gill rakers on first arch number 26 to 33, with 8 to 11 on upper limb and 17 to 22 on lower limb. Measurements are expressed as percent of SL: head length 33%–38%, postorbital head length 21%–24%, orbit diameter 9%–12%, upper jaw length 21%–25%, pelvic fin length 17%–23%, caudal fin length 20%– 25%, body depth 21%–23%. Pectoral fin has 19 to 22 rays. Dorsal fin has 10 or 11 spines and 14 to 16 rays. Tips of dorsal fin spines bear swollen fleshy tabs. Anal fin has 2 or 3 (usually 2) spines and 14 to 16 rays. Caudal fin is rounded to slightly pointed. Body is covered

with cycloid scales. Scales in horizontal series behind opercular flap number 89 to 100. Anterior section of lateral line is very near dorsal fin base and terminates below third to seventh dorsal fin ray. Vertebrae number 28 or 29: 10 or 11 precaudal and 17 to 19 caudal. Color is mottled dark brown and white, with four large dark blotches on dorsal fin; the first, between the sixth and ninth spines, is often darker than the other three. Pectoral and caudal fins have obscure bands. This species occurs from the shoreline to 12 m in the western Atlantic from southern Florida and the Bahamas to northern South America, including the eastern Gulf of Mexico and the Antilles. The only record from the Gulf of Mexico is from the Dry Tortugas. It excavates an elaborate burrow, with a terminal chamber lined with coral and shell fragments. Maximum known size is 125 mm SL. references: Jordan and Evermann 1898; Longley and Hildebrand 1941; Böhlke 1955; Robins and Ray 1986; Smith-Vaniz 1997, 2002b; Smith 1997.

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OPISTOGNATHIDAE 207

Opistognathus melachasme Yellowmouth jawfish

Smith-Vaniz, 1972

Opistognathus melachasme has posterior margin of maxilla extended as a lamina and has a dark stripe on inner lining of maxilla. Anterior naris is at end of short tube, without cirrus on posterior margin. Maxilla is produced as a thin, flexible lamina extending about one eye diameter behind posterior margin of orbit in juveniles and to end of operculum in adult males. Supramaxilla is very large and subterminally located. Jaws have a patch of symphyseal teeth that are reduced to a single row posteriorly, with those in premaxilla larger anteriorly than posteriorly and those in lower jaw enlarged and hooked laterally. Vomer lacks teeth. Gill rakers on first arch number 27 to 30, with 7 to 9 on upper limb and 20 or 21 on lower limb. Measurements are expressed as percent of SL: head length about 35%, postorbital head length 21%–22%, orbit diameter 9%–11%, upper jaw length 21%–35%, pelvic fin length 25%–26%, caudal fin length 20%–26%, body depth 17%–18%. Pectoral fin has 20 to 22

rays. Dorsal fin has 10 spines and 13 or 14 rays. Anal fin has 2 spines and 12 or 13 rays. Caudal fin is rounded to slightly pointed. Body is covered with cycloid scales. Anterior section of lateral line is very near dorsal fin base and terminates below fourth or fifth dorsal fin ray. Vertebrae number 28: 10 precaudal and 18 caudal. Color is reddish to purplish, with three dark red spots on sides and a large dark spot centered between fourth and seventh dorsal fin spines. Inside of mouth is yellow except for dark corners. This species occurs between 146 and 205 m in the western Atlantic from North Carolina and Cuba to Arrowsmith Bank off northeastern Yucatán. It has not been recorded from the Gulf of Mexico but may occur there. Maximum known size is 100 mm TL.

references: Smith-Vaniz 1972, 1997, 2002b; Robins and Ray 1986.

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208 PERCIFORMES

Opistognathus nothus

Smith-Vaniz, 1997

Opistognathus nothus has a dark stripe on inner lining of maxilla and lacks a lamina extension on maxilla. Anterior naris is at end of short tube, without cirrus on posterior margin. Maxilla extends about one eye diameter behind orbit. Supramaxilla is moderate in size and terminally located on maxilla. Jaws have a patch of symphyseal teeth that are reduced to a single row posteriorly, with those in outer row large and those of inner row hooked backward. Vomer lacks teeth. Gill rakers on first arch number 26 to 32, with 8 or 9 on upper limb and 18 or 23 on lower limb. Measurements are expressed as percent of SL: head length 29%–35%, postorbital head length 16%–20%, orbit diameter 9%–13%, upper jaw length 22%–25%, pelvic fin length 17%– 24%, caudal fin length 18%–24%, body depth

14%–17%. Pectoral fin has 20 or 21 rays. Dorsal fin has 10 spines and 14 rays. Anal fin has 2 spines and 13 rays. Caudal fin is rounded to slightly pointed. Body is covered with cycloid scales. Anterior section of lateral line is very near dorsal fin base and terminates below second to fourth dorsal fin ray. Vertebrae number 28: 10 precaudal and 18 caudal. Color is dark tan, with dark brown spots on head and dorsal fin. This species occurs between 92 and 100 m in the western Atlantic from North Carolina to Cuba and the northeastern Gulf of Mexico. Maximum known size is 79 mm SL. references: Anderson and Smith-Vaniz 1976 (in part as Opistognathus melachasme; Smith-Vaniz 1997, 2002a.

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OPISTOGNATHIDAE 209

Opistognathus robinsi Spotfin jawfish

Smith-Vaniz, 1997

Opistognathus robinsi has maxilla extended as a thin lamina and has dark bands on dorsal fin. Anterior naris is at end of short tube, with a simple cirrus on posterior rim. Maxilla extends about one eye diameter behind posterior margin of orbit in juveniles and females, and extends beyond posterior margin of operculum as a lamina in mature males. Supramaxilla is present on subterminal aspect of maxilla. Outer jaw teeth are moderately large, conical, and recurved, and inner jaw teeth are strongly recurved and arranged in two or three irregular rows. Vomer has one to five large, blunt teeth. Gill rakers on first arch number 28 to 35, with 8 to 12 on upper limb and 19 to 25 on lower limb, and number increases with growth. Measurements are expressed as percent of SL: head length 29%–39%, postorbital head length 16%–22%, orbit diameter 9%–15%, upper jaw length 19%–25% in females and 21%–32% in males, pelvic fin length 17%–

25%, caudal fin length 18%–24%, body depth 18%–24%. Pectoral fin has 18 to 21 rays. Dorsal fin has 10 or 11 spines and 15 to 18 (usually 17) rays. Anal fin has 2 or 3 (usually 3) spines and 16 to 18 rays. Caudal fin is rounded. Body is covered with cycloid scales. Scales in horizontal series number about 73 to 88. Lateral line is very near dorsal fin base and extends below first to fifth dorsal fin ray. Vertebrae number 29 to 31: 10 precaudal and 19 to 21 caudal. Color is brownish, with dark reticulations and speckles on head and nape, pale spots and blotches on body, and prominent ocellus between third and seventh dorsal spines. This species occurs from near shore to 46 m in the western Atlantic from South Carolina and the Bahamas to the northern Gulf of Mexico. Maximum known size is 131 mm SL. references: Robins and Ray 1986 (as Opistognathus sp.); Smith-Vaniz 1997, 2002a.

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210 PERCIFORMES

Opistognathus whitehursti Longley 1927 Dusky jawfish

Opistognathus whitehursti has a fleshy cirrus on anterior naris and lacks lamina extension of maxilla. Anterior naris is at end of short tube bearing a simple cirrus on posterior rim. Maxilla extends one-half to one eye diameter behind posterior margin of orbit. Jaw teeth in upper jaw are moderate to small, conical, slightly recurved, and arranged in a single row. In mature males, posteriormost premaxillary tooth is more strongly hooked than adjacent teeth. Teeth of lower jaw are in two rows anteriorly and a single row posteriorly, with those in outer row larger. Vomer has one or two (rarely three) conical teeth. Gill rakers on first arch number 24 to 32, with 8 to 11 on upper limb and 16 to 21 on lower limb. Measurements are expressed as percent of SL: head length 31%– 41%, postorbital head length 19%–25%, orbit diameter 10%–13%, upper jaw length 20%–26%, pelvic fin length 16%–26%, caudal fin length 19%–27%, body depth 21%–23%. Pectoral fin has 17 to 20 rays. Dorsal fin has 10 to 12 spines and 13 to 15 rays. Dorsal fin spines bear swollen fleshy tabs. Anal fin has 3 spines and 12 to 14 rays. Caudal fin is rounded to slightly

pointed. Body is covered with cycloid scales. Anterior section of lateral line is very near dorsal fin base and extends below last dorsal fin spine to fourth dorsal fin ray. Scales in horizontal series number 42 to 54. Vertebrae number 26 to 28: 10 or 11 precaudal and 16 to 18 caudal. Color is mottled, mostly dark brown, and occasionally with black flecks. Spinous part of dorsal fin has dark margin and bluish black spot between second and fourth spines. Rayed section of dorsal fin and caudal fin may be pale yellow. This species occurs from near the shoreline to 47.5 m in the western Atlantic from South Florida and the Bahamas to southern Brazil, including the Dry Tortugas in the eastern Gulf of Mexico and the Bahamas. It constructs burrows in sandy to gravelly bottoms, often on the edges of Thalassia beds. Maximum known size is 65 mm SL.

references: Longley and Hildebrand 1940, 1941; Böhlke 1955; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith-Vaniz 1997, 2002b.

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211

PRIACANTHIDAE

Bigeyes

Bigeyes are oblong, moderately deep bodied to deep bodied and compressed, with a strongly oblique mouth, very large eyes, and spinous cycloid scales. Dorsal profile of head is convex. Snout is short and acute. Anterior and posterior nares are close set, with anterior naris rounded and posterior naris slitlike. Mouth is terminal and oblique, with lower jaw extending distinctly beyond upper jaw and upper jaw terminating posterior to anterior margin of eye. Small conical teeth occur in jaws, vomer, and palatine. Eye is located near dorsal profile of head and possesses very reflective tapetum lucidum, producing eye shine. Preoperculum has serrated margin, with a single spine at angle. Operculum has broad, flat spine. Suboperculum and interoperculum have serrated ventral margins. Branchiostegal rays number 6. Slit is present behind fourth gill arch, and gill membranes are separate and free of isthmus. Pseudobranch on inner surface of gill cover is well developed. Gill rakers on first arch are slender and elongate and number 17 to 32. Pectoral fin has nearly vertical base, inserts on lower flank, and has 17 to 21 rays. Dorsal fin is single, is not indented, and consists of 10 spines and 11 to 15 rays. Pelvic fin is thoracic and consists of 1 spine and 5 rays. Inner ray of pelvic fin is connected to body partially to totally by membrane. Anal fin is slightly smaller than dorsal fin and consists of 3 spines and 10 to 16 rays. All fin spines are serrated, and last rays of dorsal and anal fins are split to base. Caudal fin is rounded and emarginate or lunate and consists of 16 principal rays, 14 of which are branched. Body and most of head, including maxilla, lower jaw, and branchiostegal rays, are covered with scales. Scales are adherent and cycloid, with spines on posterior margin. Spines are continuous with surface of scale and not on separate platelets, thus they are not ctenoid scales. Lateral line is continuous and highly arched anteriorly. Lateral line scales bear simple tubes and are larger than more ventrally located lateral scales. Axillary process at base of pelvic fin is absent. Gas bladder is present and possesses posterior and anterior extensions, and extrinsic muscles are present or absent. Vertebrae number 22 or 23: 9 or 10 precaudal and 13 caudal. Color is bright red in life. Bigeyes occur worldwide in tropical and warm temperate seas, with exception of waters around northern New Zealand and the Mediterranean Sea. They are generally associated with rock formations and reefs on continental and insular shelves, although some occur to 400 m. Most species are nocturnal and feed on polychaetes, cephalopods, crustaceans, and small ray-finned fishes. Eggs, larvae, and early juveniles are epipelagic, and larger juveniles and adults are associated with the bottom. There are

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212 PERCIFORMES

18 species in four genera, 4 species in separate genera in the western Central Atlantic, and 3 species in separate genera in the Gulf of Mexico. references: Starnes 1988, 2001, 2002; Nelson 1994; Tominaga et al. 1996; Smith 1997.

Key to the Species of the Gulf of Mexico (Adapted from Starnes 1988) 1a. Anal fin rays 10 or 11; dorsal fin rays 11 or 12; scales in lateral series 36 to 51. . . . . . . . . . . . Pristigenys alta p. 215 1a. Anal fin rays 13 to 16; dorsal fin rays 12 to 15; scales in lateral series 56 to 115 . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Posterior preoperculum (behind sensory canal) lacking scales and striae; membranes of rayed sections of dorsal and anal fins, and caudal fin with rows of elliptical dark spots; anterior profile of body nearly symmetrical . . . . . . . . . . . . . . . . . Heteropriacanthus cruentatus p. 213 2b. Posterior preoperculum (behind sensory canal) scaled; membranes of rayed sections of dorsal and anal fins, and caudal fin without dark spots; anterior profile of body asymmetrical. . . . . . . . . . . . . . Priacanthus arenatus p. 214

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PRIACANTHIDAE 213

Heteropriacanthus cruentatus Glasseye snapper

(Lacepède, 1801)

Heteropriacanthus cruentatus has a naked and serrated preoperculum and elliptical spots on unpaired fins. Premaxillary teeth are small and number 75 to 85, maxillary teeth number 20 to 30, vomerine teeth number 40 to 50, and palatine teeth number 20 to 25. Lachrymal and second infraorbital bones are finely serrated. Premaxilla is serrated on ventral margin. Gill rakers on first arch number 21 to 25, with 4 to 6 on upper limb and 17 to 20 on lower limb. Measurements are expressed as percent of SL: head length 31%–34%, head depth 25%– 38%, orbit diameter 13%–17%, interorbital width 8%–11%, snout length 9%–11%, lower jaw length 19%–21%, body depth 38%– 45%, caudal peduncle depth 9%–10%. Pectoral fin is of moderate length, broadly pointed, and slightly shorter than pelvic spine length and has 18 or 19 rays. Spinous section of dorsal fin increases slightly in height posteriorly and has 10 spines. Rayed section of dorsal fin has 11 to 13 rays. Pelvic fin barely extends to first anal fin spine. Anal fin has 13 or 14 rays. Caudal fin is slightly emarginate to slightly convex, with acute upper and lower corners. Scales in horizontal series number 78 to 96. Lateral line scales number 63 to 81. Head scales have relatively weak spinules. Scales are lacking from lips and posterior margin of preoperculum. Color is pink to red, including iris of eye. Occasionally body is silvery and blotched with pink and red. Spinous dorsal fin membranes

and margin of caudal fin are often dusky. Membranes of rayed sections of dorsal, anal, and caudal fins are generally patterned with elliptical dark spots. Pelagic juveniles are silvery, with dark blotches and two well-defined dark spots on base of caudal fin. This species occurs worldwide in tropical and subtropical seas on continental and insular shelves, although it is more common in insular habitats. In the western Atlantic it occurs from New Jersey and Bermuda to Argentina, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs off the west coast of Florida, on the Flower Garden Banks, and in Campeche Bay. It is most frequently captured at a depth of 20 m or less, and it is generally active at night and hides in holes and under rocks or ledges on the reef during the day. Food includes polychaetes, octopi, stomatopods, pelagic shrimps, crabs, and small ray-finned fishes. Spawning occurs in late spring and early summer in the Caribbean Sea. Maximum known size is 254 mm SL.

references: Longley and Hildebrand 1941; Caldwell 1962b; Böhlke and Chaplin 1968; Randall 1968a, 1996; Heemstra 1986c; Hoese and Moore 1977, 1998; Robins and Ray 1986 (all as Priacanthus cruentatus); Starnes 1988, 2001, 2002; Boschung 1992; Schaldach et al. 1997 (as P. cruentatus); Smith 1997; Smith-Vaniz et al. 1999.

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214 PERCIFORMES

Priacanthus arenatus Cuvier, 1829 Bigeye

Priacanthus arenatus has an asymmetrical anterior profile and scales on posterior margin of preoperculum, and lacks elliptical spots on unpaired fins. Upper jaw teeth are small and number 70 to 110, lower jaw teeth number 25 to 45, vomerine teeth number 25 to 65, and palatine teeth number 25 to 40. Lachrymal and second infraorbital bones are finely serrated. Premaxilla is smooth, although serrations often occur on either side of small spine at corner. Gill rakers on first arch number 28 to 32, with 6 to 8 on upper limb and 21 to 24 on lower limb. Measurements are expressed as percent of SL: head length 28%–35%, head depth 28%–35%, snout length 8%–10%, orbit diameter 12%–17%, interorbital width 7%– 9%, lower jaw length 17%–21%, body depth 32%– 40, caudal peduncle depth 7%–12%. Pectoral fin is of moderate length, broadly pointed, and shorter than pelvic spine length and has 17 or 19 rays. Spinous section of dorsal fin increases slightly in height posteriorly and has 10 spines. Rayed section of dorsal fin has 13 to 15 rays. Pelvic fin generally extends to third anal fin spine. Anal fin has 14 or 16 rays. Caudal fin is slightly emarginate, with acute upper and lower corners. Scales in

horizontal series number 83 to 91. Lateral line scales number 71 to 84. Head scales have relatively weak spinules. Scales are lacking from lips. Color is uniformly bright red or pink, or bright red or pink with silvery blotches. Dark red spots occasionally occur above lateral line. Unpaired fins are lightly dusted with dark melanophores. Pelvic fin is blackish, especially distally. Pelagic juveniles are silver or dusky, with three rows of dark spots on spinous section of dorsal fin. This species occurs in the tropical to subtropical Atlantic Ocean. In the western Atlantic it occurs between 20 and 250 m, from North Carolina and Bermuda to northern Argentina, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with reefs. Maximum known size is 450 mm SL.

references: Hildebrand and Schroeder 1928; Gunter and Knapp 1951; Caldwell 1962b; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hastings et al. 1976; Hoese and Moore 1977, 1998; Matsuura 1983b; Starnes 1988, 2002; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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PRIACANTHIDAE 215

Pristigenys alta Short bigeye

(Gill, 1862)

Pristigenys alta is broadly ovate in profile, deep bodied, and relatively robust, with scales on head highly modified and covered with spines, scales on posterior margin of preoperculum, and no elliptical spots on unpaired fins. Upper jaw teeth are small and number 180 to 200, lower jaw teeth number 70 to 75, vomerine teeth number 90 to 100, and palatine teeth number 90 to over 100. Lachrymal and second infraorbital bones have coarse serrations and spines on anteroventral margin. Premaxilla is serrated, and spine at corner occurs in juveniles and is vestigial in adults. Gill rakers on first arch number 24 to 28, with 7 or 8 on upper limb and 17 to 20 on lower limb. Measurements are expressed as percent of SL: head length 36%– 41%, head depth 41%– 42%, orbit diameter 20%–22%, interorbital width 6%–10%, snout length 9%–10%, lower jaw length 22%–26%, body depth 48%– 63%, caudal peduncle depth 11%–13%. Pectoral fin is broadly pointed and shorter than pelvic spine length and has 16 or 19 rays. Margin of spinous section of dorsal fin is rounded and has 10 spines. Rayed sections of dorsal fin and anal fin have pointed posterodorsal margins and have 11 or 12 and 10 or 11 rays, respectively. Pelvic fin generally extends to anal fin origin in juveniles and nearly reaches origin in adults. Caudal fin has slightly rounded posterior margin,

with acute upper and lower corners. Scales in horizontal series number 42 to 45. Lateral line scales number 36 to 40. Head scales, especially on chin, gular region, and top of head, have relatively strong spinules. Scales are lacking from lips. Color is dark red in life and pink to orange after preservation, with margins of rayed sections of dorsal and anal fins and caudal fin black. Pelagic juveniles are silvery gray, with spinous section of dorsal fin black. This species occurs between 5 and 125 m in the western Atlantic from New England and Bermuda to Costa Rica, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with rocks and burrows. Off southern Florida it spawns from mid-July through mid-September. Transformation from pelagic to benthic juvenile stages occurs between 35 and 55 mm SL. Maximum known size is 261 mm SL.

references: Longley and Hildebrand 1941 (as Pseudopriacanthus altus); Caldwell 1962a,b; Hoese and Moore 1977, 1998; Robins and Ray 1986 (all as Pseudopriacanthus altus); Shipp and Hopkins 1978; Starnes 1988, 2002; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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216

APOGONIDAE

Cardinalfishes

Cardinalfishes are oblong or moderately elongate and moderately robust to moderately compressed, with a large eye, two short dorsal fins usually, and a long caudal peduncle. Dorsal profile of head is moderately convex. Snout is short, and mouth is terminal and oblique (rarely slightly oblique), with lower jaw about equal with upper jaw or slightly protruding and upper jaw extending from about middle to posterior margin of eye. Nostril is paired, and nares are close set. Supramaxilla is present or absent on dorsal margin of maxilla. Villiform teeth occur in jaws, vomer, and often palatines, with large canines also often present in jaws. Preoperculum is serrated along posterior and ventral margins in most species, and ridge is smooth in most but serrated in some species. Operculum has a single weak spine extending from stay and running entire length. Branchiostegal rays number 7. Gill membranes are free of isthmus, and slit is present behind last (fourth) gill arch. Gill rakers are well developed. Pectoral fin insertion is below midflank, and fin is well developed, with 10 to 20 rays. Dorsal fin is subdivided: First consists of six to eight slender spines, and second consists of one spine and 8 to 14 rays, with last ray split to base. Pelvic fin inserts below pectoral fin base and has one spine and 5 rays. Anal fin is similar to second dorsal fin in shape and has two slender spines and 8 to 18 rays. Caudal fin is concave, truncate, or rounded. Scales are ctenoid, cycloid, or occasionally absent, and are usually present on nape and operculum. Lateral line is generally complete but occasionally incomplete or absent. Light organs are present in some species. Vertebrae number 24 or 25: 10 precaudal and 14 or 15 caudal. Maximum known size is about 200 mm TL. The above description applied to all but one recently described species from northwestern Australia that has a single dorsal fin, with six spines and 19 rays, an anal fin with one spine and 15 or 16 rays, and a short, deep caudal peduncle (Baldwin and Johnson 1999). Cardinalfishes occur worldwide in tropical to warm temperate seas, mostly in shallow marine waters. Some species occur in brackish to fresh water, and others are found to a depth of 290 m. They are small carnivorous fishes associated with coral and rocky reefs and sandy or weedy areas, and they are nocturnally active. Many species are mouth brooders, with either males or females brooding the eggs. A number of species are inquilines, living within the body cavities of other organisms such as sponges and mollusks, and others are closely associated with sea urchins. There are 207 species in 24 genera, 23 species in 3 genera in the western Central Atlantic, and 15 species in 3 genera in the Gulf of Mexico.

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APOGONIDAE 217

references: Böhlke and Randall 1968; Fraser 1972; Nelson 1984; Smith-Vaniz et al. 1999; Allen 2001a; Gon 2002a.

Key to the Species of the Gulf of Mexico (Adapted in part from Böhlke and Chaplin 1968) 1a. Scales on body cycloid; scales absent from midline of nape; pectoral fin rays 14 to 17; posterior margin of preoperculum entire (smooth) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Scales on body ctenoid; scales present on midline of nape; pectoral fin rays 11 to 13; posterior margin of preoperculum finely to coarsely serrate . . . . . . . . . . . . . . . . . . . . . . 4 2a. Pectoral fin rays 14 (rarely 13 or 15); pelvic fin not reaching beyond anterior third of anal fin base; pelvic fin dusky, occasionally with black tips, rarely black . . . . . . . . . . . . . . . . . . . . . . . . . . Astrapogon alutus p. 229 2b. Pectoral fin rays 15 or 16 (rarely 14 or 17); pelvic fin usually reaching middle third of anal fin base or farther; pelvic fin usually entirely black, occasionally with spotting on base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. Gill rakers on lower limb of first arch 12 to 14; pectoral fin rays 15 to 17 (usually 16 or 17); dorsal profile of head over eye rounded; pelvic fin reaching between anterior and middle third of anal fin base . . . . . . . . . . . . . . . . . . . . Astrapogon puncticulatus p. 230 3b. Gill rakers on lower limb of first arch 10 or 11; pectoral fin rays 14 or 15 (usually 15); dorsal profile of head over eye flat; pelvic fin reaching to posterior third of anal fin base . . . . . . . . . . . . . . . . . . . . . Astrapogon stellatus p. 231 4a. Large fleshy flap extending beyond preopercular corner; inner pelvic fin rays connected along length of abdomen by membrane; body color brownish . . . . . . . . . . . . . . . . . . . 5 4b. Fleshy preopercular flap absent or present but not produced beyond preopercular corner; inner pelvic rays free of abdomen; body color reddish . . . . . . . . . . . . . . . . . . . 7 5a. Jaw teeth villiform; gill rakers on lower limb of first arch 12 to 16 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5b. Some anterior jaw teeth in upper jaw and lateral teeth in lower jaw enlarged; gill rakers on lower limb of first arch 11 to 13 . . . . . . . . . . . . . . . Phaeoptyx pigmentaria p. 233 6a. Usually prominent dark stripes at bases of second dorsal fin and anal fin, and separated from bases by pale streak; gill rakers on lower limb of first arch 14 to 16 (modally 15); color pattern sharply defined with brown spots on pale background . . . . . . . . . . . . Phaeoptyx conklini p. 232 6b. Dark stripes at bases of second dorsal fin and anal fin faint or absent, and when present, not separated from bases by pale streak; gill rakers on lower limb of first arch 12 to 15;

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218 PERCIFORMES

color pattern not sharply defined with brown spots . . . . . . . . . . . . . . . . . . . . . . . . . . . Phaeoptyx xenus p. 234 7a. Jaws with prominent canine teeth; anal fin rays 9 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apogon affinis p. 220 7b. Jaws with bands of villiform teeth; anal fin rays 8 . . . . . . 8 8a. Body with dark pigmentation in definite localized blotches or bands; either vertical bar below base of second dorsal fin or roundish spot below or behind base of second dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8b. Body without distinct dark pigmentation, dark pigment limited to light peppering, faint blotches, or dusky bar on caudal fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 9a. Dark spot or saddle on upper body below or immediately behind second dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . 10 9b. Dark vertical bar behind second dorsal fin and extending to near base of anal fin . . . . . . . . . . . . . . . . . . . . . . . . . . 12 10a. Caudal peduncle with distinct dark marking; black spot on operculum; black spot below posterior section of second dorsal fin; no white spot following black spot below second dorsal fin; first dorsal fin lacking blackish area; gill rakers on lower limb of first arch 13 or 14; first dorsal fin spine 31% to 40% of second dorsal fin spine . . . . . . . . . . . . . 11 10b. Caudal peduncle without distinct dark marking; no black spot on operculum; small black saddle marking behind last dorsal fin ray; white spot following black saddle behind last dorsal fin spine; large blackish area on first dorsal fin; gill rakers on lower limb of first arch 16 or 17; first dorsal fin spine 25% to 29% of second dorsal fin spine . . . . . . . . . . . . . . . . . . . . . . . . . . . Apogon lachneri p. 223 11a. Broad blackish saddle on caudal peduncle extending below lateral line (may be faint); scales around caudal peduncle 17 to 20 . . . . . . . . . . . . . . . . . . . Apogon maculatus p. 224 11b. Small blackish round spot on upper half of caudal peduncle, free of spot on other side; scales around caudal peduncle 15 or 16 . . . . . . . . . . . . . Apogon pseudomaculatus p. 226 12a. Dark bar on caudal peduncle narrow, about width of vertical bar behind second dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . Apogon binotatus p. 222 12b. Dark bar on caudal peduncle broad, equal to or broader than eye diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13a. Dark bar on caudal peduncle with black lateral margins (except in young); dark bar between second dorsal fin and anal fin separated from base of second dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . Apogon townsendi p. 228 13b. Dark bar on caudal peduncle without black lateral margins; dark bar between second dorsal fin and anal fin in contact with last rays of second dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . Apogon planifrons p. 225

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APOGONIDAE 219

14a. Gill rakers on lower limb of first arch 12 to 14; median predorsal scales three to five; preopercular margin strongly serrated . . . . . . . . . . . . . . Apogon quadrisquamatus p. 227 14b. Gill rakers on lower limb of first arch 10 or 11; median predorsal scales five or six; preopercular margin finely serrated . . . . . . . . . . . . . . . . . . . . Apogon aurolineatus p. 221

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220 PERCIFORMES

Apogon affinis (Poey, 1875) Bigtooth cardinalfish

Apogon affinis has prominent canine teeth in jaws and a dusky stripe anterior and posterior to eye. Lower jaw extends beyond upper jaw. Upper jaw has two to four canine teeth on either side of symphysis and a narrow band of villiform teeth posterior to canines. Lower jaw has outer band of villiform teeth and inner, widely spaced row of recurved canine teeth. Vomer has several short, recurved canines on either side. Palatine has narrow band of villiform teeth. Preopercular ridge is coarsely serrate, and preopercular free margin is finely serrate and slightly expanded at angle, with fleshy posteroventral process extending slightly beyond its serrated margin. Gill rakers on first arch number 4 or 5 on upper limb and 14 or 15 on lower limb. Measurements are expressed as percent of SL: head length 36%– 40%, snout length 8%–9%, eye diameter 10%–11%, upper jaw length 19%–20%, body depth 34%– 39%. Pectoral fin is relatively long and has

12 rays. First dorsal fin has six slender spines, and second dorsal fin has one spine and 9 rays. Anal fin has 9 rays. Caudal fin is forked. Scales are ctenoid and deciduous. Color is salmon pink except for black band on either side of eye and black edge of caudal fin. Brooding males have a black fleshy chin flap. This species occurs in tropical waters of the Atlantic Ocean. In the western Atlantic it occurs at depths from 20 to 90 m, from the Florida Keys and Bermuda to Suriname, including the entire Gulf of Mexico and the Bahamas. It is generally associated with hollow coral heads, caves, and overhangs. Maximum known size is about 88 mm SL. references: Böhlke and Chaplin 1968; Smith et al. 1971; Williams and Shipp 1980; Matsuura 1983c; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Gon 2002a.

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APOGONIDAE 221

Apogon aurolineatus Bridle cardinalfish

(Mowbray, 1927)

Apogon aurolineatus has small black pigment spots on body and two dark bars below and behind eye. Jaws, vomer, and palatine have bands of villiform teeth. Preopercular ridge is smooth, and preopercular free margin is finely serrated and slightly expanded at angle, with fleshy posteroventral process extending slightly beyond its serrated margin. Gill rakers on first arch number 4 on upper limb and 10 or 11 on lower limb. Measurements are expressed as percent of SL: head length 33%–36%, snout length 7%–9%, eye diameter 10%–11%, upper jaw length 19%–20%, body depth 32%– 35%. Pectoral fin has 12 rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Caudal fin is emarginate. Scales are ctenoid and deciduous. Median predorsal

scales number 5 or 6, and circumpeduncular scales number 16 to 18. Color is pale pinkish to golden, with two dark bars radiating from below and behind eye and body marked by enlarged melanophores. This species occurs from the shoreline to 75 m in the western Atlantic from southern Florida and the Bahamas to northern South America, including the northern and southern Gulf of Mexico. It is common in grass beds and on coral reefs. Maximum known size is 65 mm TL. references: Hoese 1958; Böhlke and Randall 1968; Hoese and Moore 1977, 1998; Ogren and Brusher 1977; Williams and Shipp 1980; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Gon 2002a.

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222 PERCIFORMES

Apogon binotatus Barred cardinalfish

(Poey, 1867)

Apogon binotatus has a narrow dark bar below posterior end of second dorsal fin and on caudal peduncle. Jaws, vomer, and palatine have bands of villiform teeth. Preopercular ridge is smooth, and posterior margin of preoperculum is finely serrated and rounded at angle, with fleshy posteroventral process not extending beyond its serrated margin. Gill rakers on first arch number 4 or 5 on upper limb and 12 to 14 on lower limb. Measurements are expressed as percent of SL: head length 38%– 41%, snout length 9%–11%, eye diameter 12%–13%, upper jaw length 21%–22%, caudal peduncle length 20%–22%, body depth 32%–37%. Pectoral fin has 12 rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Caudal fin is emarginate. Scales are ctenoid and deciduous. Median

predorsal scales number 6 or 7, and circumpeduncular scales number 15 or 16. Color is pink to light red and somewhat translucent, with two distinct, narrow dark bars on body. This species occurs between the shoreline and 60 m in the western Atlantic from southeastern Florida and the Bahamas to Venezuela, including the eastern Gulf of Mexico and the Greater and Lesser Antilles. The record of this species from Bermuda is probably based on a misidentification (Smith-Vaniz et al. 1999). This species is limited to clear water and is associated with coral reefs. Maximum known size is 100 mm TL. references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Böhlke and Randall 1968; Randall 1968a, 1996; Robins and Ray 1986; Cervigón 1993b; Smith 1997; Gon 2002a.

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APOGONIDAE 223

Apogon lachneri Böhlke, 1959 Whitestar cardinalfish

Apogon lachneri has a black saddle mark behind last dorsal fin ray, a median white spot behind dark mark, and dark shading on caudal peduncle behind white spot. Jaws, vomer, and palatine have narrow bands of villiform teeth. Preopercular ridge is smooth, and posterior margin of preoperculum is serrated and rounded at angle, with fleshy posteroventral process not extending beyond its serrated margin. Gill rakers on first arch number 5 or 6 on upper limb and 16 or 17 on lower limb. Measurements are expressed as percent of SL: head length 40%– 42%, snout length 10%–11%, eye diameter 14%–16%, upper jaw length 20%–22%, caudal peduncle length 29%– 31%, body depth 31%–35%. Pectoral fin has 12 rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and

anal fin has two spines and 8 rays. Caudal fin is emarginate. Scales are ctenoid and deciduous. Median predorsal scales number 6 or 7, and scales in lateral line number 25. Color is reddish, with posterior section of first dorsal fin and anterior section and tip of second dorsal fin black. Black mark behind second dorsal fin often extends forward along base of fin. This species occurs in the western Atlantic from southern Florida and the Bahamas to Belize, including the southern Gulf of Mexico off Veracruz and the Greater and Lesser Antilles. It is associated with clear water and coral reefs. Maximum known size is 65 mm TL. references: Böhlke 1959; Böhlke and Chaplin 1968; Randall 1996; Schaldach et al. 1997; Smith 1997; Gon 2002a.

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224 PERCIFORMES

Apogon maculatus Flamefish

(Poey, 1860)

Apogon maculatus has a round dark spot below posterior end of second dorsal fin and a broad dark saddle on caudal peduncle. Dorsal profile of head is evenly convex, and snout is blunt. Jaws, vomer, and palatine have narrow bands of villiform teeth. Preopercular ridge is smooth, and posterior margin of preoperculum is finely serrated and rounded at angle, with fleshy posteroventral process not extending beyond its serrated margin. Gill rakers on first arch number 4 or 5 on upper limb and 13 or 14 on lower limb. Measurements are expressed as percent of SL: head length 39%– 41%, snout length 9%–10%, eye diameter 12%–14%, upper jaw length 21%–22%, caudal peduncle length 22%–25%, body depth 34%–37%. Pectoral fin has 12 rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Caudal fin is emarginate. Scales are ctenoid and deciduous. Median predorsal scales

number 7 or 8, and circumpeduncular scales number 19 or 20. Color is deep orangish red, with black bar behind eye on operculum, two white stripes passing through eye, blacks spot below second dorsal fin, and black saddle on caudal peduncle extending below lateral line. This species occurs from near shore to 100 m in the western Atlantic from Massachusetts and Bermuda to Venezuela, including the entire Gulf of Mexico and the Bahamas. Maximum known size is 85 mm SL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Böhlke and Randall 1968; Randall 1968a, 1996; Bright and Cashman 1974; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Hastings 1979; Williams and Shipp 1980; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; SmithVaniz et al. 1999; Gon 2002a.

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APOGONIDAE 225

Apogon planifrons Pale cardinalfish

Longley and Hildebrand, 1940

Apogon planifrons has a vertical bar below posterior end of second dorsal fin and a broad dark saddle on caudal peduncle. Jaws, vomer, and palatine have narrow bands of villiform teeth. Preopercular ridge is smooth, and posterior margin of preoperculum is finely serrated and rounded at angle, with fleshy posteroventral process not extending beyond its serrated margin. Gill rakers on first arch number 6 on upper limb and 13 to 16 on lower limb. Measurements are expressed as percent of SL: head length about 41%, snout length about 10%, eye diameter about 13%, upper jaw length about 23%, caudal peduncle length about 28%, body depth about 32%. Pectoral fin has 12 rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Caudal fin

is emarginate. Scales are ctenoid and deciduous. Circumpeduncular scales number 15 or 16. Color is pale red, with a black bar on body, extending from posterior section of second dorsal fin, and a broad black saddle on caudal peduncle. This species occurs from near shore to 30 m in the western Atlantic from southern Florida and Bermuda to Venezuela, including the eastern Gulf of Mexico and the Bahamas. It is associated with coral reefs and rocky areas. Maximum known size is about 100 mm TL. references: Longley and Hildebrand 1940, 1941; Böhlke and Chaplin 1968; Böhlke and Randall 1968; Robins and Ray 1986; Smith 1997; SmithVaniz et al. 1999; Gon 2002a.

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226 PERCIFORMES

Apogon pseudomaculatus Twospot cardinalfish

Longley, 1932

Apogon pseudomaculatus has a black spot below posterior part of second dorsal fin and a small roundish spot on upper part of caudal peduncle. Jaws, vomer, and palatine have narrow bands of villiform teeth. Preopercular ridge is smooth, and posterior margin of preoperculum is finely serrated and rounded at angle, with fleshy posteroventral process not extending beyond its serrated margin. Gill rakers on first arch number 4 or 5 on upper limb and 12 to 14 on lower limb. Measurements are expressed as percent of SL: head length 37%–38%, snout length 8%–9%, eye diameter 11%–12%, upper jaw length 20%–22%, caudal peduncle length 22%–25%, body depth 34%–37%. Pectoral fin has 12 rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Caudal fin is emarginate. Scales are ctenoid and deciduous. Circumpe-

duncular scales number 15 or 16. Color is deep orangish red, with black spot on operculum, black spot below posterior section of second dorsal fin, and black spot on caudal peduncle. This species occurs in the western Atlantic from Massachusetts and Bermuda to the Bahamas and southern Brazil, including the Gulf of Mexico. It is found throughout the Gulf of Mexico from near shore to 403 m. Maximum known size is about 110 mm TL.

references: Longley and Hildebrand 1940, 1941; Böhlke and Chaplin 1968; Böhlke and Randall 1968; Hastings et al. 1976; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Shipp and Hopkins 1978; Hastings 1979; Boschung 1992; Matsuura 1983c; Robins and Ray 1986; Cervigón 1993b; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Gon 2002a.

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APOGONIDAE 227

Apogon quadrisquamatus Sawcheek cardinalfish

Longley, 1934

Apogon quadrisquamatus has a dark smudge on caudal peduncle but no black spot or bar below posterior part of second dorsal fin. Jaws, vomer, and palatine have narrow bands of villiform teeth. Preopercular ridge is smooth, and posterior margin of preoperculum is strongly serrated and rounded at angle, with fleshy posteroventral process not extending beyond its serrated margin. Gill rakers on first arch number 3 on upper limb and 12 to 14 on lower limb. Measurements are expressed as percent of SL: head length 34%– 41%, snout length 6%– 8%, eye diameter 11%–12%, upper jaw length 22%–23%, caudal peduncle length 19%–22%, body depth 32%–34%. Pectoral fin has 12 rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Caudal fin is slightly emarginate. Scales are ctenoid and deciduous. Predorsal scales number 3 to 5 (usually 4), and circumpeduncular scales

number 16 or 18. Color is translucent pale orange to deep red and peppered with small melanophores, with black smudge on caudal peduncle. Fins are yellowish. This species occurs from 12 to 60 m in the western Atlantic from southern Florida and the Bahamas to southern Brazil, including the eastern Gulf of Mexico. It occurs on coral reefs and over grass flats and sand and rubble bottoms. Feeding takes place on or near reefs at night. This species is also associated with Diadema sea urchins, sea anemones, and long tubular sponges. Maximum known size is 58.2 mm SL. references: Longley and Hildebrand 1940, 1941; Böhlke and Chaplin 1968; Böhlke and Randall 1968; Randall 1968a, 1996; Williams and Shipp 1980; Robins and Ray 1986; Cervigón 1993b; Gilbert and Tyler 1997; Smith 1997; Gon 2002a.

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228 PERCIFORMES

Apogon townsendi Belted cardinalfish

(Breder, 1927)

Apogon townsendi has a narrow dark bar below second dorsal fin and a broad dark ring on caudal peduncle that has a distinct dark border. Dorsal profile of head is slightly convex, and snout is blunt. Jaws, vomer, and palatine have narrow bands of villiform teeth. Preopercular ridge is smooth, and posterior margin of preoperculum is strongly serrated and rounded at angle, with fleshy posteroventral process not extending beyond its serrated margin. Gill rakers on first arch number 16 to 18 on lower limb. Measurements are expressed as percent of SL: head length about 38%, snout length about 7%, eye diameter about 15%, upper jaw length about 20%, caudal peduncle length about 23%, body depth about 36%. Pectoral fin has 12 rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Caudal fin is forked. Scales are ctenoid and deciduous. Predorsal scales number 6, and scales encircling caudal peduncle number 12. Color

is pinkish red dorsally and yellowish red ventrally, with narrow dark bar at base of second dorsal fin extending to anal fin base and broad dark ring anterior to caudal fin. Narrow dark bar is separate from second dorsal fin. Anterior margin of broad dark ring is darker than posterior margin. This species occurs from 3 to 55 m in the western Atlantic from southern Florida and Bermuda to northern South America, including the eastern and western Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with coral reefs and rocky areas and often is located near drop-offs. Maximum known size is about 65 mm SL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Böhlke and Randall 1968; Randall 1968a, 1996; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Gon 2002a.

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APOGONIDAE 229

Astrapogon alutus Bronze cardinalfish

(Jordan and Gilbert, 1882)

Astrapogon alutus has a relatively short and darkly pigmented pelvic fin and a dark brown head and body. Jaws, vomer, and palatine have narrow bands of villiform teeth. Preopercular ridge is smooth, and preoperculum is smooth along posterior margin and scalloped along ventral margin. Gill rakers on first arch number 9 to 12 on lower limb. Measurements are expressed as percent of SL: head length 36%– 41%, snout length 7%–9%, eye diameter 11%–12%, upper jaw length 20%–21%, caudal peduncle length 18%–21%, pelvic fin length 26%–35%, body depth 36%–38%. Pectoral fin has 14 (rarely 13 or 15) rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Caudal fin is truncate to

slightly convex. Scales are cycloid and deciduous. Median predorsal scales are lacking. Color is dark brown, with peppering of dark melanophores on head and body. Pelvic fin is dusky and occasionally black tipped. This species occurs in shallow water in the western Atlantic from North Carolina to Venezuela, including the eastern Gulf of Mexico and the Bahamas. It is usually associated with sea-grass beds. Maximum known size is about 65 mm TL. references: Longley and Hildebrand 1941; Böhlke and Randall 1968; Hastings 1979; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Smith 1997; Gon 2002a.

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230 PERCIFORMES

Astrapogon puncticulatus Blackfin cardinalfish

(Poey, 1867)

Astrapogon puncticulatus has relatively long black pelvic fins and a dark brown head and body. Jaws, vomer, and palatine have narrow bands of villiform teeth. Preopercular ridge is smooth, and preoperculum is smooth along posterior margin and scalloped along ventral margin. Gill rakers on first arch number 4 or 5 on upper limb and 12 to 14 on lower limb. Measurements are expressed as percent of SL: head length 40%– 43%, snout length 8%–9%, eye diameter 13%–15%, upper jaw length 21%–25%, pelvic fin length 38%– 42%, body depth 37%–39%. Pectoral fin has 15 to 17 rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Caudal fin is slightly emarginate. Scales are cycloid and deciduous. Median predorsal scales are lacking.

Color is dark brown, with silvery and bronze reflections and dark bars that radiate from eye. Tips of second dorsal, anal, and caudal fins are white. This species occurs in shallow water in the western Atlantic from southern Florida and Bermuda to Brazil, including the eastern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is limited to clear water, is usually associated with sea-grass beds, and is often found in empty shells. Maximum known size is about 63 mm TL. references: Böhlke and Chaplin 1968; Böhlke and Randall 1968; Robins and Ray 1986; Cervigón 1993b; Smith 1997; Smith-Vaniz et al. 1999; Gon 2002a.

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APOGONIDAE 231

Astrapogon stellatus Conchfish

(Cope, 1867)

Astrapogon stellatus has a relatively long snout and very long and black pelvic fins. Jaws, vomer, and palatine have narrow bands of villiform teeth. Preopercular ridge is smooth, and preoperculum is smooth along posterior margin and scalloped along ventral margin. Gill rakers on first arch number 10 to 12 on lower limb. Measurements are expressed as percent of SL: head length 37%–39%, snout length 8%–9%, eye diameter 11%–12%, upper jaw length 21%–22%, pelvic fin length 34%– 43%, body depth 33%–37%. Pectoral fin has 14 or 15 rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Caudal fin is slightly truncate to slightly convex. Scales are cycloid and deciduous. Median predorsal scales are lacking. Color is brown to black, with head, body, and median fins pep-

pered with minute dark to small dark spots, and dark bars that radiate from eye. This species occurs in shallow water in the western Atlantic from Florida and Bermuda to the Bahamas and Venezuela, including the eastern and southern Gulf of Mexico and the Greater and Lesser Antilles. It lives in the mantle cavity of the queen conch (Strombus gigas), with as many as five individuals occurring within the same conch. This species feeds nocturnally on shrimps and other small crustaceans. Males brood the eggs. Maximum known size is about 72 mm TL. references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Böhlke and Randall 1968; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Randall 1996; Smith 1997; Smith-Vaniz et al. 1999; Gon 2002a.

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232 PERCIFORMES

Phaeoptyx conklini (Silvester, 1915) Freckled cardinalfish

Phaeoptyx conklini has dark stripes near bases of second dorsal fin and anal fin. Jaws, vomer, and palatine have narrow bands of villiform teeth. Preopercular ridge is smooth, and posterior margin of preoperculum is finely serrated, with fleshy posteroventral process extending to near margin of operculum. Gill rakers on first arch number 14 to 16 on lower limb. Measurements are expressed as percent of SL: head length 34%–37%, snout length 7%–9%, eye diameter 13%–14%, upper jaw length 20%– 22%, caudal peduncle length 23%–26%, body depth 32%–36%. Pectoral fin has 11 or 12 rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Inner pelvic fin ray is connected along its length to abdomen by membrane. Caudal fin is forked. Scales are ctenoid and deciduous. Median predorsal scales number 5, and circumpeduncular scales encircling caudal peduncle number 12. Color is

reddish to brownish and densely and uniformly covered with minute dark brown spots. Stripes on second dorsal fin and anal fin are separated from base by pale streak. Dark elongated blotch is located on caudal peduncle. This species occurs in clear, shallow water in the western Atlantic from southern Florida and Bermuda to Venezuela, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with empty conch shells, rubble, and empty containers. Maximum known size is about 65 mm SL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Böhlke and Randall 1968; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Randall 1996; Smith 1997; Smith-Vaniz et al. 1999; Gon 2002a.

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APOGONIDAE 233

Phaeoptyx pigmentaria Dusky cardinalfish

(Poey, 1860)

Phaeoptyx pigmentaria has some enlarged teeth in upper and lower jaws and a brownish body. Teeth of jaws, vomer, and palatine are villiform and in narrow bands except for some enlarged teeth in anterior section of upper and lateral section of lower jaw. Preopercular ridge is smooth, and posterior margin of preoperculum is finely serrated, with fleshy posteroventral process extending to near margin of operculum. Gill rakers on first arch number 11 to 13 on lower limb. Measurements are expressed as percent of SL: head length 32%–37%, snout length 7%–9%, eye diameter 10%–11%, upper jaw length 18%–20%, caudal peduncle length 23%–25%, body depth 31%–35%. Pectoral fin has 11 or 12 rays, first dorsal fin has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Inner pelvic fin ray is connected along its length to abdomen by membrane. Caudal fin is forked. Scales are ctenoid and deciduous. Median predorsal scales

number 5 or 6, and circumpeduncular scales number 16. Color is reddish to brownish and densely and uniformly covered with small dark brown spots. Dark elongated blotch is located on caudal peduncle. This species occurs in the tropical Atlantic. In the western Atlantic it occurs from near shore to 42 m, from southern Florida and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with sea-grass beds. Maximum known size is about 76 mm TL.

references: Longley and Hildebrand 1941; Caldwell and Briggs 1957; Böhlke and Chaplin 1968; Böhlke and Randall 1968; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Hastings 1979; Robins and Ray 1986; Boschung 1992; Randall 1996; Schaldach et al. 1997; Cervigón 1993b; Smith 1997; Smith-Vaniz et al. 1999; Gon 2002a.

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234 PERCIFORMES

Phaeoptyx xenus (Böhlke and Randall, 1968) Sponge cardinalfish

Phaeoptyx xenus has a brownish body and pelvic fins joined to abdomen by membrane. Jaws, vomer, and palatine have narrow bands of villiform teeth. Preopercular ridge is smooth, and posterior margin of preoperculum is finely serrated, with fleshy posteroventral process extending to near margin of operculum. Gill rakers on first arch number 12 to 15 on lower limb. Measurements are expressed as percent of SL: head length 40%– 43%, snout length 9%–11%, eye diameter 13%–14%, upper jaw length 22%–24%, caudal peduncle length 29%–31%, body depth 33%–35%. Pectoral fin has 11 rays, first dorsal has six slender spines, second dorsal fin has one spine and 9 rays, and anal fin has two spines and 8 rays. Inner pelvic fin ray is connected along its length to abdomen by membrane. Caudal fin is forked. Scales are ctenoid and deciduous.

Median predorsal scales number five to seven. Color is orangish to lavender brown, with lower section of body silvery and head yellow. Head and body are densely and uniformly covered with small dark brown spots. Dark elongated blotch is located on caudal peduncle. This species occurs from near shore to 50 m in the western Atlantic from southern Florida to Venezuela, including the eastern and northern Gulf of Mexico and the Bahamas. It is associated with coral reefs and rocky bottoms. Maximum known size is about 75 mm TL. references: Caldwell and Briggs 1957 (as Apogon pigmentarius); Böhlke and Chaplin 1968; Böhlke and Randall 1968; Hastings 1979; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Smith 1997; Gon 2002a.

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235

EPIGONIDAE

Deepwater cardinalfishes

Deepwater cardinalfishes are moderately elongate and fusiform to rather robust and compressed, with large round to oval eye, two short dorsal fins, and weak to strongly ctenoid scales. Snout is short to moderately long, and convex to straight in profile. Eye is round or oval. Mouth is terminal, oblique, and large. Upper jaw is protrusible, and maxilla is narrow and sheathed by lachrymal bone anteriorly. Supramaxilla is absent. Nostril is paired, and nares are rounded or slitlike. Teeth are absent or located in jaws, vomer, and palatine. Teeth are of variable shape but generally small. Rarely, several antrorse canine teeth are present in anterior section of upper and lower jaws. Preoperculum is double edged, and operculum is large and has one to three spines (Sphyraenops) and occasionally serrations on posterior margin. Pseudobranch is present on inner side of operculum. Gill rakers on first arch are moderately developed or occasionally long and lathlike. Branchiostegal rays number 7. Pectoral fin base is slightly below midline and strongly oblique. First dorsal fin originates slightly posterior to pectoral fin base and has six to eight slender spines. Second dorsal fin is distinctly separated from first, and has one slender spine and 8 to 11 rays. Anal fin originates distinctly posterior to second dorsal fin origin and has one to three slender spines and 7 to 10 rays. Caudal fin is forked and has several spiny procurrent rays. Body is covered with small to moderate-sized, deciduous to firmly attached scales. Second dorsal fin and anal fin are covered with scales in one genus only (Epigonus). Lateral line is complete, extends onto caudal fin, and runs parallel to dorsal profile of trunk. Occasionally lateral line scales on caudal peduncle are modified to form keel. Gas bladder is present and has welldeveloped rete mirabile and oval on posterodorsal surface. Pyloric caecae number 5 to 34. Vertebrae number 25: 10 or 11 precaudal and 14 or 15 caudal. Deepwater cardinalfishes occur worldwide over continental and insular slopes between 75 and 3,700 m. Adults of all species are thought to be benthopelagic, associated with the bottom but tending to hover above it. Juveniles are mesopelagic. There are about 30 species in five genera, 9 species in three genera in the western Central Atlantic, and 6 species in a single genus in the Gulf of Mexico. references: Mayer 1974; Johnson 1984, Johnson and Patterson 1993; Nelson 1994; Gon 2002b.

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236 PERCIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Mayer 1974) 1a. Opercular spine (ventralmost reinforced projection from posterodorsal edge of operculum) weak or absent . . . . . . 2 1b. Opercular spine strong and pungent. . . . . . . . . . . . . . . . . 5 2a. Lateral line scales 46 to 51; tongue without teeth or with scattered tooth patches . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2b. Lateral line scales 33 to 36; tongue covered with tooth patches . . . . . . . . . . . . . . . . . . . Epigonus oligolepis p. 240 3a. Gill rakers on first arch 23 to 34; premaxillary teeth short, conical, or villiform, not visible when mouth closed. . . . . 4 3b. Gill rakers on first arch 17 to 21; premaxillary teeth elongate, thin, and inwardly recurved, visible when mouth closed . . . . . . . . . . . . . . . . . . . . . Epigonus macrops p. 238 4a. Body depth 16% to 24% of SL; caudal peduncle length 25% to 32% of SL; caudal peduncle ring absent . . . . . . . . . . . . . . . . . . . . . . . Epigonus denticulatus p. 237 4b. Body depth 22% to 30% of SL; caudal peduncle length 22% to 26% of SL; caudal peduncle ring present in specimens less than 110 to 120 mm SL . . . . . . . . . . . . . . . . . . . . . . . . . Epigonus pandionis p. 241 5a. Body depth 20% to 32% of SL; second dorsal fin with 9 (rarely 10) rays; gill rakers on first arch 26 to 35 . . . . . . . . . . . . . . . . . . . . . . . . . Epigonus pectinifer p. 243 5b. Body depth 14% to 20% of SL; second dorsal fin with 10 (rarely 9) rays; gill rakers on first arch 22 to 27 . . . . . . . . . . . . . . . . . . . . . . . Epigonus occidentalis p. 239

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EPIGONIDAE 237

Epigonus denticulatus

Dieuzeide, 1950

Epigonus denticulatus has a long caudal peduncle and an opercular spine consisting of a membranous projection. Head is moderate to short. Snout is short and blunt. Maxilla extends to anterior third or half of eye. Teeth are small, conical, and arranged in one to four rows in jaws, palatine, and vomer. Tongue either lacks teeth or, rarely, has several isolated tooth patches. Preopercular angle is produced and broadly rounded, with striations radiating from inner edge. Gill rakers on first arch are simple and awl-like and number 28 to 34. Measurements are expressed as percent of SL: head length 31%–39%, head depth 16%– 20%, caudal peduncle length 25%–32%, body depth 16%–24%; and as percent of head length: eye diameter 40%– 48%. Pectoral fin has 18 to 20 rays. First dorsal fin has seven spines, and second dorsal fin has one spine and

10 rays (rarely one spine and 9 rays). Anal fin has two spines and 9 rays (rarely two spines and 8 rays). Lateral line scales number 46 to 49. Pyloric caecae number 10 to 14, and vertebrae number 25. Color is grayish brown over most of body and silvery on belly, with dorsal and caudal fins yellowish. In preservative, color is brownish black and slightly darker dorsally than ventrally. This species occurs between 200 and 830 m near the bottom in the Atlantic and western Pacific Oceans. In the western Atlantic it occurs in the northern Gulf of Mexico, in the Greater Antilles, and along the coast of Central America. Juveniles occur between 130 and 145 m. Maximum known size is 188 mm SL. references: Mayer 1974; Masuda et al. 1984; Gon 2002b.

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238 PERCIFORMES

Epigonus macrops

(Bauer, 1906)

Epigonus macrops has long premaxillary teeth and a weak opercular spine. Head is moderate to short. Snout is short and blunt. Maxilla extends to anterior one-third to two-fifths of eye length. Teeth are conical, often recurved, and arranged in a single row in upper jaw, in one or two rows in lower jaw, in two to four irregular rows in vomer, and in a single row in palatine. Teeth are lacking on tongue. Preopercular angle is weakly produced and rounded, with serrations radiating from inner edge. Operculum has short spine below 3 to 10 spinelets. Gill rakers on first arch are short and awl-like and number 17 to 21. Measurements are expressed as percent of SL: head length 34%–39%, head depth 17%–22%, interorbital width 10%– 12%, caudal peduncle length 22%–27%, body depth 20%–24%; and as percent of head length: eye diameter 40%– 48%. Pectoral fin has 18 or 19 rays. First dorsal fin has eight slender spines (rarely seven), and second dorsal fin has one slender spine and 10 rays (rarely

one spine and 9 rays). Anal fin has two slender spines and 10 rays (rarely two spines and 9 rays). Lateral line scales number 46 to 50. Pyloric caecae number eight, with one modified into luminescent organ. Vertebrae number 25: 10 precaudal and 15 caudal. Color is black in life and brownish gray in preservative. Juveniles have thin, black, anterodorsally oriented ring circling midportion of caudal peduncle. Mouth cavity lacks pigment in juveniles but is black in adults. This species occurs between 550 and 1,100 m in the western Atlantic and Indian Oceans. In the western Atlantic it occurs throughout the Gulf of Mexico, in the Lesser Antilles, and off Suriname. Pelagic juveniles have been collected between 120 and 550 m in the Caribbean Sea. Maximum known size is 206 mm SL. references: Mayer 1974; Fujii 1983b; Gon 2002b.

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EPIGONIDAE 239

Epigonus occidentalis

Goode and Bean, 1896

Epigonus occidentalis has a pungent, bony opercular spine and a moderately long caudal peduncle. Head is moderate to short. Snout is moderately short and blunt. Maxilla extends to anterior one-fourth to two-fifths of eye length. Teeth are conical, in one to four rows in jaws and in vomer, and in a single row in palatine. Tongue lacks teeth. Preopercular angle is produced and rounded or pointed and bears serrations and striations. Opercular spine is below one to three poorly ossified spinelets. Gill rakers are short and awl-like and number 22 to 27 on first arch. Measurements are expressed as percent of SL: head length 31%–38%, head depth 13%–17%, interorbital width 6%– 9%, caudal peduncle length 22%–28%, body depth 14%–20%; and as percent of head length: eye diameter 41%–52%. Pectoral fin has 19 to 21 rays. First dorsal fin has seven

slender spines, and second dorsal fin has one slender spine and 10 (rarely 9) rays. Anal fin has two spines and 9 rays (rarely two spines and 8 rays). Lateral line scales number 46 to 51. Pyloric caecae number 8 to 13. Vertebrae number 15: 10 precaudal and 15 caudal. Color is blackish brown, with dorsal, pelvic, and anal fins yellowish and caudal fin grayish brown. In preservative, color is light grayish brown. Mouth cavity lacks pigment in juveniles but is black in adults. This species occurs between 360 and 737 m in the western Atlantic from the northern and western Gulf of Mexico and throughout the Caribbean Sea. Maximum known size is 179 mm SL.

references: Mayer 1974; Gon 2002b.

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240 PERCIFORMES

Epigonus oligolepis

Mayer, 1974

Epigonus oligolepis has a weak, poorly ossified opercular spine and few or no teeth in upper jaw. Head is moderate to long. Snout is moderately long and pointed. Maxilla extends from anterior two-fifths to one-half of eye length. Teeth are small. Lower jaw has one or two rows, vomer has irregular tooth patches, and palatine has one to multiple tooth rows. Tongue has tooth patches. Preopercular angle is rectangular or lightly produced. Opercular spine is below two to six membranous spinelets and separated from spinelets by gap. Gill rakers are simple and awl-like and number 29 to 31. Measurements are expressed as percent of SL: head length 34%– 43%, head depth 17%–19%, interorbital width 9%–10%, caudal peduncle length 24%–27%, body depth

20%–25%; and as percent of head length: eye diameter 40%– 44%. Pectoral fins have 16 to 18 rays. First dorsal fin has seven spines, and second dorsal has one spine and 10 rays. Anal fin has two spines and 9 rays (rarely two spines and 8 rays). Lateral line scales number 33 to 36. Pyloric caecae number 8 to 10. Vertebrae number 25: 10 precaudal and 15 caudal. Color is gray to black in life, and slightly darker dorsally than ventrally. In preservative, specimens are light and variable in color. This species occurs between 380 and 660 m in the western Atlantic off southern Florida, in the northern Gulf of Mexico, and in the Lesser Antilles. Maximum known size is 127 mm SL. references: Mayer 1974; Gon 2002b.

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EPIGONIDAE 241

Epigonus pandionis (Goode and Bean, 1881)

Epigonus pandionis has a short, poorly ossified opercular spine and a relatively short caudal peduncle. Head is of moderate length. Snout is moderately short and blunt. Maxilla extends to anterior two-fifths to one-half of eye length. Teeth are small, conical, and occasionally recurved, and occur in one or more rows in jaws and palatine and in two chevronlike clumps in vomer. Tongue lacks teeth. Preopercular angle is moderately produced and broadly rounded. Opercular spine is below two to five poorly ossified spinelets and is separated from spinelets by gap. Gill rakers are awllike and number 26 to 30 on first arch. Measurements are expressed as percent of SL: head length 33%–39%, head depth 19%–22%, interorbital width 9%–12%, caudal peduncle length 22%–26%, body depth 22%–30%; and as percent of head length: eye diameter 37%– 49%. Pectoral fin has 17 to 19 rays. First dorsal has seven (rarely eight) spines, and second dorsal fin has one spine and 10 rays (rarely one

spine and 9 rays, one spine and 11 rays, or two spines and 10 rays). Anal fin has two spines and 9 rays (rarely two spines and 8 rays or two spines and 10 rays). Lateral line scales number 46 to 49. Pyloric caecae number 10 to 13. Vertebrae number 15: 10 precaudal and 15 caudal. Color is grayish tan. Juvenile specimens, less than 85 mm SL, have a thin, black, posterodorsally sloped ring on central portion of caudal peduncle and a broad black band on posterior end of caudal peduncle. This species occurs between 210 and 600 m in the tropical Atlantic Ocean. In the western Atlantic it ranges from New Jersey to northern Brazil, including the Gulf of Mexico and the Caribbean coast of Central and South America. It is found throughout the Gulf of Mexico within its depth range. Juveniles are pelagic between 275 and 300 m. Maximum known size is 194 mm SL. references: Mayer 1974; Fujii 1983b; Gon 2002b.

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242 PERCIFORMES

Epigonus pectinifer

Mayer, 1974

Epigonus pectinifer has a well-ossified and pungent opercular spine and a narrow caudal peduncle. Head is short to moderate in length. Snout is long and pointed, and lower jaw extends slightly beyond upper jaw. Maxilla extends to anterior one-third to one-half of eye length. Teeth are small, conical, and arranged in a single row in jaws and palatine but may be absent in upper jaw and palatine. Vomer has small patch of strong teeth. Tongue generally has teeth. Preopercular angle is narrowly produced and serrated. Opercular spine is below two or three poorly ossified spinelets and separated from spinelets by large gap. Gill rakers number 26 to 35 and are pectinate and bear nublike projections along mesial surface. Measurements are expressed as percent of SL: head length 31%–36%, head depth 14%–17%, interorbital width 8%–9%, caudal peduncle length 25%–29%, body depth 20%–32%; and

as percent of head length: eye diameter 39%– 45%. Pectoral fin has 15 to 18 rays. First dorsal fin has seven spines, and second dorsal fin has one spine and 9 (rarely 10) rays. Anal fin has two spines and 9 rays. Lateral line scales number 47 to 49. Pyloric caecae number five to seven. Vertebrae number 25: 10 precaudal and 15 caudal. Color is brownish black on upper two-thirds of body and whitish on lower third, with fin membranes black and mouth light. This species occurs between 280 and 550 m in the western Atlantic and western Pacific Oceans. In the western Atlantic it occurs in the northern Gulf of Mexico and along the Caribbean coast of Yucatán and South America. Maximum known size is 153 mm SL.

references: Mayer 1974; Masuda et al. 1984; Gon 2002b.

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243

MALACANTHIDAE

Tilefishes

Tilefishes are oblong and compressed to elongate and subcylindrical in profile, with a relatively long, continuous dorsal fin, a long anal fin, and a single opercular spine. Snout is relatively long and ranges from slightly oblique to almost vertical. Species with nearly vertical snout have predorsal ridge. Nostril is paired, and anterior and posterior nares are close set and about equal in size. Mouth is terminal to slightly subterminal, moderately large, and slightly to strongly oblique. Villiform and several canine teeth occur in jaws but are absent in vomer and palatine. Preoperculum is angular, either has or lacks serrations along margin, and either has or lacks spine at angle. Operculum has broad, flat, blunt to sharp spine along upper posterior margin. Branchiostegal rays number 6. Gill membranes are separate from isthmus. Slit is present behind last (fourth) gill arch. Gill rakers on first arch number 18 to 28. Pectoral fin is well developed and inserts on lower flank. Dorsal fin is continuous and has 1 to 10 spines and 14 to 55 rays. Pelvic fin inserts under pectoral fin base and has 1 spine and 5 rays. Anal fin has 1 or 2 weak spines and 11 to 55 rays. All fin spines are slender and rather weak. Caudal fin is truncate to forked, with 17 principal rays and 15 branched rays. Body is covered with ctenoid (rarely cycloid) scales. Lateral line is complete. Axillary process on base of pelvic fins is absent. Vertebrae number 24 to 27: 10 or 11 precaudal and 14 caudal. Tilefishes occur worldwide in tropical to temperate seas from the shoreline to 500 m. Species are closely associated with the bottom, which they burrow into. Postlarvae are distinguished by their long, serrated head spines. Food consists of polychaetes, mollusks, shrimps, crabs, and squids. There are 39 species in five genera, 10 species in three genera in the western Central Atlantic, and 6 species in three genera in the Gulf of Mexico. references: Dooley 1978, 2001, 2002; Nelson 1994; Smith 1997; Smith-Vaniz et al. 1999.

Key to the Species of the Gulf of Mexico (Adapted from Dooley 1978) 1a. Predorsal ridge or flap present; body depth 21% to 36% of SL; total dorsal fin elements (spines and rays) 22 to 36; total anal fin elements 14 to 28 . . . . . . . . . . . . . . . . . . . . 2 1b. Predorsal ridge or flap absent; body depth 12% to 36% (usually 18%) of SL; total dorsal fin elements (spines and rays) 22 to 64; total anal fin elements 14 to 56 . . . . . . . . . . . . . . . . . . . . . . . Malacanthus plumieri p. 250

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244 PERCIFORMES

2a. Operculum with strong, pointed spine; dorsal fin with 6 to 10 spines and 22 to 27 rays; anal fin with 1 or 2 spines and 20 to 26 rays; pored lateral line scales 73 to 115 . . . . . . . 3 2b. Operculum with soft, blunt tab rather than strong spine; dorsal fin with 6 to 10 spines and 14 to 16 rays; anal fin with 1 or 2 spines and 11 to 14 rays; pored lateral line scales 47 to 75. . . . . Lopholatilus chamaeleonticeps p. 249 3a. Caudal fin margin double emarginate or rounded . . . . . . . . . . . . . . . . . . . . . Caulolatilus intermedius p. 247 3b. Caudal fin margin emarginate or truncate . . . . . . . . . . . . 4 4a. Dorsal fin with 8 spines and 23 to 25 rays; anal fin with 2 spines and 22 to 24 rays; caudal fin emarginate; broad golden patch between eye and nostril . . . . . . . . . . . . . . . . . . . . . . . Caulolatilus chrysops p. 245 4b. Dorsal fin with 7 or 8 (usually 7) spines and 23 to 27 rays; anal fin with 1 or 2 spines and 20 to 24 rays; caudal fin emarginate or truncate; without broad golden patch between eye and nostril . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5a. Upper jaw extending to anterior margin of orbit; caudal fin deeply emarginate; orbit diameter 23% to 41% of head length; spinous section of dorsal fin bright orangish yellow; upper section of body with dark markings and dark line below dorsal fin; large dark blotch above pectoral fin axil; bright yellow predorsal ridge . . . . . . . . . . . . . . . . . . . . . . . . Caulolatilus cyanops p. 246 5b. Upper jaw not extending to anterior margin of orbit; caudal fin truncate to slightly emarginate; orbit diameter 15% to 29% of head length; spinous section of dorsal fin dusky; upper section of body uniformly pigmented; no dark blotch above pectoral fin axil; dark predorsal ridge . . . . . . . . . . . . . . . . . . . . . . . . Caulolatilus microps p. 248

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MALACANTHIDAE 245

Caulolatilus chrysops Goldface tilefish

(Valenciennes, 1833)

Caulolatilus chrysops is elongate and elliptical in profile, with a predorsal ridge, a welldeveloped opercular spine, and an emarginate caudal fin. Dorsal profile of head is moderately convex. Mouth is terminal, with maxilla extending to near anterior margin of orbit. Posterior naris is enclosed in tube possessing flap. Jaws have outer row of 28 to 32 canine teeth, including 1 or 2 large antrorse teeth posteriorly, and a patch of villiform teeth on either side of symphyses extending as a single row posteriorly. Preoperculum has serrae on posterior margin. Operculum has a single flat spine directed posteriorly. Gill rakers on first arch number 17 to 21. Measurements are expressed as percent of SL: head length 26%–30%, predorsal length 29%–34%, body depth 27%– 30%; and as percent of head length: snout length 28%– 46%, upper jaw length 33%– 42%, orbit diameter 19%–30%. Pectoral fin is long and pointed and has 18 or 19 rays. Dorsal fin originates over pectoral fin base and has eight spines and 23 to 25 rays. Anal fin has two spines and 22 to 24 rays. Upper lobe of caudal fin is slightly longer than lower lobe.

Scales are ctenoid on body and cycloid on head. On head, scales extend to anterior margin of orbit. Caudal fin is covered with small scales. Lateral line runs parallel to dorsal profile and consists of 82 to 91 pored scales. Rows of cheek scales number 10 to 15. Color in life is violet, with light yellow sheen dorsally and silvery to pearly white ventrally. Predorsal midline is dark, a broad bright yellow bar runs from nostril to suborbital region, and a black spot marks axil of pectoral fin. This species occurs between 90 and 192 m in the western Atlantic from North Carolina to Brazil, including the Gulf of Mexico and the Greater Antilles. In the Gulf of Mexico it occurs in the Dry Tortugas and off Louisiana, Texas, and Veracruz. It is generally associated with rubble-covered substrates. Food consists of bivalves, polychaetes, sea urchins, and crabs. Maximum known size is 540 mm TL. references: Dooley 1978, 2002; Hoese and Moore 1977, 1998; Burgess and Branstetter 1985; Robins and Ray 1986; Cervigón 1993b; Schaldach et al. 1997; Smith 1997.

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246 PERCIFORMES

Caulolatilus cyanops Blackline tilefish

Poey, 1866

Caulolatilus cyanops is elongate and elliptical in profile, with a darkly pigmented predorsal ridge, a well-developed opercular spine, and a deeply emarginate caudal fin. Dorsal profile of head is moderately convex. Mouth is terminal, with upper jaw extending to near anterior margin of orbit. Posterior naris is enclosed in tube possessing flap. Upper jaw has outer row of 17 or 18 canine teeth, and lower jaw has 15 canine teeth. Both jaws have one antrorse tooth in corner and a patch of villiform teeth on either side of symphysis. Preoperculum has fine serrae along posterior margin and coarse serrae along ventral margin. Operculum has well-developed spine about half the length of pupil diameter. Gill rakers on first arch number 17 to 21. Measurements are expressed as percent of SL: head length 27%–32%, predorsal-fin length 30%–35%, body depth 25%– 31%; and as percent of head length: snout length 27%– 41%, upper jaw length 33%– 41%, orbit diameter 23%– 41%. Pectoral fin is long and pointed and has 16 to 18 (usually 16) rays. Dorsal fin originates slightly posterior to pectoral fin base and has seven spines and

23 or 24 rays. Anal fin has one or two spines and 20 to 23 rays. Scales are ctenoid on body and cycloid on head. On top of head, scales extend to anterior margin of orbit. Caudal fin is covered with small scales. Lateral line runs parallel to dorsal profile and consists of 75 to 82 pored scales. Cheek scale rows number 8 to 13. Color in life is blue to violet dorsally, with silvery yellow sheen and chainlike pattern, and milky white ventrally. Black stripe runs below length of dorsal fin, and black spot is located on axil of pectoral fin. Spinous section of dorsal fin is orangish yellow, and bases of caudal fin lobes are yellow. This species occurs between 45 and 495 m in the western Atlantic from North Carolina to northern South America, including the Gulf of Mexico and the Greater and Lesser Antilles. It is found in the northern and southern Gulf of Mexico. Maximum known size is 342 mm SL. references: Longley and Hildebrand 1941; Dooley 1978, 2002; Castro-Aguirre and MárquezEspinoza 1981; Robins and Ray 1986; Dennis and Bright 1988a; Boschung 1992; Smith 1997.

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MALACANTHIDAE 247

Caulolatilus intermedius Anchor tilefish

Howell Rivero, 1936

Caulolatilus intermedius is elongate and elliptical in profile, with a darkly pigmented predorsal ridge, a well-developed opercular spine, and a rounded caudal fin possessing extended tips. Dorsal profile of head is moderately convex. Mouth is terminal, with upper jaw extending to anterior margin of orbit. Posterior naris is enclosed in a tube possessing a flap. Jaws have outer row of 35 canine teeth, including one large antrorse tooth at end of row, and a patch of villiform teeth on either side of symphyses. Preoperculum is finely serrated on posterior margin. Operculum has a broad spine about half the length of pupil diameter. Gill rakers on first arch number 18 to 22. Measurements are expressed as percent of SL: head length 27%–31%, pre-dorsal-fin length 30%– 36%, body depth 26%–32%; and as percent of head length: snout length 29%– 40%, upper jaw length 31%–37%, orbit diameter 26%– 40%. Pectoral fin is long and pointed and has 16 or 17 rays. Dorsal fin originates over middle of pectoral fin base and has seven spines and

24 to 26 rays. Anal fin has one or two spines and 22 or 23 rays. Upper and lower tips of caudal fin extend beyond remainder of fin. Scales are ctenoid on body and cycloid on head. On top of head, scales extend to near anterior margin of orbit. Caudal fin is covered with small scales. Color is light violet brown dorsally and white ventrally. Black anchor-shaped mark is located on nape, and a dark stripe runs from eye to maxilla. Dorsal fin has series of dark blotches forming an irregular band. Black spot is located on axil of pectoral fin. This species occurs between 45 and 290 m in the western Atlantic from the Gulf of Mexico and Cuba. In the Gulf of Mexico it is known from northwestern Florida to Yucatán. It is associated with mud bottoms. Maximum known size is 253 mm SL. references: Hoese 1958; Hoese and Moore 1977, 1998; Dooley 1978, 2002; Matsuura 1983d; Robins and Ray 1986; Boschung 1991; Schaldach et al. 1997; Smith 1997.

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248 PERCIFORMES

Caulolatilus microps Blueline tilefish

Goode and Bean, 1878

Caulolatilus microps is elongate and elliptical in profile, with a well-developed opercular spine, no dark spot on axil of pectoral fins, and a truncate caudal fin. Dorsal profile of head is moderately convex. Mouth is terminal, with upper jaw extending half the distance between posterior nostril and anterior margin of orbit. Posterior naris is enclosed in a tube possessing a flap. Jaws have outer row of 20 to 32 canine teeth, including one or two large antrorse teeth at end of rows, and a patch of villiform teeth on either side of symphyses extending as a single row posteriorly. Preoperculum has welldeveloped serrae on posterior margin. Operculum has a single flat spine about three-fourths the length of pupil. Gill rakers on first arch number 21 to 27. Measurements are expressed as percent of SL: head length 28%–32%, predorsal-fin length 30%–39%, body depth 24%– 31%; and as percent of head length: snout length 37%–51%, upper jaw length 35%– 42%, orbit diameter 15%–29%. Pectoral fin is long and pointed, reaches to anus, and has 17 or 18 rays. Dorsal fin originates over poste-

rior aspect of pectoral fin base and has seven or eight spines and 24 to 27 rays. Anal fin has two spines and 22 to 24 (usually 22 or 23) rays. Caudal fin is truncate, with upper and lower tips extending beyond remainder of fin. Scales are ctenoid on body and cycloid on head. On top of head, scales extend to anterior margin of orbit. Caudal fin is covered with small scales. Lateral line is parallel to dorsal profile and consists of 79 to 91 scales. Color is dark brown dorsally and light beige white ventrally. Head has a black predorsal ridge and a narrow gold stripe underlined by blue line that runs from tip of snout to eye. This species occurs between 30 and 130 m, from Virginia to Yucatán, including the entire Gulf of Mexico. It is associated with rubblecovered substrates. Maximum known size is 653 mm SL.

references: Hoese and Moore 1977, 1998; Dooley 1978, 2002; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997.

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MALACANTHIDAE 249

Lopholatilus chamaeleonticeps Tilefish

Goode and Bean, 1879

Lopholatilus chamaeleonticeps is elongate and quadriform in profile, with an elevated predorsal ridge forming an enlarged flap, and a truncate caudal fin. Dorsal profile of head is strongly convex. Predorsal flap develops at about 80 mm SL. Mouth is terminal, with maxilla extending to anterior margin of orbit. Posterior nostril is enclosed in a tube possessing a flap. Upper jaw has row of 13 to 19 enlarged canines, and lower jaw has 15 to 17; both jaws have patch of villiform teeth on either side of symphyses. Preoperculum has a finely serrate posterior margin, an indention at corner, and an entire lower margin. Operculum has a single blunt, poorly ossified spine. Gill rakers on first arch number 22 to 26. Measurements are expressed as percent of SL: head length 28%–35%, pre-dorsal-fin length 30%–39%, body depth 21%–31%; and as percent of head length: snout length 27%–51%, upper jaw length 37%–53%, orbit diameter 16%– 44%. Pectoral fin is long and pointed and has 16 to 18 rays. Dorsal fin originates over pectoral fin base and has seven spines and 15 rays. Anal fin has two spines and 13 or 14 rays. Upper and

lower corners of caudal fin extend beyond remainder of fin. Scales are ctenoid on body and cycloid on head. Cheek scale rows number 6 to 10. Caudal fin is covered with small scales. Lateral line forms low curve, and pored lateral line scales number 66 to 75. Color is steely bluish gray dorsally, with covering of golden spots, and milky white ventrally. Predorsal flap is yellow, dorsal and anal fins are dark with large yellow spots, and caudal fin is dark with eight or nine yellow vertical bands. This species occurs between 81 and 540 m in the western Atlantic from Nova Scotia to Suriname, including the entire Gulf of Mexico. It is associated with mud and sand bottoms, which it burrows into. Food consists of bivalve mollusks, squids, polychaetes, holothurians, decapod crustaceans, elasmobranchs, and ray-finned fishes. Maximum known size is about 1,000 mm SL. references: Bigelow and Schroeder 1947, 1953; Hoese 1958; Hoese and Moore 1977, 1998; Dooley 1978, 2002; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997.

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250 PERCIFORMES

Malacanthus plumieri (Bloch, 1786) Sand tilefish

Malacanthus plumieri is elongate and subcylindrical in profile, with very long dorsal and anal fins, and a forked caudal fin possessing elongate filaments on upper and lower corners. Dorsal profile is weakly convex. Mouth is terminal, with fleshy lips. Maxilla terminates distinctly anterior to anterior margin of orbit. Posterior naris is enclosed in a tube with a flap. Upper jaw has three or four small, anteriorly curved canines, followed by 15 to 17 large teeth and one large, anteriorly curved canine. Lower jaw has four or five anteriorly curved canines, followed by four enlarged recurved canines, 6 to 9 small teeth, and one enlarged anteriorly curved canine. Both jaws have patches of villiform teeth on either side of symphyses. Preoperculum lacks serrations on posterior and ventral margins. Operculum has one welldeveloped, sharp spine about three-fourths the length of eye. Gill rakers on first arch number 8 to 13. Measurements are expressed as percent of SL: head length 23%–28%, pre-dorsalfin length 23%–27%, body depth 13%–19%; and as percent of head length: snout length 39%–52%, upper jaw length 35%– 42%, orbit diameter 11%–25%. Pectoral fin is broad and pointed. Dorsal fin originates over pectoral fin base and has four or five weak spines and 54 to 60 rays. Anal fin has a single spine, followed by 1 unbranched and 48 to 55 branched rays.

Scales are ctenoid on most of body but cycloid on thoracic region and head. On top of head, scales extend to middle of orbit. Pored lateral line scales number 135 to 152. Color is gray to tan with bluish cast dorsally and white ventrally. Fins are yellowish, except caudal fin has orangish lobes and dusky area above center of fin. This species occurs in the western and Central Atlantic. In the western Atlantic it occurs between 10 and 153 m, from Cape Lookout, NC, and Bermuda to Uruguay, including the entire Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with sandy, rough bottoms, where it constructs burrows out of broken coral and rubble. Food consists of polychaetes, sipunculids, echinoids, ophiuroids, chitons, amphipods, stomatopods, shrimps, crabs, and ray-finned fishes. Maximum known size is 600 mm SL.

references: Longley and Hildebrand 1941; Baughman 1947, 1950b; Hoese 1958; Randall 1968a, 1996; Böhlke and Chaplin 1968; Clifton and Hunter 1972; Colin 1973; Bright and Cashman 1974; Smith et al. 1975; Hoese and Moore 1977, 1998; Dooley 1978, 2002; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith-Vaniz et al. 1999.

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251

POMATOMIDAE

Bluefishes

This family is monotypic and occurs in the Gulf of Mexico.

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252 PERCIFORMES

Pomatomus saltatrix Bluefish

(Linnaeus, 1766)

Pomatomus saltatrix is streamlined, moderately elongate, and moderately compressed, with a large head, a prominent lower jaw, and separate dorsal fins. Dorsal profile is moderately convex. Nostril is paired, and nares are close set, with posterior naris broader than anterior naris. Eye is of moderate size and laterally located. Mouth is terminal, with lower jaw slightly projecting beyond upper jaw, and upper jaw extending beyond posterior margin of orbit. Teeth occur in jaws, vomer, and palatine. Jaws have a single row of prominent, sharp, and compressed teeth. Vomerine and palatine teeth are villiform. Preoperculum has a smooth margin and a membranous flap that covers suboperculum. Operculum has a broad, flat spine on posterior margin. Branchiostegal rays number 7. Pseudobranch on inner aspect of gill cover is well developed. Gill membranes are separate and free of isthmus. Four gill arches are present, with slit behind fourth arch. Gill rakers on first arch number 3 or 4 on upper limb and 9 to 11 on lower limb. Measurements are expressed as percent of SL: head length 29%–33%, snout length 7%–9%, eye diameter 5%– 6%, upper jaw length 13%–15%, pectoral fin length 17%–20%, body depth 26%– 30%. Pectoral fin is relatively short and acutely tipped and has 16 to 18 rays. Spinous dorsal fin has seven to nine short, weak spines and is depressible into groove. Second dorsal fin has a single relatively long spine and 23 to 28 rays. Pelvic fin inserts slightly behind pectoral fin base and consists of one spine and 5 rays. Anal fin has two spines and 23 to 27 rays. Caudal fin is broad and moderately forked. Body, cheek, and operculum are covered with small, decid-

uous spinoid scales. Bases of second dorsal fin and anal fin are covered with scales. Lateral line is slightly arched over pectoral fin and consists of about 95 scales. Vertebrae number 26: 11 precaudal and 15 caudal. Color is greenish to bluish dorsally and silvery on side and ventrally, with black blotch at base of pectoral fin. Dorsal and anal fins are pale green with yellow tinge, and caudal fin is greenish with yellow tinge. This species occurs mostly over continental shelves in subtropical to warm temperate waters worldwide, with exception of the eastern Pacific and the western Pacific north of the equator. It is divided into eight major populations, and the one in the western Atlantic occurs from Nova Scotia and Bermuda to Yucatán and northern Cuba, and from Colombia to Argentina, including the entire Gulf of Mexico. Fecundity reaches to over one million eggs in large specimens, and spawning occurs from March to September. Eggs and larvae are pelagic. This species is a ravenous predator, and prey consist mostly of squids and of schooling ray-finned fishes, such as engraulids, clupeids, atherinids, sciaenids, scombrids, and stromateids. Maximum known age is about 12 years, and maximum known size is 115 cm TL.

references: Hastings and Bortone 1976; Hastings et al. 1976; Hoese and Moore 1977, 1998; Johnson 1984; Robins and Ray 1986; Boschung 1992; Roberts 1993; Nelson 1994; Goodbred and Graves 1996; Schaldach et al. 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Collette 2001a, 2002d; Klein-MacPhee 2002a.

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253

ECHENEIDAE

Remoras

Remoras are elongate and fusiform, with a flat head and a laminated, oval-shaped cephalic disc derived from the first dorsal fin. Head is wide and depressed. Lower jaw projects beyond upper jaw, and upper jaw is not protrusible. Nostril is paired, and nares are close set. Mouth is wide and horizontal, and jaw teeth are villiform and arranged in bands. Villiform teeth occur in vomer and occasionally in palatine and on tongue. Four gill arches are present, with slit behind fourth. Gill rakers on first arch are short. Gill membranes are not connected and are free of isthmus. Branchiostegal rays number 8 to 11. Pectoral fin is short, has an oblique base, inserts high on flank, and has 18 to 32 rays. First dorsal fin is modified into cephalic disc, with spines split to form 10 to 28 transverse mobile laminae inside fleshy margin. Second dorsal fin has 18 to 45 rays. Pelvic fin inserts below pectoral fin insertion and has one spine and 5 rays. Anal fin has 18 to 41 rays and is similar in shape and location to second dorsal fin. Caudal fin is forked, emarginate, or convex. Body is covered with small cycloid scales that are barely imbricate and usually embedded in skin. Lateral line is complete and nearly straight. Gas bladder is absent, and vertebrae number 26 to 40: 12 to 18 precaudal and 14 to 22 caudal. Remoras occur worldwide in tropical to warm temperate seas. They attach to sharks, rays, tarpons, barracudas, seabasses, jacks, cobia, parrotfishes, sailfishes, marlins, swordfishes, molids, dolphins, and whales by means of their adhesive discs. Some species, such as Remora osteochir and Remorina albescens, are specific to certain hosts. The former is associated only with billfishes, and the latter is generally found on rays. Individuals of these two species also attach to ships and other floating objects. Some species of Echeneis are often free swimming in shallow, inshore water. In addition to hitching a ride, they remove ectoparasites (copepods) from their hosts and feed on remnants of prey captured by their hosts. There are eight species in four genera, and all occur in the western Central Atlantic and the Gulf of Mexico. references: Jordan and Evermann 1898; Lachner 1978, 1986; Fritzsche and Fuiman 1982; Johnson 1984; Heemstra 1986k; Nelson 1994; Tominaga et al. 1996; Smith-Vaniz et al. 1999; Collette 2001b, 2002a, 2003b.

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254 PERCIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Lachner 1978; Smith-Vaniz et al. 1999) 1a. Body depth 7.1% to 12.5% of SL; pectoral fin pointed in adults; side with dark longitudinal band bordered by white band; anal fin rays 29 to 41; caudal fin lanceolate in juveniles, with middle rays filamentous, and truncate in adults, with lobes produced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Body depth 12.5% to 20% of SL; pectoral fin rounded in juveniles and adults; side without longitudinal band; anal fin rays 18 to 28; caudal fin forked in juveniles and emarginate to truncate in adults . . . . . . . . . . . . . . . . . . . . . . . . 4 2a. Cephalic disc small, with 9 to 11 lamellae . . . . . . . . . . . . . . . . . . . . . . Phtheirichthys lineatus p. 257 2b. Cephalic disc large, with 18 to 28 lamellae . . . . . . . . . . . 3 3a. Lamellae of cephalic disc 18 to 23 (usually 21); dorsal fin rays 32 to 41 (usually 36); anal fin rays 30 to 38 (usually 33); more than tips of dorsal, anal, and caudal fins white . . . . . . . . . . . . . . . . . . . . . . . Echeneis neucratoides p. 256 3b. Lamellae of cephalic disc 21 to 28 (usually 23); dorsal fin rays 33 to 45 (usually 39); anal fin rays 31 to 41 (usually 36); only tips of dorsal, anal, and caudal fins white . . . . . . . . . . . . . . . . . . . . . . . . . Echeneis naucrates p. 255 4a. Pelvic fin narrowly attached to body; lamellae of cephalic disc 12 to 14; color pale to whitish . . . . . . . . . . . . . . . . . . . . . . . . . Remorina albescens p. 262 4b. Pelvic fin broadly attached to body; lamellae of cephalic disc 15 to 20; color light to dark brown. . . . . . . . . . . . . . 5 5a. Gill rakers on lower limb of first arch 26 to 28 . . . . . . . . . . . . . . . . . . . . . . . . . . . . Remora remora p. 261 5b. Gill rakers on lower limb of first arch 12 to 14. . . . . . . . . 6 6a. Lamellae of cephalic disc 24 to 28 . . . . . . . . . . . . . . . . . . . . . . . . . . . Remora australis p. 258 6b. Lamellae of cephalic disc 15 to 20 . . . . . . . . . . . . . . . . . . 7 7a. Dorsal fin rays 27 to 34; distal two-thirds of length of pectoral fin rays flexible in specimens greater than 150 m SL; spinules of cephalic disc pointed and, in specimens greater than 150 mm SL, arranged in two or three irregular rows . . . . . . . . . . . . . . . . . . . . . . . . Remora brachyptera p. 259 7b. Dorsal fin rays 21 to 27; pectoral fin rays stiff to tips in specimens greater than 150 mm SL; spinules of cephalic disc blunt, nearly bifurcate or spatulate (especially in posterior row of each lamellae), and, in specimens greater than 150 mm SL, arranged in four irregular rows . . . . . . . . . . . . . . . . . . . . . . . . . . Remora osteochir p. 260

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ECHENEIDAE 255

Echeneis naucrates Sharksucker

Linnaeus, 1758

Echeneis naucrates is very elongate, with acutely pointed pectoral fin, narrow-based pelvic fin, and broad dark stripe on flank. Mouth is short, extends about half the distance from tip of snout to anterior margin of eye, and possesses well-developed fleshy tab on lower lip. Teeth in upper jaw are in a broad band anteriorly and a narrow band posteriorly. Teeth in lower jaw are in a broad band and are exposed when mouth is closed. Vomer and palatine have broad bands of teeth, and tongue has small granular teeth. Gill rakers on first arch number 1 to 3 on upper limb and 17 or 18 on lower limb. Branchiostegal rays number 8. Measurements are expressed as percent of SL: head length 18%–20%, snout length 8%–9%, eye diameter 3%–5%, upper jaw length 6%– 7%, pectoral fin length 12%–15%, disc length 27%–28%, pelvic fin length 11%–14%, body depth 8%–10%. Pectoral fin has 20 to 26 rays. Cephalic disc has 21 to 28 (usually 23) lamellae. Second dorsal fin has 32 to 41 rays. Anal fin has 29 to 41 rays. Caudal fin has a concave margin. Vertebrae number 30: 14 precaudal and 16 caudal. Color is slate to dark brownish gray, with dark brown stripe bordered by

white stripes along flank running from tip of snout to base of caudal fin. White stripes may be obscure in large specimens. Median fins are dark with light to white margins. This species occurs worldwide in tropical to warm temperate seas, except for the eastern Pacific. In the western Atlantic it occurs over continental and insular shelves from Nova Scotia and Bermuda to Uruguay, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It commonly swims free of hosts but attaches to a variety of large pelagic vertebrates, including sharks, ray-finned fishes, sea turtles, and occasionally humans and boats. Unlike some of the other species of echeneids, it always attaches to the outside surfaces of its host. Maximum known size is 900 mm SL.

references: Bigelow and Schroeder 1953; Böhlke and Chaplin 1968; Randall 1968a, 1996; Cressey and Lachner 1970; Hoese and Moore 1977, 1998; Lachner 1978, 1986; Heemstra 1986k; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Collette 2001b, 2002c,f.

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256 PERCIFORMES

Echeneis neucratoides Whitefin sharksucker

Zuiew, 1786

Echeneis neucratoides is elongate, with a moderately pointed pectoral fin, a narrow-based pelvic fin, and upper and lower margins of medial fins white. Mouth extends about twothirds of the distance from tip of snout to anterior margin of eye and has poorly developed fleshy tab on lower lip. Teeth in upper jaw are in a broad band anteriorly and a narrow band posteriorly. Teeth in lower jaw are in a broad band and are exposed when mouth is closed. Vomer and palatine have broad bands of teeth, and tongue has small granular teeth. Gill rakers on first arch number 1 to 3 on upper limb and 14 on lower limb. Branchiostegal rays number 8. Measurements are expressed as percent of SL: head length 19%–21%, snout length 8%–9%, eye diameter 3%– 4%, upper jaw length 6%–7%, pectoral fin length 15%– 17%, disc length 25%–28%, pelvic fin length 12%–14%, body depth 7%–11%. Pectoral fin

has an acute tip and 22 rays. Cephalic disc has 18 to 23 (usually 21) lamellae. Second dorsal fin has 32 to 41 (usually 36) rays. Anal fin has 30 to 38 (usually 33) rays. Caudal fin has a truncate margin. Color is dark brown to blackish in adults, with dark brown stripe bordered by white stripe along flank running from tip of snout to base of caudal fin. Stripe may be obscured by ground color in some specimens. Median fins are dark, with relatively broad light to white margins. This species is restricted to the western Atlantic from Massachusetts to northern South America in coastal waters, including the northern Gulf of Mexico, the Bahamas, and the Antilles. Maximum known size is 750 mm TL. references: Böhlke and Chaplin 1968; Lachner 1978; Robins and Ray 1986; Boschung 1992; Smith 1997, Collette 2002c.

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ECHENEIDAE 257

Phtheirichthys lineatus Slender suckerfish

(Menzies, 1791)

Phtheirichthys lineatus is very elongate, with an acutely pointed (lanceolate) pectoral fin, a small cephalic disc, and a narrow-based pelvic fin. Mouth extends about three-fourths of the distance from tip of snout to anterior margin of eye. Teeth in upper jaw are sharp pointed, recurved, and arranged in a band. Teeth in lower jaw are sharp pointed and recurved and are exposed when mouth is closed. Vomer and palatine have broad bands of teeth, and tongue has small granular teeth. Gill rakers on first arch number 11 to 13 on lower limb. Measurements are expressed as percent of SL: head length 15%–21%, snout length 6%– 8%, eye diameter about 3%, upper jaw length 5%– 7%, pectoral fin length 10%–12%, disc length 15%–28%, pelvic fin length 10%–12%, body depth 6%–7%. Pectoral fin has an acute tip and 18 to 21 rays. Cephalic disc has 9 to 11 lamellae. Second dorsal fin has 29 to 33 rays. Anal fin has 29 to 34 rays. Caudal fin has convex margin. Vertebrae number 40. Color is

dark brown to blackish dorsally and white ventrally, with a faint dark stripe on midflank bordered by narrow light stripes in large specimens. Margins of dorsal and anal fins are dark, and upper and lower margins of caudal fin are light. This species occurs worldwide in tropical to subtropical seas. In the western Atlantic it occurs from South Carolina and Bermuda to northern South America, including the northern and southern Gulf of Mexico. It attaches to bodies or enters gill cavities of large fishes. Sphyraena barracuda is the most common host, but sea turtles also serve as hosts. This species is occasionally free swimming. Maximum known size is 435 mm SL. references: Cressey and Lachner 1970; Hoese and Moore 1977, 1998; Lachner 1986; Heemstra 1986k; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Collette 2002c.

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258 PERCIFORMES

Remora australis Whalesucker

(Bennett, 1840)

Remora australis is relatively stout and depressed anteriorly, with a rounded pectoral fin, a broad-based pelvic fin, and a relatively large cephalic disc. Mouth is short and extends about two-thirds of the distance from tip of snout to anterior margin of eye. Teeth in upper jaw are in a broad band anteriorly and a narrow band posteriorly, with outer teeth enlarged. Lower jaw has villiform teeth arranged in a band, with outermost teeth slightly enlarged. Vomer has a narrow band of teeth, and tongue lacks teeth. Gill rakers on first arch number 17 to 20. Branchiostegal rays number 9. Measurements are expressed as percent of SL: head length about 25%, snout length about 13%, eye diameter about 2%, upper jaw length about 10%, pectoral fin length about 13%, disc length about 48%, pelvic fin length about 13%, body depth about 12%. Pectoral fin has a rounded margin and 22 to 24 rays.

Cephalic disc has 24 to 28 lamellae. Second dorsal fin has 23 to 26 rays. Anal fin has 24 to 26 rays. Caudal fin has a concave margin. Spinules on lamellae of cephalic disc are arranged in two rows, with those of anterior row compressed and bladelike and those of posterior row conical. Vertebrae number 27: 12 precaudal and 15 caudal. Color is dark brown or grayish brown and slightly darker ventrally, with light border along dorsal and anal fins. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs from the northern Gulf of Mexico to Brazil. It is associated with cetaceans. Maximum known size is 403 mm SL.

references: Hoese and Moore 1977, 1998; Lachner 1986; Heemstra 1986k; Robins and Ray 1986; Boschung 1992; Collette 2002c.

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ECHENEIDAE 259

Remora brachyptera (Lowe, 1839) Spearfish remora Remora brachyptera is relatively stout and depressed anteriorly, with a rounded pectoral fin, a broad-based pelvic fin, and a relatively small cephalic disc. Mouth is relatively large and extends about to anterior margin of eye. Teeth in upper jaw are villiform and arranged in a broad band, with some near symphysis enlarged. Lower jaw has villiform teeth arranged in a band, with outermost teeth slightly enlarged. Vomer has a narrow band of teeth, and tongue has a tooth patch. Gill rakers on first arch number 11 to 17. Branchiostegal rays number 9. Measurements are expressed as percent of SL: head length 24%–25%, snout length 10%–11%, eye diameter 3%– 4%, upper jaw length 9%–11%, pectoral fin length 14%–16%, disc length 31%–35%, pelvic fin length 12%–13%, body depth 13%–14%. Pectoral fin has a rounded margin and 23 to 27 rays, with distal two-thirds of rays flexible in specimens greater than 150 mm SL. Cephalic disc has 15 to 18 lamellae. Second dor-

sal fin has 27 to 34 rays. Anal fin has 22 to 28 rays. Caudal fin has a concave margin. Spinules on lamellae of cephalic disc are pointed and, in specimens longer than 150 mm SL, are arranged in two or three irregular rows per lamella. Color is dark brown to reddish brown, with margins of dorsal and anal fins lighter. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs from Nova Scotia to Brazil, including the northern and southern Gulf of Mexico. It is associated with billfishes (Xiphiidae) but also occurs on sharks and mollids (Masturus lanceolatus) and occasionally is free swimming. Food consists of parasitic copepods. Maximum known size is 260 mm SL. references: Cressey and Lachner 1970; Hoese and Moore 1977; Lachner 1986; Heemstra 1986k; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Collette 2002c,f.

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260 PERCIFORMES

Remora osteochir Marlinsucker

(Cuvier, 1829)

Remora osteochir is relatively stout and depressed anteriorly, with a relatively stiff, rounded pectoral fin and a broad-based pelvic fin. Mouth is relatively small and extends about two-thirds of the distance from tip of snout to anterior margin of eye. Teeth in upper jaw are villiform and arranged in a broad band, with some near symphysis slightly enlarged. Lower jaw has villiform teeth arranged in a band, with outermost teeth slightly enlarged. Vomer has a narrow band of teeth, and tongue has a tooth patch. Gill rakers on first arch number 11 to 17. Branchiostegal rays number 9. Measurements are expressed as percent of SL: head length 20%–22%, snout length 9%–10%, eye diameter about 2%, upper jaw length about 7%, pectoral fin length 16%–18%, disc length 41%– 45%, pelvic fin length 13%–14%, body depth 7%–10%. Pectoral fin has rounded margin and 20 to 24 rays.

Cephalic disc has 15 to 19 lamellae. Second dorsal fin has 20 to 26 rays. Anal fin has 20 to 25 rays. Caudal fin has concave margin. Spinules on lamellae of cephalic disc are blunt and, in specimens longer than 150 mm SL, are arranged in four irregular rows per lamella. Color is dark gray to black and darker dorsally than ventrally, with margins of dorsal and anal fins also dark. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs from the northern and southern Gulf of Mexico and from Bermuda to Brazil. It is associated with billfishes (Xiphiidae). Maximum known size is 386 mm SL. references: Cressey and Lachner 1970; Hoese and Moore 1977; Lachner 1986; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Collette 2002c,f.

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ECHENEIDAE 261

Remora remora Remora

(Linnaeus, 1758)

Remora remora is relatively stout and depressed anteriorly, with a rounded pectoral fin, a broad-based pelvic fin, and a relatively short cephalic disc. Mouth is relatively large and extends about to anterior margin of eye. Teeth in upper jaw are villiform and arranged in a broad band. Lower jaw has villiform teeth arranged in a band, with outermost teeth slightly enlarged and recurved. Vomer has a narrow band of villiform teeth, and tongue has a tooth patch. Gill rakers on first arch are relatively long and slender and number 28 to 37. Branchiostegal rays number 9. Measurements are expressed as percent of SL: head length 25%–29%, snout length 11%–13%, eye diameter 3%– 4%, upper jaw length 10%–12%, pectoral fin length 14%–18%, disc length 35%– 42%, pelvic fin length 15%–16%, body depth 13%–14%. Pectoral fin has a rounded margin and 25 to 32 rays. Cephalic disc has 16 to 20 lamellae. Sec-

ond dorsal fin has 21 to 27 rays. Anal fin has 20 to 24 rays. Caudal fin has a concave margin. Spinules on lamellae of cephalic disc are bluntly tipped. Vertebrae number 27. Color is uniformly brackish, blackish, or sooty. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs from Nova Scotia and Bermuda to Argentina, including the northern and southern Gulf of Mexico. It is associated with a wide variety of pelagic sharks, including Carcharhinus floridanus and Prionace glauca. Maximum known size is 618 mm SL. references: Cressey and Lachner 1970; Hoese and Moore 1977, 1998; Lachner 1986; Heemstra 1986k; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Collette 2002c,f.

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262 PERCIFORMES

Remorina albescens White suckerfish

(Temminck and Schlegel, 1850)

Remorina albescens is relatively stout and very depressed anteriorly, with a rounded pectoral fin and a relatively small and narrow-based pelvic fin. Mouth is relatively large and extends about to anterior margin of eye. Upper jaw has small, conical, and recurved teeth arranged in a broad band, with teeth in outer row recurved and cardiform. Lower jaw has small, conical, and recurved teeth arranged in a broad band, with outermost teeth largest. Vomer has a broad band of recurved, conical teeth, and tongue has a tooth patch. Gill rakers on first arch are relatively poorly developed and number 10 to 14. Measurements are expressed as percent of SL: head length 23%–26%, snout length 12%–14%, eye diameter 2%–3%, upper jaw length 10%–12%, pectoral fin length 16%–21%, disc length 34%–36%, pelvic fin length 10%–12%, body depth 11%–13%. Pectoral fin has a rounded margin and 18 to 21 rays. Cephalic disc has 12 to 14 lamellae. Sec-

ond dorsal fin has 18 to 23 rays. Anal fin has 18 to 24 rays. Caudal fin has a convex margin. Spinules on lamellae of adhesive disc are compressed and acutely tipped. Color is uniformly pale gray to bluish white. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs from Florida and Bermuda to Brazil, including the northern and southern Gulf of Mexico. It is associated with Manta birostra and has been observed with the anterior half of its body in the cloaca of its host. Other hosts include various species of sharks and Makaira sp. Maximum known size is 225 mm TL. references: Cressey and Lachner 1970; Hoese and Moore 1977, 1998; Lachner 1986; Heemstra 1986k; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Collette 2002c.

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263

RACHYCENTRIDAE

Cobia

This family is monotypic and occurs in the Gulf of Mexico.

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264 PERCIFORMES

Rachycentron canadum Cobia

(Linnaeus, 1766)

Rachycentron canadum is elongate and subcylindrical in cross section, with a broad, depressed head and first dorsal fin consisting of free spines and membranes. Snout is long and acute. Nostril is paired, and nares are close set, with a flap on posterior margin of anterior naris and on anterior margin of posterior naris. Mouth is terminal and nearly horizontal, with upper jaw extending to near anterior margin of orbit and lower jaw projecting beyond upper. Jaw teeth are villiform and arranged in bands. Vomer, palatine, and tongue have patches or bands of villiform teeth. Preopercular margin is membranous, and opercular margin lacks spine. Four gill arches are present, with slit behind fourth arch, and gill membranes are separate and free of isthmus. Gill rakers on first arch are lathlike and number five on upper limb and eight on lower limb. Measurements are expressed as percent of SL: head length 23%–27%, snout length 8%– 11%, eye diameter 3%– 4%, upper jaw 10%– 12%, pectoral fin length 20%–22%, pelvic fin length 13%–14%, body depth 17%–18%. Pectoral fin is pointed and has 21 or 22 rays. First dorsal fin consists of seven to nine short spines, and second dorsal fin has one spine and 27 to 33 rays. Anal fin is similar in shape to second dorsal fin and has two short spines and 23 to 27 rays. Caudal fin is truncate in juve-

niles and deeply emarginate in adults, with upper lobe longer than lower. Scales are small and embedded in thick skin. Lateral line is wavy anteriorly and extends to caudal fin base. Gas bladder is absent. Vertebrae number 25: 12 precaudal and 13 caudal. Color is dark brown on back and side and yellowish ventrally, with two silvery bands on side. This species occurs worldwide, with exception of the eastern Pacific and Pacific Plate (Springer 1982), in tropical and warm temperate seas. In the western Atlantic it occurs from Massachusetts and Bermuda to Argentina, including the entire Gulf of Mexico. It is pelagic but is also associated with coral reefs, rocky substrates, and estuaries. Individuals are attracted to shadows produced by large, nearsurface objects such as buoys and flotsam. Food consists of crabs, squids, elasmobranchs, and ray-finned fishes. Maximum known age is eight years, and maximum known size is 200 cm TL.

references: Randall 1968a, 1996; Dawson 1971; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith 1986c; Shaffer and Nakamura 1989; Boschung 1992; Ditty and Shaw 1992; Smith 1995; Tominaga et al. 1996; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; SmithVaniz et al. 1999; Collette 2001c, 2002e.

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265

CORYPHAENIDAE

Dolphinfishes

Dolphinfishes are elongate, compressed, and subrectangular in profile, with dorsal fin originating on the head, extending to near caudal peduncle, and consisting solely of rays. Profile of head is strongly elevated and rounded, especially in adult males. Mouth is terminal and moderately oblique, with lower jaw projecting beyond upper jaw and upper jaw extending to pupil. Nostril is paired, and nares are close set, with a broad flap on posterior margin of anterior naris and on anterior margin of posterior naris. Eye is large, is located at mid-depth of head, and lacks adipose eyelid. Jaw teeth are small and arranged in narrow bands. Vomer and palatine have narrow bands of small teeth, and a patch of teeth occurs on tongue. Operculum lacks spines. Gill membranes are separate and free of isthmus. Gill rakers on first arch are short. Four gill arches are present, with slit behind fourth. Branchiostegal rays number 7. Pectoral fin is short and small and has an oblique base. Single dorsal fin is moderately higher anteriorly than posteriorly, extends from nape to caudal peduncle, and consists of 48 to 65 rays. Pelvic fin inserts below pectoral fin base, has one spine and 5 rays, and fits into groove on abdomen. Anal fin is elongate and consists of 23 to 30 rays. Caudal fin is deeply forked. Body is covered with small cycloid scales. Lateral line is complete and curved upward over pectoral fin. Gas bladder is absent, and vertebrae number 30 to 34. Upper section of body is brilliant metallic blue, with many dark spots in life, but fades to a uniform silver shortly after death. Juveniles less than 15 cm TL have vertical bars. Maximum known size is 160 cm TL. Dolphinfishes occur worldwide in tropical to warm temperate seas. They are epipelagic in oceanic waters and over or near continental and insular shelves, and are often associated with flotsam or other structures on or near the water surface. They feed on pelagic small crustaceans, squids, and ray-finned fishes. There are two species, and both occur in the Gulf of Mexico. references: Jordan and Evermann 1898; Gibbs and Collette 1959; Fritzsche and Fuiman 1982; Johnson 1984; Smith 1986b; Nelson 1994; Smith-Vaniz et al. 1999; Collette 2001d, 2002a.

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Key to the Species of the Gulf of Mexico (Adapted from Collette 1986a) 1a. Specimens up to 200 mm TL . . . . . . . . . . . . . . . . . . . . . . 2 1b. Specimens greater than 200 mm TL . . . . . . . . . . . . . . . . . 3 2a. Sides of body patterned with alternating light and dark bars that extend onto dorsal, anal, and caudal fins . . . . . . . . . . . . . . . . . . . . . . . Coryphaena hippurus p. 268 2b. Sides of body uniformly colored, dorsal and anal fins weakly barred. . . . . . . . . . . . . Coryphaena equiselis p. 267 3a. Body depth in adults less than 25% of SL; pectoral fin longer than half of head length; dorsal fin rays 58 to 66; lateral line scales 200 or more; tooth patch on tongue oval and covering middle third of tongue . . . . . . . . . . . . . . . . . . . . . . . Coryphaena hippurus p. 268 3b. Body depth in adults greater than 25% of SL; pectoral fin usually shorter than half of head length; dorsal fin rays 52 to 59; lateral line scales 200 or fewer; tooth patch on tongue trapezoidal in shape, wider anteriorly than posteriorly, and covering most of tongue . . . . . . . . . . . . . . . . . . . . . . . . Coryphaena equiselis p. 267

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Coryphaena equiselis Pompano dolphin

Linnaeus, 1758

Coryphaena equiselis is elongate, compressed, and relatively deep bodied, with relatively small pectoral fins and a convex anal fin. Head profile is slightly convex in young specimens, strongly convex in adults, and slightly more convex in males than in females. Tooth patch on tongue is trapezoid shaped, wider anteriorly than posteriorly, and covers most of tongue. Gill rakers on first arch number 1 or 2 on upper limb and 10 or 11 on lower limb in juveniles, and none on upper limb and 8 to 10 on lower limb in adults. Pectoral fin is acute, is about half of head length, and has 18 to 21 rays. Dorsal fin has 52 to 59 rays, and anal fin has 23 to 29 rays. Lateral line scales number 160 to 200. Vertebrae number 33 or (rarely) 34: 13 or 14 precaudal and 19 or 20 caudal. Color is brilliant metallic bluish green dorsally and silvery with golden sheen and small

black spots laterally and ventrally, with indistinct darker bars that do not extend onto fins. Shortly after death, color changes to gray with a green tinge. Juveniles are uniformly colored, and dorsal and anal fins are weakly barred. This species occurs worldwide in tropical and warm temperate seas in both oceanic and coastal waters. It is widespread in the Gulf of Mexico. Food consists of crustaceans, squids, and small ray-finned fishes. Maximum known size is 75 cm TL.

references: Gibbs and Collette 1959; Böhlke and Chaplin 1968; Collette et al. 1969; Hoese and Moore 1977, 1998; Palko et al. 1982; Collette 1986a, 2001d, 2002a,b; Robins and Ray 1986; Smith 1986b; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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Coryphaena hippurus Linnaeus, 1758 Dolphin

Coryphaena hippurus is elongate, compressed, and relatively slender, with relatively large pectoral fins and a concave anal fin. Body is relatively slender, and head profile of juvenile specimens is slightly convex. Head profile becomes increasingly convex with growth and in large males is nearly vertical. Tooth patch on tongue is small and oval shaped and covers about onethird of tongue surface. Gill rakers on first arch number 1 or 2 on upper limb and 6 to 11 on lower limb in juveniles, and none on upper limb and 8 or 9 on lower limb in adults. Pectoral fin is acute, is longer than half of head length, and has 17 to 20 rays. Dorsal fin has 58 to 66 rays, and anal fin has 25 to 30 rays. Lateral line scales number 200 to 280. Vertebrae number 30 or (rarely) 31: 13 or 14 precaudal and 17 or 18 caudal. Color is bright bluish dorsally,

golden yellow with dark flecks laterally, and whitish ventrally. Fins are bluish or greenish. Juvenile specimens are patterned with broad light and dark bars that extend onto dorsal and anal fins. Colors fade quickly after death. This species occurs worldwide in tropical and warm temperate seas in both oceanic and coastal waters. It is widespread in the Gulf of Mexico. Food consists of crustaceans, squids, and ray-finned fishes. Maximum known size is 200 cm TL.

references: Gibbs and Collette 1959; Böhlke and Chaplin 1968; Hoese and Moore 1977, 1998; Palko et al. 1982; Collette 1986a, 2001d, 2002a,b; Robins and Ray 1986; Smith 1986b; Boschung 1992; Schaldach et al. 1997; Smith 1997; Oxenford 1999; Smith-Vaniz et al. 1999.

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269

CARANGIDAE

Jacks

Jacks are deep bodied and compressed to elongate or fusiform, usually with two dorsal fins and a slender caudal peduncle. Head is short, deep, and compressed to long and rounded. Nostril is paired. Mouth is terminal and of moderate size. Jaw teeth are small and arranged in one or more bands, caninelike and in a single row, or, rarely, absent. Teeth are present or absent in vomer, palatine, and tongue. Operculum has spines in larvae and small juveniles but lacks them in large juveniles and adults. Gill rakers vary in size and number, and number decreases with growth in some species. Gill membranes are not connected and are free of isthmus. Branchiostegal rays number 6 to 10. Pectoral fin is short and pointed or long and falcate and has 14 to 24 rays. First dorsal fin has four to eight spines, and second has one spine and 17 to 44 rays. In some species, first dorsal fin is replaced by four to eight short, free spines that may be embedded. Pelvic fin inserts below pectoral fin insertion and has one spine and 5 rays. Two species lack pelvic fins. Anal fin has two (rarely one) free spines preceding membranous section of fin, and membranous section consists of one spine and 15 to 28 rays. Anal spines may be embedded in large specimens of some species. Anterior sections of rayed dorsal fin and anal fin are greatly elongated in some species. Caudal fin is high and deeply forked. Caudal peduncle varies from moderately deep to slender and often has one or two lateral keels. Body is at least partly covered with small cycloid (rarely ctenoid or needlelike) scales. Lateral line is complete, broadly curved over pectoral fin, and extends onto caudal fin. Scales of straight part of lateral line are often modified into enlarged, thickened spiny scutes. Gas bladder is present in most species, and when present, extends into caudal myomeres. Vertebrae number 24 to 26: 10 or 11 precaudal and 14 to 16 caudal. Most species are countercolored: dark (green or blue to blackish) dorsally and pale (silvery to white or yellowish) ventrally. Some species are uniformly silvery or uniformly dark in life. Juveniles of some species are barred or spotted. Maximum known size is 150 mm TL. Jacks occur worldwide in tropical to warm temperate seas. They are pelagic over continental and insular shelves or are oceanic. Some, especially juveniles, enter estuaries. Most are schooling fishes that pursue largely pelagic prey such as shrimps, squids, and ray-finned fishes. Spawning occurs offshore, and eggs and larvae are pelagic. Body shape and coloration change considerably with growth. Juveniles generally are deeper bodied, have shorter fins, and are more brightly colored than adults. There are 140 species in 32 genera worldwide, 32 species in 15 genera in the western Central Atlantic, and 28 or 29 species in 14 or 15 genera in the Gulf of Mexico.

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references: Fritzsche and Fuiman 1982; Johnson 1984; Smith-Vaniz 1984, 1986a, 2001b, 2002c; Nelson 1994; Tominaga et al. 1996; Smith-Vaniz et al. 1999.

Key to the Species of the Gulf of Mexico (Adapted from Berry 1959; Smith-Vaniz et al. 1999) 1a. Posterior straight part of lateral line with enlarged, thickened spiny scutes; pectoral fins longer than head and falcate (equal to head in Selar and Trachurus and shorter than head in Decapterus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Posterior straight part of lateral line without scutes; pectoral fins shorter than head . . . . . . . . . . . . . . . . . . . . . . 20 2a. Maximum body depth 20% to 28% of fork length (FL); pectoral fins equal to or shorter than head length. . . . . . . 3 2b. Maximum body depth 28% to 67% of fork length (FL); pectoral fins longer than head length . . . . . . . . . . . . . . . . 7 3a. Single detached finlet behind dorsal and anal fins. . . . . . . 4 3b. No finlet behind dorsal and anal fins (posterior ray of dorsal fin may be slightly separated in Trachurus) . . . . . . . . . 6 4a. Curved section of lateral line with 10 to 14 dark spots; curved section of lateral line with 37 to 56 scales and one to five scutes; total lateral line scales anterior to caudal peduncle 72 to 98 . . . . . . . . . Decapterus punctatus p. 283 4b. Curved section of lateral line with no dark spots; curved section of lateral line with 61 or more scales but no scutes; total lateral line scales anterior to caudal peduncle 103 or more. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5a. Curved section of lateral line moderately arched; anterior scales in straight section of lateral line 0 to 8; straight section of lateral line with 34 to 44 scutes . . . . . . . . . . . . . . . . . . . . . . . . . . . . Decapterus tabl p. 284 5b. Curved section of lateral line weakly arched; anterior scales in straight section of lateral line 19 to 33; straight section of lateral line with 23 to 32 scutes . . . . . . . . . . . . . . . . . . . . . . Decapterus macarellus p. 282 6a. Scales in curved section of lateral line expanded dorsoventrally (scutelike); cleithral furrow along posteroventral margin of gill cavity absent . . . . . Trachurus lathami p. 302 6b. Scales in curved section of lateral line not expanded dorsoventrally; cleithral furrow along posteroventral margin of gill cavity present . . . . . . . Selar crumenophthalmus p. 290 7a. Body appears naked, scales are minute and embedded when present (except some scales on posterior section of lateral line modified into small- to moderate-sized scutes) . . . . . . 8 7b. Body with small scales over most of body; posterior section of lateral line with or without scutes . . . . . . . . . . . . . . . 11 8a. Pelvic fins longer than upper jaw in large juveniles and adults; dorsal profile of head in front of eye broadly

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rounded; anterior dorsal fin spines not elongated in small specimens . . . . . . . . . . . . . . . . . . . . . Alectis ciliaris p. 274 8b. Pelvic fins about one-fourth or one-third the length of upper jaw in large juveniles and adults; dorsal profile of head in front of eye nearly straight to slightly concave; anterior dorsal fin spines distinctly elongated . . . . . . . . . . 9 9a. Anterior profile of head nearly vertical; anterior lobes of second dorsal fin and anal fin not greatly elongated . . . . 10 9b. Anterior profile of head oblique; anterior lobes of second dorsal fin and anal fin greatly elongated into filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Selene vomer p. 293 10a. Gill rakers on upper limb and lower limb of first gill arch 6 to 8 and 24 to 28, respectively; body depth 53% to 63% of FL. . . . . . . . . . . . . . . . . . . . . . . . . Selene brownii p. 291 10b. Gill rakers on upper limb and lower limb of first gill arch 7 to 10 and 27 to 35, respectively; body depth 43% to 56% of FL . . . . . . . . . . . . . . . . . . . . . . . Selene setapinnis p. 292 11a. Longest fin spines of first dorsal fin longer than lobe of second dorsal fin. . . . . . . . . . . . . Pseudocaranx dentex p. 289 11b. Longest fin spines of first dorsal fin shorter than lobe of second dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12a. Upper caudal fin lobe about 25% to 35% longer than lower caudal fin lobe; upper jaw 33% to 35% of head length; body strongly compressed . . . . . . . . . . . . . . . . . . . . . . . 13 12a. Upper caudal fin lobe about equal to lower caudal fin lobe; upper jaw more than 34% of head length; body not strongly compressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13a. Scutes in straight part of lateral line well developed and number 38 to 56 . . . . . Hemicaranx amblyrhynchus p. 286 13b. Scutes in straight part of lateral line poorly developed and number 6 to 12 . . . . . . . Chloroscombrus chrysurus p. 281 14a. Tongue, roof, and floor of mouth white, remainder of mouth dark; anal fin spines reduced or reabsorbed . . . . . . . . . . . . . . . . . . . . . . . . . . . Uraspis secunda p. 303 14b. Lining of mouth not distinctly light and dark; anal fin spines present and movable . . . . . . . . . . . . . . . . . . . . . . 15 15a. Lower limb of first gill arch with 31 to 35 gill rakers; adults with dark stripe extending from back to lower caudal fin lobe . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx ruber p. 280 15b. Lower limb of first gill arch with fewer than 29 gill rakers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16a. Lower limb of first gill arch with 25 to 28 gill rakers; straight section of lateral line with 46 to 56 scutes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx crysos p. 276 16b. Lower limb of first gill arch with 16 to 21 gill rakers; straight section of lateral line with 22 to 39 scutes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 17a. Dorsal fin rays 25 to 27; anal fin rays 21 to 24 . . . . . . . . . . . . . . . . . . . . . . . Caranx bartholomaei p. 275

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17b. Dorsal fin rays 19 to 23; anal fin rays 16 to 19 . . . . . . . 18 18a. Head and body dark brown to sooty black; upper jaw in adults extends to about middle of eye; lobe of second dorsal fin longer than head length . . . . Caranx lugubris p. 279 18b. Head and body green, yellow, and silver in adults; dark bars on body of juveniles; upper jaw in adults extends to posterior margin of eye; lobe of second dorsal fin shorter than head length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 19a. Chest completely scaled in specimens longer than 20 to 25 mm SL; first dorsal fin with pigment concentrated between first and fourth spines in specimens 17 to 45 mm SL; pigment of last three bars on body terminates immediately below lateral line on specimens between 20 and 60 mm SL . . . . . . . . . . . . . . . . . . . . . . Caranx latus p. 278 19b. Chest with only small patch of scales before pelvic fin in specimens longer than 25 mm SL; first dorsal fin with pigment evenly distributed over entire fin in specimens 17 to 45 mm SL; pigment of last three bars on body extends to anal fin base on specimens between 20 and 60 mm SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx hippos p. 277 20a. Caudal peduncle with dorsal and ventral grooves; base of rayed section of anal fin about 45% to 70% of base of second dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 20b. Caudal peduncle without grooves; base of rayed section of anal fin equal to or only slightly shorter than base of second dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 21a. Two rayed finlets behind dorsal and anal fins; upper jaw extends to anterior margin of eye (juveniles) or fails to reach anterior margin of eye (adults) . . . . . . . . . . . . . . . . . . . . . . . . Elagatis bipinnulata p. 285 21b. No finlets behind dorsal and anal fins; upper jaw extends at least to anterior margin of eye . . . . . . . . . . . . . . . . . . 22 22a. Upper jaw extending to anterior margin of eye; caudal fin lobes with distinct white tips; caudal peduncle of adults with well-developed medial cutaneous keel . . . . . . . . . . . . . . . . . . . . . . . . . . . Naucrates ductor p. 287 22b. Upper jaw extending from anterior margin to middle of eye; caudal fin lobes without distinct white tips; caudal peduncle of adults with slightly developed or no medial cutaneous keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23a. Total gill rakers on first gill arch fewer than 20 in specimens greater than 200 mm TL; fin rays in second dorsal fin 29 to 40 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 23b. Total gill rakers on first gill arch more than 20; fin rays in second dorsal fin 27 to 33 . . . . . . . . . . . . . . . . . . . . . . . 25 24a. Second dorsal fin with 29 to 35 rays; anal fin base 59% to 71% of second dorsal fin base; juvenile specimens have five dark bars on body that become irregularly split vertically

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CARANGIDAE 273

and do not extend to fin membranes, and a sixth bar on caudal peduncle . . . . . . . . . . . . . . . Seriola dumerili p. 295 24b. Second dorsal fin with 33 to 40 rays; anal fin base 48% to 63% of second dorsal fin base; juvenile specimens have six solid dark bars on body, with third, fourth, and fifth extending to fin membranes . . . . . . . Seriola zonata p. 298 25a. First dorsal fin with eight spines (first or last missing in large specimens); anterior lobe of second dorsal fin 12% to 12.5% of FL; anal fin base 53% to 59% of second dorsal fin base; juveniles with nuchal stripe extending from eye to nape, not extending to dorsal fin base, and seven irregular and broken bars on body, with third through seventh extending onto fin membranes . . . . . Seriola fasciata p. 296 25b. Dorsal fin with seven spines (first missing in large specimens); anterior lobe of second dorsal fin 19% to 22% of FL; anal fin base 63% to 70% of second dorsal fin base; juveniles with nuchal stripe extending through eye to dorsal fin base, and seven dark bars on body bisected by light pigment and not extending onto fin membranes . . . . . . . . . . . . . . . . . . . . . . . . . . . Seriola rivoliana p. 297 26a. Snout pointed; posterior sections of dorsal and anal fins consist of semidetached finlets; scales needlelike and partially embedded . . . . . . . . . . . . . . Oligoplites saurus p. 288 26b. Snout rounded, lower jaw included; posterior sections of dorsal and anal fins not consisting of semidetached finlets; scales cycloid and oval shaped . . . . . . . . . . . . . . . . . . . . 27 27a. Dorsal fin rays 17 to 21; anal rays 16 to 19 . . . . . . . . . . 28 27b. Dorsal fin rays 22 to 27; anal fin rays 20 to 24 . . . . . . . . . . . . . . . . . . . . . . . Trachinotus carolinus p. 299 28a. Anterior dorsal and anal fin rays greatly prolonged, reaching beyond caudal peduncle in adults; series of narrow dark bars on upper sides; teeth absent on tongue . . . . . . . . . . . . . . . . . . . . . . . . . Trachinotus goodei p. 301 28b. Anterior dorsal and anal fin rays not greatly prolonged, reaching base of caudal fin in adults; upper sides without narrow dark bars; teeth present on tongue . . . . . . . . . . . . . . . . . . . . . . . . Trachinotus falcatus p. 300

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Alectis ciliaris (Bloch, 1787) African pompano

Alectis ciliaris is moderately deep bodied as an adult and very deep bodied as a juvenile and compressed, with bony scutes on straight section of lateral line and posterior section of body triangular shaped. Dorsal profile of head is strongly arched. Snout is blunt. Eye possesses adipose eyelid. Maxilla extends to or beyond middle of eye. Jaw teeth are small, clawlike, depressible, and arranged in narrow bands, and become obsolete with age. Preopercular margin is membranous. Gill rakers on first arch are lathlike and number 4 to 6 on upper limb and 12 to 17 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and has 20 rays. First dorsal fin consists of seven free spines in juveniles but is absent in adults. Second dorsal fin has one spine and 18 or 19 rays, with anterior rays lobelike in adults. Pelvic fin is very elongate in juveniles. Anal fin consists of two free spines followed by one spine and 15 to 17 rays, with anterior rays elongated and forming lobe. Free anal spines are not visible in adults. In juveniles, anterior lobed section of dorsal and anal fins is greatly extended and filamentous. Scales are very small and cycloid, with

parts of head and body naked. Lateral line is strongly arched anteriorly and possesses 12 to 30 scutes on posterior straight section. Caudal peduncle has two keels. Color is silvery, with light metallic bluish tinge on upper section of head and body. Juveniles have three chevronshaped dark patterns on body. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs from Massachusetts and Bermuda to Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. Juveniles are pelagic drifters, and adults occur near the bottom to a depth of 60 m. It is a solitary species. Food consists of squids, small crabs, and ray-finned fishes. Maximum known size is 130 cm to 150 cm FL.

references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Matsuura 1983e; Robins and Ray 1986; Smith-Vaniz 1986a, 2001b; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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CARANGIDAE 275

Caranx bartholomaei Yellow jack

Cuvier, 1833

Caranx bartholomaei is elongate and moderately compressed, with a relatively long snout, a relatively long second dorsal fin, and bony scutes on straight portion of lateral line. Dorsal profile of head is evenly convex. Eye possesses adipose eyelid. Maxilla does not extend to anterior margin of eye. Jaw teeth are arranged in narrow bands, with those in outer row enlarged and recurved, and those in upper jaw larger than those of lower jaw. Preopercular margin is smooth in specimens greater than 40 mm SL. Gill rakers on first arch are lathlike and number 6 to 9 on upper limb and 18 to 21 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and has 20 or 21 rays. First dorsal fin has eight spines, and second dorsal fin has one spine and 25 to 28 rays, with anterior rays slightly longer than remainder of rays. Anal fin consists of two free spines followed by one spine and 21 to 24 rays, with anterior rays slightly longer than remainder of rays. Chest is scaled. Lateral line is moderately arched anteriorly and possesses 91 to 114 scales, including 22 to 28 scutes on poste-

rior straight section. Caudal peduncle has two keels. Vertebrae number 24: 10 precaudal and 14 caudal. Color is pale greenish blue dorsally and silvery with a yellowish cast ventrally. Juveniles have five vertical bars on body. This species occurs in the western Atlantic from Massachusetts and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with reefs along the outer continental and insular shelves. Juveniles are pelagic and associated with jellyfishes and rafts of Sargassum weed, and also occur in mangrove-lined lagoons. It is usually a solitary species. Food consists mostly of benthic ray-finned fishes. Maximum known size is about 90 cm FL.

references: Berry 1959; Böhlke and Chaplin 1968; Randall 1968a, 1996 (as Carangoides bartholomaei); Hoese and Moore 1977, 1998; CastroAguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Boschung 1992; Cervigón 1993a; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Smith-Vaniz 2002c.

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276 PERCIFORMES

Caranx crysos Blue runner

(Mitchill, 1815)

Caranx crysos is elongate and moderately compressed, with a relatively short second dorsal fin and bony scutes on straight portion of lateral line. Dorsal profile of head is evenly convex, and snout is moderately acute. Eye possesses adipose eyelid. Maxilla extends to about middle of eye. Jaw teeth are arranged in narrow bands, with those in outer row enlarged and recurved. Preopercular margin is smooth in specimens greater than 40 mm SL. Gill rakers on first arch are lathlike and number 10 to 14 on upper limb and 23 to 28 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and has 21 to 23 rays. First dorsal fin has eight spines, and second dorsal fin has one spine and 22 to 25 rays, with anterior rays longer than remainder and forming acutely tipped lobe. Anal fin consists of two free spines followed by one spine and 19 to 21 rays, with anterior rays slightly longer than remainder of rays and forming acutely tipped lobe. Chest is scaled. Lateral line is moderately arched anteriorly. Lateral line scales number 86 to 98, with last 46 to 56 forming scutes. Caudal pe-

duncle has two keels. Vertebrae number 25: 10 precaudal and 15 caudal. Color is dark olive to bluish dorsally and silvery to brassy ventrally. Juveniles have seven vertical bars on body. This species occurs in the western Atlantic from Nova Scotia and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is a schooling species along the outer continental and insular shelves and is not associated with reefs. Juveniles are pelagic and associated with rafts of Sargassum weed. Food consists of shrimps, crabs, other invertebrates, and rayfinned fishes. Maximum known size is about 630 mm FL.

references: Berry 1959; Böhlke and Chaplin 1968; Randall 1968a (as C. fusus), 1996 (as Carangoides crysos); Hoese and Moore 1977, 1998; Matsuura 1983e; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; SmithVaniz 2002c.

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CARANGIDAE 277

Caranx hippos Crevalle jack

(Linnaeus, 1766)

Caranx hippos is elongate and moderately compressed, with a relatively short second dorsal fin, a largely scaleless chest, and bony scutes on straight portion of lateral line. Dorsal profile of head is broadly convex, and snout is bluntly pointed. Eye possesses adipose eyelid. Maxilla extends to or behind posterior margin of eye. Jaw teeth are arranged in narrow bands, with those in outer row enlarged and recurved. Preopercular margin is smooth in specimens greater than 40 mm SL. Gill rakers on first arch are lathlike and number 6 to 9 on upper limb and 16 to 19 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and has 20 to 21 rays. First dorsal fin consists of eight spines, and second dorsal fin has one spine and 19 to 21 rays, with anterior rays longer than remainder of rays and forming acutely tipped lobe. Anal fin consists of two free spines followed by one spine and 16 or 17 rays, with anterior rays slightly longer than remainder of rays and forming acutely tipped lobe. Chest is naked except for median patch of scales anterior to pelvic fin. Lateral line is moderately arched anteriorly and possesses 23 to 35 scutes on posterior straight section. Caudal peduncle has two keels. Vertebrae number 24: 10 pre-

caudal and 14 caudal. Color is greenish, bluish, or bluish black dorsally and silvery white or golden ventrally. Elongate black spot is located on margin of operculum, and broad black spot is located on lower rays of pectoral fin. Juveniles have five vertical bars on body. This species occurs in the tropical to warm temperate Atlantic. In the western Atlantic it occurs from Nova Scotia to Uruguay, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater Antilles. It is a schooling species along continental and insular shelves and occurs in brackish water. Large individuals occur in deep offshore waters. Juveniles are pelagic and associated with rafts of Sargassum weed. Food consists of shrimps, other invertebrates, and ray-finned fishes. Maximum known size is at least 101 cm TL.

references: Berry 1959; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977 (as C. fuscus), 1998; Castro-Aguirre and Márquez-Espinoza 1981; Matsuura 1983e; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz 2002c.

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278 PERCIFORMES

Caranx latus Agassiz, 1831 Horse-eye jack

Caranx latus is elongate and moderately compressed, with a relatively short second dorsal fin, a completely scaled chest, and bony scutes on straight portion of lateral line. Dorsal profile of head is evenly convex, and snout is bluntly pointed. Eye possesses adipose eyelid. Maxilla extends to or behind posterior margin of eye. Jaw teeth are arranged in narrow bands, with those in outer row enlarged and recurved. Preopercular margin is smooth in specimens greater than 40 mm SL. Gill rakers on first arch are lathlike and number 6 or 7 on upper limb and 16 to 18 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and has 19 to 21 rays. First dorsal fin consists of eight spines, and second dorsal fin has one spine and 19 to 22 rays, with anterior rays longer than remainder of rays and forming acutely tipped lobe. Anal fin consists of two free spines followed by one spine and 16 to 18 rays, with anterior rays slightly longer than remainder of rays and forming acutely tipped lobe. Chest is fully scaled. Lateral line is moderately arched anteriorly and consists of 84 to 92 scales, the last 32 to 39 of which are scutes. Caudal pe-

duncle has two keels. Vertebrae number 24: 10 precaudal and 14 caudal. Color is dark blue to bluish green dorsally and silvery white to golden ventrally. Dorsal fin lobe and occasionally posterior scutes are dark to black. Juveniles have five dark bars on body. This species occurs in the tropical and warm temperate Atlantic. In the western Atlantic it occurs from New Jersey and Bermuda to Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is a schooling species along the continental and insular shelves and occurs in brackish and freshwater with connections to the sea. Food consists of shrimps; other invertebrates, including pteropods; and especially ray-finned fishes. Maximum known size is about 800 mm TL.

references: Longley and Hildebrand 1941; Berry 1959; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Smith-Vaniz 2002c.

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CARANGIDAE 279

Caranx lugubris Black jack

Poey, 1860

Caranx lugubris is oblong and moderately compressed, with a relatively high second dorsal fin and bony scutes on straight portion of lateral line. Dorsal profile of head is straight anteriorly and strongly convex at level of eye, and snout is bluntly pointed. Eye possesses prominent adipose eyelid. Maxilla extends to about middle of eye. Jaw teeth are arranged in narrow bands, with those in outer row of upper jaw enlarged and recurved. Outer teeth of lower jaw are enlarged but smaller than those of upper jaw. Preopercular margin is smooth in specimens greater than 40 mm SL. Gill rakers on first arch are lathlike and number 6 to 8 on upper limb and 18 to 21 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and has 20 to 22 rays. First dorsal fin consists of eight spines, and second dorsal fin has one spine and 20 to 23 rays, with anterior rays greatly elongate and forming long, acutely tipped lobe. Anal fin consists of two free spines followed by one spine and 17 to 20 rays, with anterior rays greatly elongated and forming

long, acutely tipped lobe. Chest is fully scaled. Lateral line is strongly arched anteriorly and possesses 26 to 33 scutes on posterior straight section. Caudal peduncle has two keels. Vertebrae number 24: 10 precaudal and 14 caudal. Color is gray to brown, with fins and posterior scutes dark brown to black. Color of juveniles is unknown. This species is circumtropical. In the western Atlantic it occurs from southern Florida and Bermuda to Brazil, including the Bahamas, the Greater and Lesser Antilles, and offshore waters of the Gulf of Mexico. It is uncommon at depths less than 24 to 65 m and is associated with clear oceanic water. Food consists mostly of ray-finned fishes. Maximum known size is about 991 mm FL. references: Longley and Hildebrand 1941; Berry 1959; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith-Vaniz 1986a, 2001b; Cervigón 1993b; Smith 1997; Smith-Vaniz et al. 1999.

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280 PERCIFORMES

Caranx ruber Bar jack

(Bloch, 1793)

Caranx ruber is elongate and moderately compressed, with a relatively long second dorsal fin, and bony scutes on straight portion of lateral line. Dorsal profile of head is evenly convex, and snout is moderately pointed. Eye possesses adipose eyelid. Maxilla extends nearly to anterior margin of eye. Jaw teeth in upper jaw consist of an outer row of enlarged, recurved teeth and an inner band of villiform teeth. Preopercular margin is smooth in specimens greater than 44 mm SL. Gill rakers on first arch are lathlike and number 10 to 14 on upper limb and 31 to 38 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and has 19 to 21 rays. First dorsal fin consists of eight spines, and second dorsal fin has one spine and 26 to 30 rays, with anterior rays longer than remainder of rays and forming acutely tipped lobe. Anal fin consists of two free spines followed by one spine and 23 to 26 rays, with anterior rays slightly longer than remainder of rays and forming acutely tipped lobe. Chest is fully scaled. Lateral line is moderately arched anteriorly and possesses 97 to 104 scales, including 23 to 29 scutes on posterior straight section. Caudal peduncle has two

keels. Vertebrae number 24: 10 precaudal and 14 caudal. Color is silvery gray dorsally and silvery white ventrally, with dark bar extending from second dorsal fin through lower lobe of caudal fin. Juveniles have six vertical bars on body. This species occurs in the western Atlantic from New Jersey and Bermuda to Venezuela, including the entire Gulf of Mexico, the Bahamas, and the Antilles. In the northern Gulf of Mexico it is limited to offshore waters. It is associated with clear shallow water, usually near coral reefs, and often forms small to large schools. Juveniles are associated with rafts of Sargassum weed. Food consists of shrimps, other invertebrates, and especially ray-finned fishes. Maximum known size is about 500 mm TL.

references: Longley and Hildebrand 1941; Berry 1959; Böhlke and Chaplin 1968; Randall 1968a, 1996 (as Carangoides ruber); Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999 (as Carangoides ruber); Smith-Vaniz 2002c.

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CARANGIDAE 281

Chloroscombrus chrysurus Atlantic bumper

(Linnaeus, 1766)

Chloroscombrus chrysurus is ovate and compressed, with ventral profile more convex than dorsal profile, dorsal fins close together and about the same height, and upper caudal fin lobe longer than lower caudal fin lobe. Dorsal profile of head is slightly convex. Anterior naris is oval shaped, with a flap on posterior margin, and posterior naris is oval shaped and just behind anterior naris. Eye has poorly developed adipose eyelid. Maxilla extends nearly to anterior margin of eye. Jaw teeth are villiform, slightly recurved, and arranged in narrow bands. Preopercular margin is smooth. Gill rakers on first arch are lathlike and number 9 to 12 on upper limb and 30 to 37 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and has 18 to 20 rays. First dorsal fin consists of eight spines, and second dorsal fin consists of one spine and 25 or 28 rays, with anterior rays slightly elongated. Anal fin consists of two free spines followed by one spine and 25 to 28 rays, with anterior rays slightly elongated. Chest is fully scaled. Lateral

line is strongly arched anteriorly and possesses 6 to 12 weak scutes on posterior straight section. Caudal peduncle lacks keels. Vertebrae number 24: 10 precaudal and 14 caudal. Color is metallic blue dorsally and silvery white ventrally, with black saddle on caudal peduncle. This species occurs in the tropical and warm temperate Atlantic. In the western Atlantic it occurs in coastal waters from Massachusetts and Bermuda to Uruguay, including the northern and southern Gulf of Mexico and the Greater and Lesser Antilles. It is a schooling species and occurs in both coastal marine and brackish waters. Juveniles occur far offshore and are frequently associated with jellyfishes. Maximum known size is about 367 mm TL. references: Longley and Hildebrand 1941; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith-Vaniz 1986a, 2002c; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999.

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282 PERCIFORMES

Decapterus macarellus Mackerel scad

(Cuvier, 1833)

Decapterus macarellus is elongate and oval in cross section, with two distinctly separated dorsal fins, widely separated finlets behind second dorsal fin and anal fin, and scutes on posterior part of straight section of lateral line. Dorsal profile of head is very slightly convex, and snout is relatively long and acutely pointed. Eye has well-developed adipose eyelid. Maxilla extends beyond anterior margin of eye. Jaw teeth are minute and arranged in a single row in juveniles but are absent in adults. Preopercular margin is smooth. Gill rakers on first arch are lathlike and number 9 to 13 on upper limb and 31 to 39 on lower limb. Branchiostegal rays number 7. Anterior margin of shoulder girdle, forming posterior margin of gill opening, has two small papillae that project into gill opening. Pectoral fin is falcate and has 22 to 24 rays. First dorsal fin consists of eight spines, and second dorsal fin consists of one spine and 31 or 37 rays, with last ray separated from remainder of fin. Anal fin consists of two free spines followed by one spine and 27 to 31 rays, with last ray separated from remainder of fin. Chest is fully scaled. Lateral line is weakly arched anteriorly and possesses 68 to 79 scales

and 0 scutes on curved section and 19 to 33 scales and 23 to 32 scutes on straight section. Total lateral line scales and scutes range from 119 to 133. Caudal peduncle lacks keels. Vertebrae number 24: 10 precaudal and 14 caudal. Color is metallic blue to bluish black dorsally and silvery to white ventrally, with small black spot on upper margin of operculum. Caudal fin is yellowish green to yellow. This species occurs worldwide in open water in the tropical and warm temperate seas. In the western Atlantic it occurs from the Gulf of Maine and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it is apparently limited to offshore waters and is rare to absent in the northern Gulf. It is a schooling species and feeds on planktonic invertebrates. Maximum known size is about 300 mm FL. references: Berry 1968; Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986; Smith-Vaniz 1986a, 2001b, 2002c; Retzer 1990; Boschung 1992; Cervigón 1993b; SmithVaniz et al. 1999.

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CARANGIDAE 283

Decapterus punctatus Round scad

(Cuvier, 1829)

Decapterus punctatus is elongate and oval in cross section, with two distinctly separated dorsal fins, widely separated finlets behind second dorsal fin and anal fin, and scutes on entire straight section of lateral line. Dorsal profile of head is very slightly convex, and snout is relatively long and acutely pointed. Eye possesses well-developed adipose eyelid. Maxilla extends beyond anterior margin of eye. Jaw teeth are minute and arranged in a single row. Preopercular margin is smooth. Gill rakers on first arch are lathlike and number 11 to 16 on upper limb and 32 to 40 on lower limb. Branchiostegal rays number 7. Anterior margin of shoulder girdle, forming posterior margin of gill opening, has two small papillae that project into gill opening. Pectoral fin is falcate and has 19 to 21 rays. First dorsal fin consists of eight spines, and second dorsal fin has one spine and 29 to 34 rays, with last ray separated from remainder of fin. Anal fin consists of two free spines followed by one spine and 25 to 30 rays, with last ray separated from remainder of fin. Chest is fully scaled. Lateral line is moderately arched anteriorly and possesses 37 to 56 scales, 0 to 6 scutes on curved section, and 32 to 46 scutes on straight section. Total lateral line scales and scutes range from 77 to

98, excluding scutes on caudal peduncle. Caudal peduncle lacks keels. Vertebrae number 25: 10 precaudal and 15 caudal. Color is greenish to greenish blue dorsally and silvery to white with a dusky cast ventrally. Narrow bronze or olive stripe extends from tip of snout to caudal peduncle, small black spot is located on upper margin of operculum, and 1 to 14 dark spots mark pores of lateral line scales on curved section of lateral line. This species occurs in the tropical and warm temperate Atlantic. In the western Atlantic it occurs in coastal waters from Massachusetts and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is a schooling species closely associated with the bottom to about 90 m. It feeds on planktonic invertebrates, such as copepods, gastropod larvae, ostracods, and pteropods. Maximum known size is about 183 mm FL.

references: Longley and Hildebrand 1941; Berry 1968; Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986; Smith-Vaniz 1986a, 2002c; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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284 PERCIFORMES

Decapterus tabl Redtail scad

Berry, 1968

Decapterus tabl is elongate and oval in cross section, with two distinctly separated dorsal fins, widely separated finlets behind second dorsal fin and anal fin, and scutes on most of straight section of lateral line. Dorsal profile of head is very slightly convex, and snout is relatively long and acutely pointed. Eye has well-developed adipose eyelid. Mouth is terminal and slightly oblique, and maxilla extends beyond anterior margin of eye. Jaw teeth are minute and arranged in a single row, with those of upper jaw becoming obsolete with age. Preopercular margin is smooth. Posterodorsal margin of opercular membrane is finely serrated in adults. Gill rakers on first arch are lathlike and number 10 to 12 on upper limb and 30 to 33 on lower limb. Branchiostegal rays number 7. Anterior margin of shoulder girdle, forming posterior margin of gill opening, has two small papillae that project into gill opening. Pectoral fin is falcate and has 20 to 22 rays. First dorsal fin consists of eight spines, and second dorsal fin has one spine and 29 to 34 rays, with last ray separated from remainder of fin. Anal fin consists of two free spines followed by one spine and 24 to 27 rays, with last ray separated from remainder of fin. Chest is fully scaled. Lateral line is moderately arched anteriorly and possesses 62 to 78 scales and 0 scutes

on curved section, and 0 to 8 scales and 34 to 44 scutes on straight section. Total lateral line scales and scutes range from 103 to 118, excluding scutes on caudal peduncle. Caudal peduncle lacks keels. Vertebrae number 24: 10 precaudal and 14 caudal. Color is metallic blue to bluish black dorsally and silvery to white ventrally. Small black spot is located on upper margin of operculum, and spots are absent on pored scales of curved section of lateral line. Caudal fin is bright red, and tips of second dorsal fin rays are tinged with red. This species occurs in the tropical to warm temperate Atlantic, Indian, and western and Central Pacific Oceans. In the western Atlantic it occurs from North Carolina and Bermuda to Brazil, including the Gulf of Mexico off Louisiana and Tamaulipas. In the Gulf of Mexico it is associated with topographic highs such as salt domes (Bruce Thompson, pers. comm., July 18, 1998). It is a schooling species in midwater or near the bottom between 150 and 200 m. Food consists of planktonic invertebrates, primarily copepods. Maximum known size is about 480 mm FL. references: Berry 1968; Robins and Ray 1986; Smith-Vaniz 1986a, 2001b, 2002c; Retzer 1990; Cervigón 1993b; Smith-Vaniz et al. 1999.

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CARANGIDAE 285

Elagatis bipinnulata Rainbow runner

(Quoy and Gaimard, 1825)

Elagatis bipinnulata is fusiform, with two slightly separated dorsal fins, extremely long second dorsal and anal fins, and widely separated finlets behind second dorsal and anal fins. Dorsal profile of head is very slightly convex, and snout is long and acutely pointed. Eye possesses poorly developed adipose eyelid. Maxilla fails to reach anterior margin of eye. Jaw teeth are minute and arranged in bands. Minute teeth occur on roof of mouth and tongue. Preopercular margin is smooth. Gill rakers on first arch are lathlike and number 10 to 12 on upper limb and 25 to 28 on lower limb. Pectoral fin is falcate and has 20 rays. First dorsal fin consists of six spines, and second dorsal fin has one spine and 25 to 30 rays, with last two rays separated from remainder of fin. Anal fin consists of one free spine followed by one spine and 18 to 22 rays, with last two rays separated from remainder of fin. Scales are ctenoid and cover body and parts of operculum, cheek, pectoral fin, pelvic fin, and caudal fin. Lateral line is slightly arched anteriorly and possesses about 100 scales. Caudal peduncle has groove

dorsally and ventrally. Vertebrae number 24: 10 precaudal and 14 caudal. Color is dark olive blue or green dorsally and white ventrally. Two narrow light blue or bluish white stripes run from eye to caudal peduncle, separated by broad olive or yellowish stripe. Fins are dark, with olive or yellowish tint. This species is circumtropical. In the western Atlantic it occurs from Massachusetts and Bermuda to northeastern Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It schools near the surface, both over reefs and offshore. It feeds on planktonic invertebrates and ray-finned fishes. Maximum known size is 107 cm FL.

references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith-Vaniz 1986a, 2001b, 2002c; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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286 PERCIFORMES

Hemicaranx amblyrhynchus Bluntnose jack

(Cuvier, 1833)

Hemicaranx amblyrhynchus is moderately elongate and compressed, with bony scutes on straight portion of lateral line, first dorsal fin shorter than lobe of second dorsal fin, and a relatively short snout. Dorsal profile of head is evenly convex, and snout is blunt. Eye possesses weak adipose eyelid. Maxilla extends to anterior margin of eye. Jaw teeth are narrow and arranged in a single row and form a continuous band. Preopercular margin is smooth in specimens greater than 44 mm FL. Gill rakers on first arch are lathlike and number 7 to 10 on upper limb and 18 to 23 on lower limb. Branchiostegal rays number 6. Pectoral fin is moderately falcate and has 20 to 22 rays. First dorsal fin consists of seven spines, and second dorsal fin has one spine and 25 to 30 rays, with anterior rays slightly longer than remainder of rays and forming short, acutely tipped lobe. Anal fin consists of two free spines followed by one spine and 21 to 26 rays, with anterior rays slightly longer than remainder of rays and forming short, acutely tipped lobe. Up-

per lobe of caudal fin is distinctly longer than lower lobe. Chest is fully scaled. Lateral line is strongly arched anteriorly and possesses 38 to 56 scutes on posterior straight section. Caudal peduncle lacks keels. Vertebrae number 26: 10 precaudal and 16 caudal. Color is dark bluish green dorsally and silvery ventrally, with large black blotch on operculum and dark margin on dorsal fins and upper lobe of caudal fin. Juveniles have four or five dark bars on body. This species occurs in coastal waters in the western Atlantic from North Carolina to Brazil, including the northern and southern Gulf of Mexico, Cuba, and Trinidad. It is associated with the bottom, enters brackish water, and does not form large schools. Juveniles are associated with jellyfishes. Maximum known size is about 290 mm FL. references: Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Castro-Aguirre et al. 1999; Smith-Vaniz 2002c.

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CARANGIDAE 287

Naucrates ductor (Linnaeus, 1758) Pilotfish

Naucrates ductor is fusiform and slightly compressed, with first dorsal fin greatly reduced and fleshy keel on lateral aspect of caudal peduncle. Dorsal profile of head is moderately convex, and snout is moderately long and blunt. Maxilla extends to anterior margin of eye. Jaw teeth are minute and arranged in bands. Adipose eyelid is well developed. Preopercular margin is smooth. Gill rakers on first arch are lathlike and number 6 or 7 on upper limb and 15 to 20 on lower limb. Pectoral fin is falcate and has 19 rays. First dorsal fin consists of four or five very short, free spines, with first minute and last covered with flesh in specimens larger than 200 mm FL. Second dorsal fin has one spine and 25 to 29 rays. Anal fin consists of two free spines followed by one spine and 15 to 17 rays. Anterior free anal fin spine is covered with flesh in adult specimens. Scales are very small and ctenoid. Lateral line is moderately arched anteriorly. Caudal peduncle has a groove dorsally and ventrally. Vertebrae number 25: 10 precaudal and 15 cau-

dal. Color is silvery, with head dark and five to seven black bars on body that extend to dorsal and ventral midlines. Pectoral, pelvic, and caudal fins are blackish, and tips of caudal fin lobes are white. This species is circumtropical. In the western Atlantic it occurs from Nova Scotia and Bermuda to Argentina, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is pelagic in oceanic waters and is associated with large fishes such as sharks and rays, sea turtles, and driftwood. Juveniles are often associated with seaweeds and jellyfishes. It feeds on pieces of large prey captured by the host species, on ectoparasites on the host, and on small ray-finned fishes and invertebrates. Maximum known size is about 630 mm FL, and maturity is reached at about 230 mm FL. references: Robins and Ray 1986; Smith-Vaniz 1986a, 2001b, 2002c; Cervigón 1993b; Schaldach et al. 1997; Smith-Vaniz et al. 1999.

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288 PERCIFORMES

Oligoplites saurus Leatherjack

(Bloch and Schneider, 1801)

Oligoplites saurus is elongate and greatly compressed, with upper and lower profiles similar in shape, two slightly separated dorsal fins, and needlelike scales. Dorsal profile of head is slightly concave over eye, and snout is long and acutely pointed. Maxilla extends to posterior margin of eye. Jaw teeth are small and arranged in two rows. Teeth in outer row of upper jaw are irregular and smaller anteriorly than posteriorly, and teeth in lower jaw are conical. Juveniles have outwardly hooked spatulate teeth in outer row that are lost later in life. Preopercular margin is smooth. Gill rakers on first arch are lathlike and number 5 to 8 on upper limb and 13 to 15 on lower limb. Pectoral fin is relatively short and has 15 to 17 rays. First dorsal fin consists of four to six short spines. Second dorsal fin has one spine and 19 to 21 rays. Anal fin consists of two free spines followed by one spine and 19 to 22 rays. The last 11 to 15 rays of second dorsal fin and anal fin form semidetached finlets. Scales are needlelike (aricular) and partially embedded.

Lateral line is slightly arched over pectoral fin. Vertebrae number 26: 10 precaudal and 16 caudal. Color is bluish dorsally and silver to white laterally and ventrally. Dorsal fin spines are dusky, and membrane is transparent. This species occurs in the western Atlantic and eastern Pacific. In the western Atlantic it occurs from Massachusetts to southern Brazil or Uruguay, including the northern and southern Gulf of Mexico and the Greater and Lesser Antilles. It is associated with sandy beaches, bays, and inlets, often in turbid water. It also occurs in freshwater. Adults form large schools, and juveniles are often associated with the surface. Food consists of crustaceans and rayfinned fishes. Juveniles have incisor-like outer teeth and clean other fishes of ectoparasites. Maximum known size is 297 mm FL. references: Randall 1968a, 1996; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz 2002c.

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CARANGIDAE 289

Pseudocaranx dentex

(Bloch and Schneider, 1801)

Pseudocaranx dentex is moderately elongate and moderately compressed, with bony scutes on straight portion of lateral line, first dorsal fin higher than lobe of second dorsal fin, and a relatively long snout. Dorsal profile of head is evenly convex, and snout is long and moderately pointed. Eye has weak adipose eyelid. Maxilla fails to reach anterior margin of eye and has straight posterior margin. Jaw teeth are blunt and conical and arranged in a single row in both jaws. Juveniles have inner row of conical teeth in upper jaw. Gill rakers on first arch are lathlike and number 11 to 14 on upper limb and 23 to 28 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and longer than head. Dorsal fins are close together, and first consists of eight spines that are longer than lobe of second. Second dorsal fin has one spine and 25 to 27 rays. Anal fin consists of two free rays followed by one spine and 21 to 26 rays, with anterior rays slightly longer than remainder of rays and forming acutely tipped lobe. Terminal rays of second dorsal fin and anal fin are enlarged and slightly separated but not detached from penultimate rays. Chest is fully scaled or with naked spot anteroven-

trally. Scales are small and cycloid. Lateral line is moderately arched anteriorly and possesses 57 to 78 scales in curved section, and 2 to 27 scales and 16 to 30 scutes on posterior straight section. Caudal peduncle lacks keels. Vertebrae number 25: 10 precaudal and 15 caudal. Color is pale greenish to steely blue dorsally and silvery ventrally, with yellow stripe along side, base of second dorsal fin, and anal fin. Black spot is located on posterodorsal margin of operculum. Caudal and second dorsal fins are yellow. This species occurs in the subtropical Atlantic, Pacific, and Indian Oceans between 80 and 200 m. In the western Atlantic it occurs from North Carolina, Bermuda, and southern Brazil, with an unconfirmed account from Little Bahama Bank. There is an unconfirmed record of this species from the southern Gulf of Mexico off Tabasco, Mexico (UNAM 6042, 1856N, 9201W). This is a schooling species that feeds on the bottom. Maximum known size is 820 mm FL. references: Smith-Vaniz 1986a, 2001b, 2002c; Smith-Vaniz et al. 1999.

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290 PERCIFORMES

Selar crumenophthalmus Bigeye jack

(Bloch, 1793)

Selar crumenophthalmus is elongate and moderately compressed, with a deep furrow and two papillae on cleithrum of shoulder girdle and scutes along straight section of lateral line. Lower profile of body is slightly more convex than upper profile, and snout is relatively long and acutely pointed. Eye is very large and possesses well-developed adipose eyelid. Maxilla extends to anterior margin of pupil. Jaw teeth are small and recurved, with those in upper jaw arranged in a narrow band tapering posteriorly and those in lower jaw in a single row. Preopercular margin is smooth. Gill rakers on first arch are lathlike and number 9 to 12 on upper limb and 27 to 31 on lower limb. Branchiostegal rays number 7. Shoulder girdle has two broad-based papillae projecting into posterior wall of opercular cavity, with lower larger than upper, and a large cleithral furrow below lower papilla. Pectoral fin is falcate and has 20 or 22 rays. First dorsal fin consists of eight spines. Second dorsal fin has one spine and 24 to 27 rays, with anterior rays elongated. Anal fin consists of two free spines followed by one spine and 21 to 23 rays, with anterior rays elongated. Chest is fully scaled except for area anterior to pectoral fin. Lateral line is slightly arched anteriorly and possesses 48 to 56 pored scales and 0 to 4 scutes on curved section, and 0 to 11 pored scales and 29 to 44 scutes

on straight section. Total lateral line scales and scutes range from 84 to 94, excluding scutes on caudal peduncle. Caudal peduncle lacks keels. Accessory lateral line runs to origin of first dorsal fin. Vertebrae number 24: 10 precaudal and 14 caudal. Color is metallic blue to bluish green dorsally and silvery to white ventrally. Snout is dusky or blackish; a narrow yellowish stripe extends from operculum to upper part of caudal peduncle; and a small, elongated black spot is located on upper margin of operculum. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs in coastal waters to 170 m from Nova Scotia and Bermuda to Brazil, including the northern and southern Gulf of Mexico and the Greater and Lesser Antilles. It is a schooling species and feeds on planktonic and benthic invertebrates, such as shrimps and crabs, and small ray-finned fishes. Maximum known size is at least 270 mm FL.

references: Longley and Hildebrand 1941 (as Trachurops crumenophthalma); Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Castro-Aguirre and MárquezEspinoza 1981; Robins and Ray 1986; Smith-Vaniz 1986a, 2001b, 2002c; Boschung 1992; Cervigón 1993b; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999.

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CARANGIDAE 291

Selene brownii

(Cuvier, 1816)

Selene brownii is very deep bodied and extremely compressed, with ventral profile more convex than dorsal, pelvic fins very short, and scales very small and embedded. Dorsal profile of head is nearly vertical and concave to above eye and strongly rounded to nape. Snout is blunt. Eye possesses poorly developed adipose eyelid. Maxilla extends to below anterior margin of eye. Jaw teeth are relatively small and arranged in narrow, irregular bands. Preopercular margin is membranous. Gill rakers on first arch are lathlike and number 6 to 8 on upper limb and 24 to 28 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and has 17 rays. First dorsal fin has eight spines, with first four spines elongated and second equal to body depth in specimens shorter than 60 mm FL. Second dorsal fin has one spine and 21 to 23 rays, with anterior rays forming acute lobe. Anal fin consists of two free spines followed by one spine and 17 to 19 rays. Free anal

fin spines are absent in specimens greater than 130 mm FL. Pelvic fins are very short in juveniles and rudimentary in adults. Scales are very small and cycloid, cover most of lower half of body, but are mostly absent from area anterior to pelvic fin base and junction between curved and straight sections of lateral line. Straight section of lateral line has 7 to 12 weak scutes on caudal peduncle. Caudal peduncle has two keels. Vertebrae number 24: 10 precaudal and 14 caudal. Color is silvery, with faint dark spot on edge of operculum and on upper section of caudal peduncle. Juveniles have black spot on straight section of lateral line. This species occurs in the western Atlantic over the continental shelf from the southern Gulf of Mexico off Veracruz and the Greater and Lesser Antilles (Guadeloupe only) to Brazil. Maximum known size is 230 mm FL. references: Smith 1997; Smith-Vaniz 2002c.

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292 PERCIFORMES

Selene setapinnis Atlantic moonfish

(Mitchill, 1815)

Selene setapinnis is very deep bodied and extremely compressed, with ventral profile more convex than dorsal, pelvic fins very short, and scales very small and embedded. Dorsal profile of head is nearly vertical and concave to above eye and strongly rounded to nape. Snout is blunt. Eye possesses poorly developed adipose eyelid. Maxilla extends to below anterior margin of eye. Jaw teeth are relatively small and arranged in narrow, irregular bands. Preopercular margin is membranous. Gill rakers on first arch are lathlike and number 7 to 10 on upper limb and 27 to 35 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and has 17 or 18 rays. First dorsal fin has eight spines, with first four spines elongated and second equal to body depth in specimens shorter than 60 mm FL. Second dorsal fin has one spine and 21 to 24 rays, with anterior rays only slightly longer than posterior rays and not forming lobe. Anal fin consists of two free spines followed by one spine and 16 to 19 rays. Free anal spines are absent in specimens greater than 130 mm FL. Pelvic fins are very short in juveniles and rudimentary in adults. Scales are very small and cycloid, cover most of lower half of body, but are mostly absent from area

anterior to pelvic fin base and junction between curved and straight sections of lateral line. Straight section of lateral line has 7 to 17 weak scutes. Caudal peduncle has two keels. Vertebrae number 24: 10 precaudal and 14 caudal. Color is silvery, with faint dark spot on edge of operculum and on upper section of caudal peduncle. Juveniles have black spot on straight section of lateral line. This species occurs to about 54 m in the western Atlantic from Nova Scotia to Argentina, including the northern and southern Gulf of Mexico and the Antilles. It is apparently absent from the Bahamas. It is a schooling species associated with the bottom. Juveniles to 30 mm FL are pelagic drifters and found near the surface up to 180 km offshore. Food consists of crustaceans and small ray-finned fishes. Maximum known size is 332 mm FL, and sexual maturity is reached at about 130 mm FL.

references: Randall 1968a; Hoese and Moore 1977, 1998; Castro-Aguirre and MárquezEspinoza 1981; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; SmithVaniz 2002c.

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CARANGIDAE 293

Selene vomer Lookdown

(Linnaeus, 1758)

Selene vomer is very deep bodied and extremely compressed, with dorsal and ventral profiles parallel to abdominal margin, pelvic fins very short, and scales very small and embedded. Dorsal profile of head is very obtuse and concave to above eye and strongly rounded to nape. Snout is blunt. Eye possesses poorly developed adipose eyelid. Maxilla extends to below anterior margin of eye. Jaw teeth are relatively small, conical, and recurved and arranged in narrow bands. Preopercular margin is membranous. Gill rakers on first arch are lathlike and number 6 to 9 on upper limb and 23 to 27 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and has 20 to 22 rays. First dorsal fin has eight spines, with first four spines elongated and second spine equal to 2.5 times FL in specimens shorter than 35 mm FL. Second dorsal fin has one

spine and 20 to 23 rays, with anterior rays forming elongate filament. Anal fin consists of two free spines followed by one spine and 17 to 20 rays, with anterior rays elongated and forming acute lobe. Free anal fin rays are absent in specimens greater than 110 mm FL. Pelvic fins are very short in adults but greatly elongated in juveniles to 90 mm FL. Scales are very small and cycloid, cover most of body, but are mostly absent above lateral line anterior to second dorsal fin. Straight section of lateral line has 7 to 12 weakly developed scutes. Caudal peduncle lacks keels. Vertebrae number 24: 10 precaudal and 14 caudal. Color is silvery to golden, and dorsal surface has metallic bluish tinge. Prolonged dorsal and anal fin rays are black. Juveniles have dusky, slightly oblique crossband and band over eye. This species occurs in shallow coastal wa-

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294 PERCIFORMES

ters in the western Atlantic from Maine to Uruguay, including the northern and southern Gulf of Mexico and the Greater Antilles. It occurs in schools and feeds near the bottom on crustaceans and ray-finned fishes. Maximum known size is 400 mm FL.

references: Bigelow and Schroeder 1953; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Smith-Vaniz 2002c.

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CARANGIDAE 295

Seriola dumerili (Risso, 1810) Greater amberjack

Seriola dumerili is elongate and slightly compressed, with relatively short pectoral fins, no scutes on lateral line, and a dorsal and ventral caudal peduncle groove. Upper profile of body is slightly more convex than lower profile, and snout is moderately long and acutely pointed. Eye possesses poorly developed adipose eyelid. Maxilla extends to about middle of pupil. Supramaxilla is extremely broad. Jaw teeth are minute and arranged in broad bands. Gill rakers on first arch are lathlike and number 20 to 24 in specimens between 20 and 70 mm FL, and 11 to 19 in specimens greater than 200 mm FL. Branchiostegal rays number 7. Pectoral fin is acutely tipped and has 20 or 21 rays. First dorsal fin consists of six or seven spines, with last spine reduced and covered with skin in specimens greater than 600 mm FL. Second dorsal fin has one spine and 29 to 34 rays, with anterior rays forming short acute lobe, about 13% to 14% of FL. Anal fin consists of two free spines followed by one spine and 18 to 22 rays, with anterior rays forming acute lobe. Free anal fin rays are reduced and covered with flesh in specimens greater than 600 mm FL. Scales are small and cycloid. Lateral line is slightly arched anteriorly and consists of 141 to 163 scales. Caudal peduncle has upper and lower grooves. Vertebrae number 24: 10 precaudal and 14 caudal. Color is blue to oliva-

ceous dorsally and silvery to whitish on sides and ventrally. Dark nuchal stripe passes from eye to origin of first dorsal fin, and amber stripe runs from eye to along midflank of body. Caudal fin is dark or dusky with light and narrow posterior margin. Juveniles between 20 and 170 mm FL have five dark bars on side that become irregularly split vertically, and one bar on caudal peduncle. Bars on body do not extend to medial fin membranes. Dark nuchal stripe is well developed. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs over continental and insular shelves from Nova Scotia and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is generally a schooling species. Juveniles are associated with floating plants and debris over oceanic and neritic waters. Food consists of invertebrates and ray-finned fishes. Maximum known size is 188 cm FL.

references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Mather 1971; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith-Vaniz 1986a, 2001b, 2002c; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; SmithVaniz et al. 1999.

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296 PERCIFORMES

Seriola fasciata (Bloch, 1793) Lesser amberjack

Seriola fasciata is elongate and slightly compressed, with relatively short pectoral fins, no scutes on lateral line, and a dorsal and ventral caudal peduncle groove. Upper profile of body is slightly more convex than lower profile, and snout is moderately long and acutely pointed. Eye possesses poorly developed adipose eyelid. Maxilla extends to about anterior margin of pupil. Supramaxilla, on dorsal margin of maxilla, is narrow. Jaw teeth are minute and arranged in broad bands. Gill rakers on first arch are lathlike, number 6 to 8 on upper limb and 16 to 18 on lower limb, and do not decrease with growth. Branchiostegal rays number 7. Pectoral fin is acutely tipped and has 20 rays. First dorsal fin consists of eight spines, with first and last spines reduced and covered with flesh in large specimens. Second dorsal fin has one spine and 28 to 33 rays, with anterior rays forming short, acute lobe, about 12% to 12.5% of FL. Anal fin consists of two free spines followed by one spine and 17 to 20 rays, with anterior rays forming short, acute lobe. Free anal fin spines are reduced and covered with flesh in specimens greater than 600 mm FL. Scales are small and cycloid. Lateral line

is slightly arched anteriorly. Caudal peduncle has upper and lower grooves. Vertebrae number 24: 10 precaudal and 14 caudal. Color is dark pinkish to violet dorsally and silvery to whitish laterally and ventrally. Faint nuchal stripe passes from eye to nape but does not reach base of first dorsal fin. Juveniles between 40 and 260 mm FL have seven dark, irregular, and broken bars. Bars on body extend to medial fin membranes. Dark nuchal stripe is well developed. This species occurs in the tropical to warm temperate Atlantic Ocean. In the western Atlantic it occurs on or near the bottom between 55 and 150 m from Massachusetts and Bermuda to Venezuela, including the Gulf of Mexico and the Greater Antilles. It is generally a schooling species. Juveniles are epipelagic in oceanic and neritic waters. Food consists of squids and probably ray-finned fishes. Maximum known size is 675 mm FL. references: Mather 1971; Hoese and Moore 1977, 1998; Robins and Ray 1986; Cervigón 1993a,b; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Smith-Vaniz 2002c.

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CARANGIDAE 297

Seriola rivoliana Valenciennes, 1833 Almaco jack

Seriola rivoliana is elongate and slightly compressed, with relatively short pectoral fins, no scutes on lateral line, and a dorsal and ventral caudal peduncle groove. Upper profile of body is slightly more convex than lower profile, and snout is moderately long and acutely pointed. Eye possesses poorly developed adipose eyelid. Maxilla extends to about anterior margin of pupil. Supramaxilla, on dorsal margin of maxilla, is very broad. Jaw teeth are minute and arranged in broad bands. Gill rakers on first arch are lathlike and number 24 to 29 in specimens between 20 and 70 mm FL, and 22 to 26 in specimens greater than 200 m FL. Branchiostegal rays number 7. Pectoral fin is acutely tipped and has 21 rays. First dorsal fin consists of seven spines, with first spine reduced and covered with flesh in large specimens. Second dorsal fin has one spine and 27 to 33 rays, with anterior rays forming long, acute lobe, about 19% to 22% of FL. Anal fin consists of two free spines followed by one spine and 18 to 22 rays, with anterior rays forming long, acute lobe. Free anal fin spines are reduced and covered with flesh in specimens greater than 750 mm FL. Scales are small and cycloid. Lateral line is slightly arched anteriorly and

consists of 122 to 143 scales. Caudal peduncle has upper and lower grooves. Vertebrae number 24: 10 precaudal and 14 caudal. Color is brown, olivaceous, or bluish green dorsally and silvery to whitish laterally and ventrally. Faint nuchal stripe may be present from eye to nape. Caudal fin is dark with light, narrow posterior margin. Juveniles between 20 and 180 mm FL have seven dark bars subdivided by light irregular bars. Bars on body do not extend to medial fin membranes. Dark nuchal stripe is well developed. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs from Cape Cod and Bermuda to Argentina, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. Adults are pelagic in oceanic and outer neritic waters. Food consists of ray-finned fishes. Maximum known size is 123 cm FL. references: Böhlke and Chaplin 1968; Randall 1968a; Mather 1971; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith-Vaniz 1986a, 2001b, 2002c; Boschung 1992; Cervigón 1993a,b; Smith-Vaniz et al. 1999.

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298 PERCIFORMES

Seriola zonata (Mitchill, 1815) Banded rudderfish Seriola zonata is elongate and slightly compressed, with relatively short pectoral fins, no scutes on lateral line, and a dorsal and ventral caudal peduncle groove. Upper profile of body is slightly more convex than lower profile, and snout is moderately long and acutely pointed. Eye possesses poorly developed adipose eyelid. Maxilla extends to posterior margin of eye. Supramaxilla, on dorsal margin of maxilla, is broad. Jaw teeth are minute and arranged in broad bands. Gill rakers on first arch are lathlike and number 20 to 25 in specimens smaller than 100 mm FL, and 14 to 17 in specimens greater than 400 mm FL. Branchiostegal rays number 7. Pectoral fin is acutely tipped and has 18 rays. First dorsal fin consists of eight spines, with first spine reduced and often covered with flesh in specimens greater than 600 mm FL. Second dorsal fin has one spine and 33 to 39 rays, with anterior rays forming moderately long, slightly acute lobe, about 13% to 14% of FL. Anal fin consists of two free spines followed by one spine and 19 to 21 rays, with anterior rays forming moderately long, acute lobe. Free anal fin spines are reduced and located in groove in specimens greater than

400 mm FL. Scales are small and cycloid, and scutes are absent from lateral line. Lateral line is slightly arched anteriorly. Caudal peduncle has upper and lower grooves. Vertebrae number 24: 11 precaudal and 13 caudal. Color is bluish green dorsally and silvery to whitish on sides and ventrally. Dark nuchal stripe may be present from eye to first dorsal fin origin. Juveniles between 20 and 300 mm FL have six dark bars on body, with fourth, fifth, and sixth bars extending onto median fin membranes. Dark nuchal stripe extends from eye to first dorsal fin origin. This species occurs in coastal waters in the western Atlantic from Nova Scotia and Bermuda to Brazil, including the Gulf of Mexico and the Greater Antilles. Adults are pelagic. Juveniles are pelagic and associated with floating algae, jellyfishes, and large pelagic fishes. Food consists of shrimps and ray-finned fishes. Maximum known size is 800 mm FL. references: Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Smith-Vaniz et al. 1999; Smith-Vaniz 2002c.

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CARANGIDAE 299

Trachinotus carolinus (Linnaeus, 1766) Florida pompano

Trachinotus carolinus is deep bodied and compressed, with relatively short pectoral fins, no scutes on lateral line, and a relatively short anal fin base. Upper and lower profiles of body are about equally convex, and snout is very blunt. Eye possesses poorly developed adipose eyelid. Upper jaw projects slightly beyond lower jaw, and maxilla extends to below middle of eye. Jaw teeth are small, conical, and recurved in juveniles to 200 mm FL, but absent in larger specimens. Tongue lacks teeth. Gill rakers on first arch are lathlike and number 5 to 7 on upper limb and 8 to 14 on lower limb. Branchiostegal rays number 7. Specimens develop relatively shorter heads and deeper bodies with growth. Pectoral fin is acutely tipped and has 18 to 20 rays. First dorsal fin consists of six separate spines, with first spine reduced and often covered with flesh in specimens greater than 300 mm FL. Second dorsal fin has one spine and 22 to 27 rays, with anterior rays forming moderately long, acute lobe that fails to reach caudal fin base in adults. Anal fin consists of two free spines followed by one spine and 20 to 24 rays, with anterior rays forming relatively short, acute lobe. Base of anal fin is shorter

than base of second dorsal fin. Scales are small, cycloid, and partially embedded. Anterior section of lateral line is slightly wavy. Caudal peduncle lacks upper and lower grooves. Vertebrae number 24: 10 precaudal and 14 caudal. Color is dark silvery and metallic greenish to bluish dorsally and white laterally and ventrally. Tips of second dorsal fin and anal fin are dark. This species occurs between the shoreline and 40 m in the western Atlantic from Massachusetts to Brazil, including the northern and southern Gulf of Mexico. It is absent in the Antilles except for Jamaica, Puerto Rico, Tobago, and Trinidad. Adults are pelagic and are associated with sandy beaches around inlets and bays. Juveniles form large schools during the summer in the surf zone along beaches. Food consists of mollusks, crabs, shrimps, and small ray-finned fishes. Maximum known size is 350 mm FL. references: Fields 1962; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz 2002c.

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300 PERCIFORMES

Trachinotus falcatus Permit

(Linnaeus, 1758)

Trachinotus falcatus is deep bodied and compressed, with relatively short pectoral fins, no scutes on lateral line, and a relatively long anal fin base. Upper and lower profiles of body are about equally convex, and snout is very blunt. Eye possesses poorly developed adipose eyelid. Upper jaw projects slightly beyond lower jaw, and maxilla extends to below middle of eye. Jaw teeth are small, conical, and recurved in juveniles to 200 mm FL, but absent in larger specimens. Tongue has irregular patch of teeth. Gill rakers on first arch are lathlike and number 5 to 9 on upper limb and 11 to 15 on lower limb. Branchiostegal rays number 7. Specimens develop relatively shorter heads and deeper bodies with growth. Pectoral fin is acutely tipped and has 17 to 20 rays. First dorsal fin consists of five separate spines, with first spine reduced and often covered with flesh in specimens greater than 430 mm FL. Second dorsal fin has one spine and 17 to 21 rays, with anterior rays forming long, acute lobe extending to base of caudal fin in adults. Anal fin consists of two free spines followed by one spine and 16 to 19 rays, with anterior rays forming relatively long, acute lobe. Base of anal fin is about

equal to base of second dorsal fin. Scales are small, cycloid, and partially embedded. Lateral line is wavy anteriorly and consists of about 135 scales. Caudal peduncle lacks upper and lower grooves. Vertebrae number 24: 10 precaudal and 14 caudal. Color is bluish gray, with iridescent blue to bluish green sheen dorsally, and silvery laterally and ventrally. This species occurs in coastal waters to about 36 m in the western Atlantic from Massachusetts and Bermuda to southern Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. Adults are pelagic and are associated with channels and depressions over sandy and mud bottoms and reefs. Juveniles form large schools during the summer in the surf zone along beaches. Food consists of mollusks, crabs, shrimps, and small ray-finned fishes. Maximum known size is 106 cm FL. references: Fields 1962; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Smith-Vaniz 2002c.

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CARANGIDAE 301

Trachinotus goodei Palometa

Jordan and Evermann, 1896

Trachinotus goodei is deep bodied and compressed, with relatively short pectoral fins, no scutes on lateral line, and a relatively long anal fin base. Upper and lower profiles of body are about equally convex but slightly asymmetrical, and snout is very blunt. Eye possesses poorly developed adipose eyelid. Upper jaw projects slightly beyond lower jaw, and maxilla extends to below middle of eye. Jaw teeth are small, conical, and recurved, and decrease in number with growth but are always present. Tongue lacks teeth. Gill rakers on first arch are lathlike and number 4 to 9 on upper limb and 8 to 14 on lower limb. Branchiostegal rays number 7. Specimens develop relatively shorter heads and more slender bodies with growth. Pectoral fin is acutely tipped and has 16 to 19 rays. First dorsal fin consists of six separate spines. Second dorsal fin has one spine and 19 or 20 rays, with anterior rays forming very long, acute lobe that extends beyond base of caudal fin in adults. Anal fin consists of two free spines followed by one spine and 16 to 18 rays, with anterior rays forming very long, acute lobe that extends beyond base of caudal fin. Base of anal fin is about equal to base of sec-

ond dorsal fin. Scales are small, cycloid, and partially embedded, and scutes are absent from lateral line. Lateral line is wavy anteriorly and consists of 107 scales. Caudal peduncle lacks upper and lower grooves. Vertebrae number 24: 10 precaudal and 14 caudal. Color is bluish silver dorsally, silvery laterally, and yellowish ventrally, with four prominent dark, narrow bars on upper flank. This species occurs in coastal waters to about 36 m in the western Atlantic from Massachusetts and Bermuda to Argentina, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. Adults are pelagic and are associated with the surf zone over sandy beaches, reefs, and rocky areas. Food consists of invertebrates and small ray-finned fishes. Maximum known size is 506 mm FL. references: Fields 1962; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1999; Smith-Vaniz et al. 1999; Smith-Vaniz 2002c.

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302 PERCIFORMES

Trachurus lathami Nichols, 1920 Rough scad

Trachurus lathami is elongate and slightly compressed, with two distinctly separated dorsal fins and all scales of lateral line scutelike. Dorsal and ventral profiles are about equally convex, and snout is relatively long and acutely pointed. Eye possesses well-developed adipose eyelid. Maxilla extends beyond anterior margin of eye. Jaw teeth are minute and arranged in a single row. Gill rakers on first arch are lathlike and number 12 to 16 on upper limb and 33 to 41 on lower limb. Shoulder girdle has furrow along upper margin but lacks papillae. Branchiostegal rays number 7. Pectoral fin is falcate and has 20 to 21 rays. First dorsal fin has eight spines, and second dorsal fin has one spine and 28 to 34 rays. Anal fin consists of two free spines followed by one spine and 24 to 30 rays. Last fin ray of second dorsal fin and anal fin is connected by membrane to remainder of fins but is displaced posteriorly from penultimate ray by a distance equal to one-half greater than the distance between any other fin rays. Scales are moderately small and cycloid and cover body except for area behind

pectoral fin. Lateral line is moderately arched anteriorly and possesses 68 to 77 scutes. Accessory lateral line is present on dorsolateral surface between eighth dorsal fin spine and fourth dorsal fin ray. Caudal peduncle lacks keels. Vertebrae number 24: 10 precaudal and 14 caudal. Color is light or dark bluish to bluish green dorsally and silvery to whitish on side and ventrally. Area above eye is blackish, and a small, oval black spot is present on operculum. This species occurs in the western Atlantic from the Gulf of Maine to northern Argentina, including the northern and southern Gulf of Mexico. It is a schooling species on continental shelves from 50 to 90 m. Food consists of small invertebrates. Maximum known size is about 330 mm SL. references: Berry and Cohen 1974; Hoese and Moore 1977, 1998; Castro-Aguirre and MárquezEspinoza 1981; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1997; Smith-Vaniz 2002c.

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CARANGIDAE 303

Uraspis secunda (Poey, 1860) Cottonmouth jack Uraspis secunda is oblong and moderately compressed, with bony scutes on straight portion of lateral line, first dorsal fin lower than lobe of second dorsal fin, and chest naked. Dorsal and ventral profiles are about equally convex but slightly asymmetrical, and snout is bluntly pointed. Eye possesses weakly developed adipose eyelid. Maxilla extends to about anterior margin or middle of eye. Jaw teeth are arranged in two to four irregular rows in specimens smaller than 280 mm FL and in a single row in larger specimens. Gill rakers on first arch are lathlike and number 3 to 8 on upper limb and 13 to 16 on lower limb. Branchiostegal rays number 7. Pectoral fin is falcate and has 22 to 24 rays. First dorsal fin has eight spines, and second dorsal fin has one spine and 27 to 32 rays, with anterior rays slightly longer than remainder of rays. Anal fin consists of two free spines followed by one spine and 19 to 23 rays, with anterior rays slightly longer than remainder of rays. The two free anal spines are reabsorbed in specimens greater than 150 mm FL. Scales are small and cycloid. Chest is naked ventrally to half the distance to pectoral fin

base. Lateral line is moderately arched anteriorly and possesses 26 to 40 well-developed scutes on posterior straight section, with some scutes antrorse, or pointing anteriorly. Caudal peduncle has two poorly developed keels. Vertebrae number 24: 10 precaudal and 14 caudal. Color is leaden, bluish black or dusky dorsally and dark olive to bluish in specimens greater than 300 mm FL. Juveniles and specimens up to 300 mm FL have six or seven dark bars on body. Tongue and floor of mouth are white or cream colored, and remainder of mouth is black. This species occurs in the Atlantic and Pacific Oceans. In the western Atlantic it occurs from New Jersey and Bermuda to São Paulo, Brazil, including the eastern and north-central Gulf of Mexico and Santo Domingo in the Greater Antilles. It is largely an oceanic species. Maximum known size is about 435 mm FL.

references: Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith-Vaniz 1986a, 2001b, 2002c; Cervigón 1993a,b.

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304

BRAMIDAE

Pomfrets

Pomfrets are elliptical to teardrop shaped in profile, deep bodied, and moderately to very compressed, with a single elongate dorsal fin and an elongate anal fin that lacks spines. Dorsal profile of head is moderately to very strongly convex. Snout is shorter to slightly longer than orbit diameter. Nostril is paired, with posterior naris slitlike and slightly larger than anterior naris. Eye is moderate to large and laterally located. Mouth is oblique and superior or terminal, with lower jaw projecting beyond upper jaw. Upper jaw extends at least to center of eye. Jaw teeth are conical, villiform, and recurved and occur in bands. Small, conical teeth are present or absent in palatine and vomer. Margin of preoperculum ranges from entire (lacking serrations) to strongly serrated but is entire in adults. Operculum lacks spines. Branchiostegal rays number 6 to 8. Gill rakers on first arch number 7 to 24. Pseudobranch on inner surface of operculum is well developed. Pectoral fin is pointed and falcate in adults and has 17 to 23 rays. Dorsal fin originates from near anterior margin of head to behind pectoral fin base, with first several rays unbranched, and has 31 to 57 rays. Pelvic fin is thoracic and short to very short and has one spine and 5 rays. Anal fin is elongate and similar in shape to dorsal fin and has 21 to 50 rays, with several anterior rays unbranched. Caudal peduncle is narrow, and caudal fin is forked to crescent shaped. Scales are adherent and have spines attached to posterior margin. Scales cover body and most of head, including maxilla, preoperculum, and operculum. Large axillary process is present on base of pelvic fin. Lateral line is poorly developed or absent in adults. Gas bladder is absent. Vertebrae number 36 to 47. Maximum known size is about 100 cm TL. Pomfrets occur worldwide in tropical to temperate oceanic waters. Most species are pelagic between the surface and 600 m, although some are benthic. Most species school and are found at less than 200 m. Food consists of squids, amphipods, shrimps, salps, and small ray-finned fishes. There are about 21 species in seven genera, 9 species in six genera in the western Central Atlantic, and 5 species in four genera in the Gulf of Mexico. references: Mead 1972; Johnson 1984; Smith 1986a; Nelson 1994; Last 2001a; Thompson 2002a.

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BRAMIDAE 305

Key to the Species of the Gulf of Mexico (Adapted from Mead 1972) 1a. Dorsal and anal fins free of scales, flexible, and completely depressible into sheaths of elongated scales inserted along bases of fins . . . . . . . . . . . . . . . Pterycombus brama p. 308 1b. Dorsal and anal fins of adults with scales; dorsal and anal fins of juveniles and adults moderately stiff and not depressible; basal sheath of elongate scales absent from bases of dorsal and anal fins . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Dorsal profile of head above eye not strongly convex, distance between dorsal margin of orbit and nearest point on dorsal profile less than half of eye diameter . . . . . . . . . . . . . . . . . . . . . . . . . . Taractes rubescens p. 309 2b. Dorsal profile of head above eye strongly convex, distance between dorsal margin of orbit and nearest point on dorsal profile greater than half of eye diameter . . . . . . . . . . . . . . 3 3a. Head strongly compressed, with lower edge of lower jaw in contact along ventral midline; scales on caudal peduncle forming graded size series with scales of midcaudal rays .............................................4 3b. Head moderately compressed, with lower edge of lower jaw not in contact along ventral midline (part of isthmus separates lower edge); scales on caudal peduncle abruptly larger than those of midcaudal rays . . . . . . . . . . . . . . . . . . . . Taractichthys longipinnis p. 310 4a. Pelvic fin less than 25% of SL; pectoral fin relatively high on flank, in juveniles distance between lowermost pectoral fin ray and insertion of pelvic fin is greater than 42% of head length, or more than about 12% of SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brama caribbea p. 306 4b. Pelvic fin greater than 25% of SL, pectoral fin relatively low on flank, in juveniles distance between lowermost pectoral fin ray and insertion of pelvic fin is less than 42% of head length, or about 12% of SL . . . . . . . . . . . . . . . . . . . . . . . . . . Brama dussumieri p. 307

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306 PERCIFORMES

Brama caribbea

Mead, 1972

Brama caribbea is relatively short bodied and elliptical, with dorsal fin relatively stiff, covered with scales, and originating over base of pectoral fin. Anterior profile of head is strongly convex. Posterior naris is slitlike. Mouth is terminal, with lower jaw projecting slightly if at all beyond upper jaw and upper jaw extending to center of eye. Premaxilla is separated from antimere by edentulous fold of skin. Teeth of upper jaw consist of outer row of enlarged, recurved canines and inner band of villiform teeth. Lower jaw has outer row of enlarged canines and inner band of villiform teeth on either side of symphysis. Vomer has small patch of villiform teeth, and palatine has a single row of small, depressible teeth. Posterior margins of preoperculum, interoperculum, and operculum are finely serrated. Gill rakers on first arch number 2 to 5 on upper limb and 8 to 11 on lower limb. Measurements are expressed as percent of SL: head length 28%–33%, snout length 6%– 8%, eye diameter 9%–12%, snout to origin of dorsal fin 41%– 49%, snout to origin of anal fin 58%– 65%, body depth 51%– 66%. Pectoral fin has about 20 rays. Dorsal fin is slightly elevated anteriorly in adults and has 32 to 35 rays. Pelvic fin has prominent axillary

scale. Anal fin is low over length and has 27 to 30 rays. Upper lobe of caudal fin is considerably longer than lower lobe. Scales are elongated and imbricate along posterior margin. Scales in horizontal series number about 51, pored lateral line scales number 56, and predorsal scale rows number about 34. Vertebrae number 36 to 38: 15 or 16 precaudal and 20 to 21 caudal. Color is brownish black and slightly darker dorsally than ventrally. Interradial membranes of dorsal and anal fin are black. Lining of mouth is black. This species occurs in the western and possibly eastern Atlantic. In the western Atlantic it occurs between the surface and about 400 m from North Carolina and Bermuda to northern Brazil, including the northern and southwestern Gulf of Mexico and the Greater and Lesser Antilles. Larvae are most abundant between the surface and 50 m. Maximum known size is 270 mm TL. references: Mead 1957 (as Collybus drachme), 1972; Murdy et al. 1983; Shimizu 1983a; Robins and Ray 1986; Thompson and Russell 1996; Smith-Vaniz et al. 1999; Thompson 2002a.

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BRAMIDAE 307

Brama dussumieri Cuvier, 1831

Brama dussumieri is relatively short bodied and elliptical, with dorsal fin relatively stiff, covered with scales, and originating over base of pectoral fin. Anterior profile of head is strongly convex. Posterior naris is slitlike. Mouth is terminal, with lower jaw projecting slightly beyond upper jaw and upper jaw extending to center of eye. Premaxilla is separated from its antimere by edentulous fold of skin. Jaw teeth consist of outer row of enlarged, recurved canines and inner band of villiform teeth. Villiform teeth are either present or absent in palatine and vomer. Posterior margin of preoperculum lacks spines but has fine serrae in adults, and has six or more spines in juveniles. Gill rakers on first arch number 3 to 7 on upper limb and 8 to 12 on lower limb. Measurements for adults and large juveniles are expressed as percent of SL: head length 26%– 29%, snout length 6%– 8%, eye diameter 7%– 8%, snout to origin of dorsal fin 36%– 42%, snout to origin of anal fin 50%– 60%, body depth 44%–57%. Small juveniles have relatively long heads, large eyes, and deep bodies.

Pectoral fin has 21 rays. Dorsal fin is slightly elevated anteriorly in adults and has 33 to 37 rays. Pelvic fin has a prominent axillary process. Anal fin is slightly elevated anteriorly and has 27 to 32 rays (for Atlantic specimens). Upper lobe of caudal fin is slightly longer than lower lobe. Scales are elongated and imbricate along posterior margin. Scales in horizontal series number 57 to 65, and pored lateral line scales number 65. Vertebrae number 42: 16 precaudal and 26 caudal. Color is dark gray, with metallic sheen. Dorsal and anal fins are dark. This species occurs worldwide in tropical seas. In the western Atlantic it occurs between the surface and at least 200 m from Cape Hatteras to northern Brazil, including the northern and southwestern Gulf of Mexico and the Antilles. Maximum known size is 370 mm TL.

references: Mead 1972; Robins and Ray 1986; Thompson and Russell 1996; Last 2001a; Thompson 2002a.

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308 PERCIFORMES

Pterycombus brama

Fries, 1837

Pterycombus brama is teardrop shaped in lateral profile, with dorsal fin originating over head and dorsal and anal fins high and sail-like. Dorsal profile of head is moderately rounded. Posterior naris is slitlike and slightly larger than anterior naris. Mouth is terminal, with lower jaw projecting beyond upper jaw and upper jaw extending to middle of eye. Jaw teeth consist of outer irregular row of enlarged, recurved canines and inner band of villiform teeth, with band narrowing posteriorly. Posterior margin of preoperculum is entire to slightly serrated in adults but consists of a series of graduated spines in juveniles. Gill rakers on first arch number one or two on upper limb and five to seven on lower limb. Measurements are expressed as percent of SL: head length 24%– 29%, snout length 5%–7%, eye diameter 7%– 11%, snout to dorsal fin origin 14%–22%, snout to anal fin origin 28%– 40%, body depth 23%–36%. Pectoral fin has 20 to 22 rays. Dorsal fin originates over posterior rim of orbit in adults and over pectoral fin base in juveniles and has 48 to 53 rays. Pelvic fin is short

and inserts anterior to pectoral fin base. Anal fin originates under pectoral fin base in adults, but under posterior end of abdominal cavity in small juveniles, and has 40 to 43 rays. Dorsal and anal fins are fully depressible into grooves formed by elongated scales along bases of fins. Scales on head lack vertical spines, and scales on body either lack or possess spines. Lateral scale rows number 48 to 53, and predorsal scales number 10 or 11. Lateral line is absent in adults. Vertebrae number 48 to 51: 21 to 23 precaudal and 25 to 28 caudal. Color of dorsum and flanks is dark with silvery sheen. This species occurs in the tropical to temperate Atlantic. In the western Atlantic it occurs between 25 and 300 m from Newfoundland to the Greater Antilles, including the northern and southern Gulf of Mexico and the Bahamas. Maximum known size is about 450 mm TL. references: Mead 1972; Murdy et al. 1983; Robins and Ray 1986; Thompson and Russell 1996; Thompson 2002a.

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BRAMIDAE 309

Taractes rubescens

(Jordan and Evermann, 1887)

Taractes rubescens is elliptical in profile and moderately deep bodied, with anterior section of dorsal and anal fins elevated into lobes, dorsal and anal fins covered with scales, and caudal peduncle bearing well-developed keel. Dorsal contour of head between eyes is flat to slightly arched, and forehead is slightly concave. Mouth is terminal, with lower jaw projecting slightly beyond upper jaw and upper jaw extending to about center of eye. Posterior naris is slitlike and larger than anterior naris. Eye is vertically elliptical. Jaw teeth are small, recurved, and arranged in short bands. Teeth are absent from symphyses of jaws. Palatine has short band of small teeth. Margins of preoperculum and interoperculum are entire (smooth) in adults but serrated or toothed in juveniles. Gill rakers on first arch number 9 to 12, excluding rudiments. Measurements are expressed as percent of SL: head length 28%– 33%, snout length 7%–10%, eye diameter 5%–7%, snout to dorsal fin origin 38%– 45%,

snout to anal fin origin 59%– 67%, body depth 36%– 41%. Pectoral fin has 19 to 22 rays. Dorsal fin originates over pectoral fin base and has 30 to 31 rays. Anal fin has 21 to 23 rays. Caudal fin possesses poorly developed transverse precaudal grooves. Scales on caudal peduncle are abruptly larger than those on proximal part of caudal fin. Lateral line scales become indistinct with growth. Scales lack spines except in juveniles. Color of body and fins is black except for white crescent region on caudal fin. This species occurs in the tropical Atlantic and Central and eastern Pacific. In the western Atlantic it occurs in the northern Gulf of Mexico and off Trinidad. Maximum known size is 850 mm TL. references: Mead 1957, 1972; Mead and Maul 1958; Haedrich 1986a; Robins and Ray 1986; Thompson and Russell 1996; Last 2001a; Thompson 2002a.

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310 PERCIFORMES

Taractichthys longipinnis Bigscale pomfret

(Lowe, 1843)

Taractichthys longipinnis is short and deep bodied, with dorsal and anal fins relatively stiff, covered with scales, and falcate. Anterior profile of head is strongly to moderately convex, becoming less convex with age. Interorbital region is arched. Posterior naris is slitlike and closer to eye than to anterior naris. Mouth is terminal, with lower jaw projecting beyond upper jaw and upper jaw extending beyond center of eye to posterior margin of orbit. Jaw teeth are pointed, conical, recurved, and arranged in bands, with anterior inner teeth largest. Palatine has narrow band of teeth, but vomer lacks teeth. Gill rakers on first arch number one to three on upper limb, one in corner, and five to eight on lower limb. Measurements are expressed as percent of SL: head length 28%– 34%, snout length 7%–10%, eye diameter 5%–10%, snout to dorsal fin origin 41%– 47%, snout to anal fin origin 53%– 60%, body depth 54%– 61%. Pectoral fin has 21 rays. Dorsal fin originates between pectoral fin base and anal fin origin and has 33 to 38 rays. Anal

fin has 27 to 30 rays. Caudal fin is emarginate in juveniles and forked in adults and has welldeveloped precaudal pits. Scales of head and belly lack attached spines on posterior margin, but remainder of scales have spines. Scales in horizontal series number 39 to 46. Lateral line is indistinct. Vertebrae number 44 to 47: 19 to 22 precaudal and 24 or 25 caudal. Color is black, with pectoral and distal part of pelvic fin white and crescent border on caudal fin light. This species occurs in the tropical to temperate Atlantic. In the western Atlantic it occurs from Browns Bank, off Nova Scotia, and Bermuda to northern South America, including the northern and southern Gulf of Mexico and Puerto Rico. Maximum known size is about 910 mm TL. references: Mead 1957, 1972; Robins and Ray 1986; Smith 1986a; Boschung 1992; Thompson and Russell 1996; Smith-Vaniz et al. 1999; Thompson 2002a.

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311

CARISTIIDAE

Manefishes

Manefishes are deep bodied and compressed, with a high and long dorsal fin, long pelvic fins, and an anal fin lacking spines. Profile of head is very steep, and snout is truncated. Mouth is terminal, moderately oblique, and moderately large. Maxilla is partially to completely covered by lachrymal bone when mouth is closed. Nostril is paired and located in front of eye. Preopercular margin is entire, and opercular margin has two weak spines. Branchiostegal rays number six. Pectoral fin is fan shaped, inserts on lower flank, and has a very oblique base. Dorsal fin is sail-like, originates on head, and fits into groove on dorsum. Pelvic fin is very long, inserts anterior to pectoral fin base, has one spine and five rays, and fits into groove that runs along midline of belly to origin of anal fin. Anal fin is elongate and fits into wide, thin skin flap. Anterior rays of dorsal and anal fins are not segmented but are bilaterally paired. Caudal peduncle is short, and caudal fin is truncate. Body and side of head are covered with small, deciduous scales. Lateral line is present (one species) or absent (remaining species). Gas bladder is absent. Vertebrae number 35 to 40. Maximum known size is 265 mm SL. Manefishes occur worldwide in tropical to warm temperate oceanic waters. There are four known species in two genera and perhaps several additional undescribed species. All species are epipelagic to bathypelagic between 100 and 2,000 m but usually between 300 and 800 m. They are often associated with siphonophores. Development is oviparous, and eggs are pelagic. One species occurs in the western Atlantic, including the Gulf of Mexico. references: Fujii 1983c; Johnson 1984; Heemstra 1986i; Post 1986, 1990; Janssen et al. 1989; Tolley et al. 1990; Nelson 1994; Tominaga et al. 1996; Paxton 2001a; McEachran 2002a.

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312 PERCIFORMES

Caristius

sp. cf. maderensis Maul, 1949

Caristius sp. cf. maderensis is deep bodied, squarish in lateral profile, and compressed, with a long and high dorsal fin, long pelvic fins, and lateral line absent. Posterior naris is about twice the size of anterior naris. Maxilla extends to pupil of eye and is covered by lachrymal bone when mouth is closed. Jaw teeth are needlelike, short, slightly recurved, and uniserial. Vomer, palatine, and tongue lack teeth. Gill rakers on first arch are in double rows, with those of outer row long, bladelike, and possessing bristles on lateral surfaces, and those of inner row clublike and possessing terminal bristles. Outer gill rakers number 6 or 7 on upper limb, 1 in corner, and 15 on lower limb. Measurements are expressed as percent of SL: head length 34%–38%, pre-dorsal-fin length 12%–18%, pre-anal-fin length 50%– 53%, body depth 62%–72%; and as percent of head length: snout length 20%–25%, orbit length 36%– 44%, upper jaw length 38%– 44%. Pectoral fin has 16 to 18 rays. Dorsal fin originates over posterior margin of eye and has 29 or 30 rays. Pelvic fin inserts under preoper-

culum, anterior to pectoral fin base. Anal fin originates below midlength of dorsal fin base and has 18 or 19 rays. Caudal peduncle is short and slender. Caudal fin is truncate and has 17 principal rays and 6 or 7 procurrent rays. Body is covered with cycloid scales that vary considerably in size. Juveniles are light brown with three dark brown bars, and adults are light brown with operculum dark brown and fin membranes dark. This species possibly occurs worldwide in tropical and subtropical seas; however, the number of species within the family is unknown, and populations in different areas may represent unrecognized species. Specimens from the Gulf of Mexico differ slightly from the type description of the species and may represent a new species. In the western Atlantic this species is known from the eastern Gulf of Mexico, off Alabama and Florida. Maximum known size of specimens of the western Atlantic population is 247 mm SL. references: Post 1986; Tolley et al. 1990.

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313

EMMELICHTHYIDAE Rovers

Rovers are elongate, oval in profile, and slightly compressed, with very protrusible jaws and maxilla expanded posteriorly. Dorsal and ventral profiles of body are symmetrical. Nostril is paired, and nares are close set. Mouth is terminal, slightly oblique, and moderate in size, with lower jaw projecting slightly beyond upper jaw. Premaxilla has well-developed ascending and midlateral processes and is firmly attached to large rostral cartilage. Supramaxilla is long and slender and partially concealed under lachrymal bone when mouth is closed. Jaws either lack teeth or have small, conical teeth anteriorly, and vomer and palatine lack teeth. Preoperculum is broadly rounded, is slightly projected at corner as thin lamina, and has either smooth or slightly crenate margin. Operculum has two or three flat spines on posterior margin. Branchiostegal rays number 7. Gill rakers on first arch are long and numerous. Pectoral fin is relatively short, inserts on lower flank, and has 18 to 20 rays. Dorsal fin is deeply notched or divided into spinous and rayed sections and has 7 to 13 spines and 9 to 12 rays. Pelvic fin is thoracic and has axillary process, 1 spine, and 5 rays. Midventral scaly process is located between pelvic fins. Anal fin is similar in shape to rayed section of dorsal fin and has 3 spines and 9 to 11 rays. Caudal fin is forked and has 19 principal rays, 17 of which are branched. Most of head, including maxilla, and body is covered with weak ctenoid scales. Scales form sheath at base of rayed section of dorsal and anal fins, and sheath is expanded posteriorly to cover last several rays. Lateral line is continuous and consists of 68 to 73 tubed scales. Vertebrae number 24: 10 precaudal and 14 caudal. Rovers occur worldwide in tropical to warm temperate seas near the bottom between 57 and 500 m. They are schooling fishes that usually occur over drop-offs around islands or deep reefs. Adults and large juveniles (100 mm SL) are benthopelagic, and postlarvae and small juveniles (70 mm SL) are epipelagic. Food consists of zooplankton, especially colonial salps. There are 15 species in three genera, and two species in separate genera occur in the western Central Atlantic and the Gulf of Mexico. references: Heemstra and Randall 1977; Johnson 1984; Heemstra 1986j, 2002d; Nelson 1994; Smith-Vaniz et al. 1999; Carpenter 2001a.

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314 PERCIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Heemstra and Randall 1977; Heemstra 2002d) 1a. Dorsal fin divided to base between spinous and rayed sections, but without distinct gap between sections; base length of spinous section of dorsal fin 24% to 26% of SL; head length 29% to 33% of SL; dorsal fin spines 11 . . . . . . . . . . . . . . . . . . . . . . . . . Erythrocles monodi p. 316 1b. Dorsal fin divided to base between spinous and rayed sections, with distinct gap between sections bearing short, isolated, or buried spines; base length of spinous section of dorsal fin 28% to 31% of SL; head length 25% to 27% of SL; dorsal fin spines 12 or 13, with 7 to 9 spines connected by membrane . . . . . . . . . . . . . . Emmelichthys ruber p. 315

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EMMELICHTHYIDAE 315

Emmelichthys ruber

(Trunov, 1976)

Emmelichthys ruber has a first dorsal fin that exceeds head length and a gap between dorsal fins. Anterior naris is oval and has a broadbased flap on posterior margin, and posterior naris is nearly circular. Maxilla extends to near anterior margin of pupil. Teeth are absent from upper jaw, but a few minute, conical teeth are present in front of lower jaw. Gill rakers on first arch number 8 to 12 on upper limb and 24 to 27 on lower limb. Rear margin of gill cavity bears shallow groove on upper section of cleithrum. Measurements are expressed as percent of SL: head length 25%–27%, orbit diameter 9%–13%, interorbital width 6%–7%, pre-dorsal-fin length 36%– 43%, first dorsal fin base 28%–31%, caudal peduncle depth 9%–12%, body depth 22%–28%. Pectoral fin is moderately pointed and has 19 or 20 rays. First dorsal fin has seven to nine spines connected by membranes, followed by three to five spines free of membranes and buried in flesh in large specimens. Second dorsal fin has one spine and 10 rays. Anal fin has three spines and

9 rays. Scales occur on maxilla of specimens greater than 35 mm SL. Lateral line scales number 71 to 74. Scale rows between dorsal fin origin and lateral line number 5 or 6, scale rows between lateral line and anal fin origin number 14 or 15, and scales encircling caudal peduncle number 28 to 30. Color is reddish dorsally and silvery white laterally and ventrally. Maxilla, lower jaw, and ventrolateral section of head are silvery white. Fins are light pink to transparent, except caudal fin is reddish. This species occurs between 180 and 200 m in the tropical to warm temperate waters of the Atlantic Ocean from St. Helena, Jamaica, Bermuda, and the eastern Gulf of Mexico. In the Gulf of Mexico it has been reported once from Naples, FL (2620N, 8410W). Postlarvae and small juvenile specimens have been collected between the surface and 100 m in deep water. Maximum known size is 230 mm TL. references: Trunov 1976; Heemstra and Randall 1977; Ruiz-Carus et al. 2002; Heemstra 2002d.

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316 PERCIFORMES

Erythrocles monodi Crimson rover

Poll and Cadenat, 1954

Erythrocles monodi has a midlateral keel on caudal peduncle and lacks a space between dorsal fins. Anterior naris is elliptical or slit, and posterior naris is nearly circular. Orbit is slightly larger than snout length in juveniles but slightly smaller than snout in adults. Jaws are toothless or possess a few minute, conical teeth in anterior sections. Juveniles have a narrow band of cardiform teeth on each side of vomer and two or three teeth on palatine, but these are absent in adults. Preopercular margin is smooth, crenulate, or weakly serrate. Gill rakers on first arch number 9 to 12 on upper limb and 27 to 29 on lower limb. Measurements are expressed as percent of SL: head length 29%– 33%, orbit diameter 8%–10%, interorbital width 7%–9%, pre-dorsal-fin length 36%– 39%, caudal peduncle depth 7%–9%, body depth 26%–27%. Small specimens (114 mm SL) have a relatively short head (27%–32%) and shallow body (24%–26%). Pectoral fin

has 18 to 20 (rarely 18) rays, dorsal fin has 11 spines and 11 or 12 rays, and anal fin has 3 spines and 9 or 10 rays. Lateral line scales number 68 to 72. Scale rows between dorsal fin origin and lateral line number 7 to 9, scale rows between lateral line and anal fin origin number 15 to 18, and scales encircling caudal peduncle number 30 to 32. Color is reddish dorsally and silvery laterally and ventrally. Dorsal, pelvic, and anal fins are pinkish white, and pectoral and caudal fins are red. This species occurs in the tropical to warm temperate waters of the Atlantic Ocean between 100 and 300 m. In the western Atlantic it occurs from South Carolina and the Bahamas to Venezuela, including the northern Gulf of Mexico and the Windward Islands of the Lesser Antilles. Maximum known size is 550 mm TL. references: Heemstra 1972, 2002d; Heemstra and Randall 1977; Robins and Ray 1986.

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317

LUTJANIDAE Snappers

Snappers are relatively slender to relatively deep bodied and moderately compressed, with a single dorsal fin, generally large canine teeth in jaws, and maxilla covered (more or less) by lachrymal bone when mouth is closed. Dorsal profile of head is more or less straight. Snout is short to relatively long. Nostril is paired, and nares are close set, with posterior naris elongated and slightly larger than anterior naris. Mouth is terminal, moderately large to large, and moderately to strongly oblique. Upper jaw is moderately protrusible, premaxilla has broad midlateral process, and supramaxilla is absent. Villiform teeth occur in jaws, vomer (rarely absent), and palatines. Large conical or canine teeth generally also occur in jaws. Preoperculum is generally serrated along posterior and ventral margins and may be notched along posterior margin. Operculum has 2 flat spines directed posteriorly. Branchiostegal rays number 7. Gill membranes are separate and free of isthmus, and last full arch (fourth) has slit posteriorly. Pectoral fin is well developed and narrowly pointed, with nearly vertical base and upper rays considerably longer than lower rays. Dorsal fin is unnotched to slightly notched and consists of 10 to 12 (rarely 9 or 13) spines and 10 to 19 rays. Pelvic fin is well developed but smaller than pectoral fin, inserts below pectoral fin base, and has 1 spine and 5 rays. Anal fin is equal to or shorter than rayed section of dorsal fin and has 3 spines and 7 to 11 rays. Caudal fin is truncate to deeply forked and consists of 16 to 26 procurrent rays and 17 principal rays. Scales are ctenoid and of moderate size and cover body, cheeks, operculum, and occasionally maxilla, but not snout, preorbital region, or lower jaw. Axillary process on base of pelvic fin is well developed. Lateral line is complete. Vertebrae number 24: 10 precaudal and 14 caudal. Maximum known size 160 cm TL Snappers occur worldwide in tropical to warm temperate seas between the shoreline and 550 m on continental and insular shelves and slopes. Several Indo-Pacific species of Lutjanus occur only in fresh and brackish waters, and juvenile stages of many species occur in fresh and brackish waters. However, most species are associated with marine reefs. All species are thought to be gonochoristic and display no sexual dimorphism. Eggs and larvae are pelagic. Eggs range from 0.65 to 1.02 mm in diameter and have a single oil droplet. Larvae have well-developed head spines and elongated dorsal and pelvic spines. Maximum known life span ranges from 4 to 42 years. Most species are nocturnal and feed on either benthic organisms or plankton. Food consists of gastropods, cephalopods, shrimps, crabs, urochordates, and rayfinned fishes. Ray-finned fishes are by far the most numerous food resource. There are about 125 species in 21 genera, 18 species in

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318 PERCIFORMES

6 genera occur in the western Central Atlantic, and 16 or 17 species in 6 genera occur in the Gulf of Mexico. Lutjanus ambiguous is regarded as a hybrid of L. synagris and Ocyurus chrysurus, following Loftus (1992) and Domeier and Clarke (1992). references: Anderson 1967, 1986, 2002c; Johnson 1980, 1984; Allen 1985; Nelson 1994.

Key to the Species of the Gulf of Mexico (Adapted from Anderson 1967; Allen 1985) 1a. Bases of dorsal and anal fin rays with covering of scales; last dorsal and anal fin rays not produced . . . . . . . . . . . . 2 1b. Bases of dorsal and anal fin rays without covering of scales; last dorsal and anal fin rays somewhat produced . . . . . . 14 2a. Gill rakers on lower limb of first arch (excluding rudiments) 17 to 22 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2b. Gill rakers on lower limb of first arch (excluding rudiments) 16 or fewer (some specimens of Lutjanus buccanella may have 17) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3a. Dorsal fin with 12 or 13 spines and 10 or 11 (usually 11) rays; caudal fin forked, without pointed lobes; vomerine tooth patch with broad, medial posterior extension . . . . . . . . . . . . . . . . . . . . Rhomboplites aurorubens p. 337 3b. Dorsal fin with 10 spines and 12 to 14 rays; caudal fin deeply forked, with pointed lobes; vomerine tooth patch with narrow, medial posterior extension . . . . . . . . . . . . . . . . . . . . . . . . . Ocyurus chrysurus p. 334 4a. Dorsal fin with 10 spines and 12 (rarely 11 or 13) rays; black spot on upper flank below anterior dorsal fin rays (present on specimens of all sizes but occasionally absent in Lutjanus synagris) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4b. Dorsal fin with 10 (rarely 9 or 11) spines and 14 (rarely 13 or 15) rays; black spot on upper flank below anterior dorsal fin rays present or absent . . . . . . . . . . . . . . . . . . . . . . 6 5a. About one-fourth to one-half of black lateral spot extending below lateral line; 7 or 8 gill rakers on upper limb and 15 to 17 gill rakers on lower limb of first arch . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus mahogoni p. 330 5b. Less than one-fourth to none of black lateral spot extending below lateral line in specimens above 60 mm SL; 6 or 7 gill rakers on upper limb and 13 or 14 (rarely 12 or 15) gill rakers on lower limb of first gill arch . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus synagris p. 332 6a. Large, distinct black spot at base and in axil of pectoral fin; no black spot below anterior rays of dorsal fin; anal fin rounded; dark area on scales at base of dorsal fin rays; iris of eye golden yellow to orange in life . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus buccanella p. 325

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LUTJANIDAE 319

6b. No large, distinct black spot at base and in axil of pectoral fin; black spot below anterior rays of dorsal fin present or absent; anal fin rounded or angular; no dark area at base of dorsal fin rays; iris of eye not golden yellow to orange in life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7a. Anal fin rounded, with middle rays less than half the length of head (throughout life); no black spot below anterior rays of dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7b. Anal fin angular, with middle rays almost half to more than half of head length (in specimens greater than 40 mm SL for L. analis, 50 mm SL for L. campechanus, and 60 mm SL for L. vivanus); black spot below anterior dorsal fin rays, at least in small specimens (to 465 mm SL in L. analis and to 200 to 300 mm SL in L. campechanus and L. vivanus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 8a. Vomerine tooth patch without distinct medial posterior extension; canine teeth in upper and lower jaws strong and equally developed; cheek scales in 8 to 10 (usually 9) rows . . . . . . . . . . . . . . . . . . . . . . . Lutjanus cyanopterus p. 327 8b. Vomerine tooth patch with distinct medial posterior extension; canine teeth in upper jaw usually larger than those of lower jaw; cheek scales 5 to 9 (usually 7 or 8) rows . . . . . 9 9a. Pectoral fin length about equal to distance from tip of snout to posterior margin of preoperculum, or 24% to 39% of SL; body depth 31% to 39% of SL (usually 32% to 37% of SL) . . . . . . . . . . . . . . . . Lutjanus griseus p. 328 9b. Pectoral fin length longer than distance between tip of snout and posterior margin of preoperculum, or 29% to 33% of SL (in L. apodus between 75 to 96 mm SL, pectoral fin is 24% to 39% of SL); body depth 36% to 43% of SL (usually 37% to 42% SL) . . . . . . . . . . . . . . . . . . . 10 10a. Lateral line scales 40 to 45; scales between dorsal fin base and lateral line number 5 to 7; no whitish bar below eye . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus apodus p. 324 10b. Lateral line scales 46 to 49; scales between dorsal fin base and lateral line number 8 to 11; whitish bar between eye and area posterior to maxilla . . . . . . . Lutjanus jocu p. 329 11a. Vomerine tooth patch without distinct posteromedial extension; anal fin rays eight (rarely seven); iris red in life; spot on flank below anterior dorsal fin rays, large in small specimens and small but distinct in larger specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus analis p. 323 11b. Vomerine tooth patch with distinct posteromedial extension; anal fin rays seven to nine (usually eight or nine) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12a. Anal fin rays nine (rarely eight); lateral line scales 46 to 51 (usually 47 to 49); scales from dorsal fin base to lateral line 7 to 10 (usually 8 or 9); scales on anterior part of body below lateral line distinctly larger than those

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320 PERCIFORMES

on posterior part of body below lateral line . . . . . . . . . . . . . . . . . . . . . . Lutjanus campechanus p. 326 12b. Anal fin rays eight (rarely seven or nine); lateral line scales 49 to 53 (usually 50 or 51); scales from dorsal fin base to lateral line 9 to 12 (usually 10 to 12); scales on anterior part of body below lateral line not distinctly larger than those on posterior part of body below lateral line . . . . . 13 13a. Scales from lateral line to anal fin base 16 to 19; scales from dorsal fin base to lateral line 9 to 11 (usually 10); cheek scale rows six (rarely five or seven); scales on anterior part of body above lateral line smaller than those on anterior part of body below lateral line; pelvic fin length 53% to 62% of SL; spot on flank under anterior dorsal fin rays of young specimens equal to or larger than eye; iris red in life . . . . . . . . . . . . . . . . . Lutjanus purpureus p. 331 13b. Scales from lateral line to anal fin base 20 to 24 (usually 21 to 23); scales from dorsal fin base to lateral line 10 to 12 (usually 11 or 12); cheek scale rows seven (rarely eight); scales on anterior part of body above lateral line similar in size to those on anterior part of body below lateral line; pelvic fin length 63% to 76% of SL; spot on flank under anterior dorsal fin rays of young specimens smaller than eye; iris yellow in life . . . . . . . . . . Lutjanus vivanus p. 333 14a. Dorsal fin deeply notched between spinous and rayed sections; maxilla scaled . . . . . . . . . . . . . Etelis oculatus p. 322 14b. Dorsal fin not deeply notched between spinous and rayed sections; maxilla scaleless . . . . . . . . . . . . . . . . . . . . . . . . 15 15a. Interorbital region convex; last ray of dorsal and anal fins slightly produced; color in life dusky violet dorsally and pale ventrally; color in preservative dark grayish brown . . . . . . . . . . . . . . . . . . . . . . . . . . . Apsilus dentatus p. 321 15b. Interorbital region flattened; last ray of dorsal and anal fins moderately produced; color in life reddish; color in preservative pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16a. Lateral line scales 48 to 52; total gill rakers on first arch 24 to 28 . . . . . . . . . . . . . . . Pristipomoides aquilonaris p. 335 16b. Lateral line scales 54 to 57; total gill rakers on first arch 19 to 25 . . . . . . . . . . . Pristipomoides macrophthalmus p. 336

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LUTJANIDAE 321

Apsilus dentatus Black snapper

Guichenot, 1853

Apsilus dentatus is violet to dark brown and has a strongly convex interorbital region and slightly produced last dorsal and anal fin rays. Lower jaw slightly projects beyond upper jaw, and upper jaw extends to near anterior margin of eye. Jaw has inner band of villiform teeth and outer band of conical teeth, with anterior teeth in upper and lower jaws caninelike. Vomerine tooth patch is V-shaped or triangular shaped, without medial posterior extension. Palatine teeth are arranged in band. Gill rakers on lower limb of first arch (including rudiments) number 15 or 16. Pectoral fin is long and pointed and has 15 or 16 rays. Dorsal fin is not deeply incised between spinous and rayed sections and has 10 spines and 9 or 10 rays. Anal fin has 8 rays. Caudal fin is forked to emarginate. Tubed lateral line scales number 58 to 63. Membranes of dorsal and anal fins lack scales. Scale rows on back are parallel with lateral line. Dark coloration of dorsum shades to pale violet to brown ventrally. Fins are brown

to black, and caudal fin has pale distal margin. Fins of some individuals have blue cast. Iris of eye is bronze surrounded with a dark ring. Small juveniles are bright blue. This species occurs between 40 and 242 m in the western Atlantic from the northern Gulf of Mexico and the Florida Keys to Venezuela, including the Bahamas and the Greater and Lesser Antilles. In the Gulf of Mexico it has been captured near the West Flower Garden Bank at 40 m. It is generally associated with rocky bottoms. In Jamaica this species spawns throughout most of the year, but the most active spawning occurs in the colder months. Food consists of benthic cephalopods, tunicates, and ray-finned fishes. Maximum known size is 550 mm SL.

references: Böhlke and Chaplin 1968; Allen 1985; Robins and Ray 1986; Trimm and Searcy 1989; Anderson 2002c.

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322 PERCIFORMES

Etelis oculatus (Valenciennes, 1828) Queen snapper

Etelis oculatus has a scaled maxilla, a notched dorsal fin, and naked dorsal and anal fin membranes. Snout is shorter than eye diameter in juvenile specimens and about equal to eye diameter in adults. Interorbital space is flat. Lower jaw projects slightly beyond upper jaw, and maxilla extends to posterior margin of eye. Jaw teeth are small, conical, and arranged in bands, with those of outer row slightly enlarged and relatively widely spaced. One or two teeth on each side of upper jaw and frequently on each side of lower jaw are caninelike. Vomerine teeth are in a narrow V-shaped patch or, rarely, in a narrow triangular patch. Gill rakers on first arch, including rudiments, number 7 to 11 on upper limb and 14 to 18 on lower limb. Measurements are expressed as percent of SL: head length 33%–34%, snout length about 9%, eye diameter 11%–12%, upper jaw length 15%– 16%, pectoral fin length 24%–26%, pelvic fin length 18%–21%, body depth 27%–29%. Pectoral fin is slender and has 15 or 16 rays. Dorsal fin has 10 spines and 11 rays. Anal fin has 8 rays. Last ray of dorsal fin and anal fin is considerably longer than penultimate rays.

Caudal fin is deeply forked, with upper lobe slightly longer than lower lobe. In some large specimens, the upper caudal lobe is appreciably longer than lower lobe. Lateral line scales number 47 to 50. Scale rows on back are parallel with lateral line. Color is deep pink to red on dorsal half of body and pale pink to silvery on remainder of body. Rayed section of dorsal fin, pectoral fin, pelvic fin, and anal fin is pink. Spinous section of dorsal fin and caudal fin is brilliant red. This species occurs between 135 and 450 m in the western Atlantic from North Carolina and Bermuda to Brazil, including the eastern and northern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is rare in the northern Gulf of Mexico. It is associated with rocky bottoms. Food consists of squids, crustaceans, and small ray-finned fishes. Maximum known size is about 700 mm SL.

references: Anderson 1972, 1981, 2002c; Allen 1985; Burgess and Branstetter 1985; Robins and Ray 1986; Smith-Vaniz et al. 1999.

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LUTJANIDAE 323

Lutjanus analis (Cuvier, 1828) Mutton snapper

Lutjanus analis has relatively small canine teeth in both jaws, a small dark spot on upper flank, and blue lines on snout. Interorbital space is convex. Lower jaw is level with upper jaw, and maxilla extends to about anterior margin of orbit. Jaws have an outer row of small canine teeth and an inner band of villiform teeth. Vomerine tooth patch is V-shaped or crescentic, without a posteromedial extension. Preoperculum has coarsely serrated posterior margin. Gill rakers on first arch, including rudiments, number 18 to 21, with 6 to 8 on upper limb and 12 or 13 on lower limb. Measurements are expressed as percent of SL: head length 36%– 41%, snout length 11%–17%, eye diameter 5%–10%, maxilla length 13%–17%, pectoral fin length 27%–31%, body depth 36%– 40%. Pectoral fin extends to level of anus and has 15 to 17 rays. Dorsal fin has 10 (rarely 9 or 11) spines and 14 (rarely 13) rays. Anal fin is angular and has 8 (rarely 7) rays, with middle rays at least half of head length in specimens greater than 40 mm SL. Caudal fin is lunate to moderately forked. Scales in horizontal series number 66 to 70, and pored lateral line scales number 47 to 51 (usually 48 or 49). Scale rows on back rise obliquely above lateral line. Scales between dorsal fin base and lateral line number 4, and scale rows on cheek number five or six. Scales cover proximal parts of membranes of rayed sections of dorsal and anal fins. Color is olive green on dorsal half of body and whit-

ish with red tinge on remainder of body. Black spot, about the size of pupil of eye, is located on flank above lateral line and below rayed section of dorsal fin. A pair of blue stripes occurs on snout and cheek, with upper one extending to opercular margin. Dorsal fin and upper lobe of caudal fin are olive, and other fins and lower lobe of caudal fin are red. Iris of eye is red. This species occurs in the western Atlantic from Massachusetts and Bermuda (waif occurrence or introduced) to southeastern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the coastal waters of the Gulf of Mexico and is associated with coral reefs, sandy bottoms, and sea grasses, including estuaries and bays with mangroves. It is a solitary species except during spawning season, when it occurs in large aggregations. Food consists of gastropods, cephalopods, shrimps, crabs, and ray-finned fishes. Spawning takes place during the winter in the northeastern Caribbean Sea. Maximum known age is 14 years. Maximum known size is 800 mm SL.

references: Longley and Hildebrand 1941 (as Lutianus analis); Anderson 1967, 2002c; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999.

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324 PERCIFORMES

Lutjanus apodus Schoolmaster

(Walbaum, 1792)

Lutjanus apodus has a pair of enlarged canines in upper jaw, a rounded anal fin, and narrow bars on side. Interorbital space is convex. Maxilla extends to anterior margin of orbit. Upper jaw has an outer row of canine teeth, four of which are large and two of which are equal to orbital length, and an inner row of villiform teeth. Lower jaw has an outer row of large teeth, including several canines on side of jaw, and an inner row of villiform teeth. Vomerine teeth are in a V-shaped or crescentic patch, with a posteromedial extension. Preoperculum has serrated posterior margin and corner but is entire on ventral margin. Gill rakers on first arch, including rudiments, number 5 to 7 on upper limb and 11 to 15 on lower limb. Measurements are expressed as percent of SL: head length 40%– 42%, snout length 16%–17%, eye diameter 8%–10%, upper jaw length 15%– 16%, pectoral fin length 29%–33%, body depth 36%– 43%. Pectoral fin has 16 or 17 rays. Dorsal fin is shallowly notched and has 10 spines and 14 rays. Anal fin is rounded and has 8 rays, with middle rays less than half of head length. Caudal fin is slightly emarginate to truncate. Tubed lateral line scales number 40 to 45 (usually 42 to 44). Scale rows on anterior section of back are parallel to lateral line. Scale rows of cheek number five. Scales cover

proximal parts of membranes of rayed sections of dorsal and anal fins. Color in life is olive gray to brownish on upper half of body, with eight narrow, pale vertical bars on flank, and whitish with reddish cast on remainder of body. Solid or broken blue line runs under eye from upper jaw to posterior margin of operculum. Bars and blue line may disappear with growth. This species occurs in the tropical to warm temperate Atlantic. In the western Atlantic it occurs from Massachusetts and Bermuda to northeastern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the shallow coastal waters of the Gulf of Mexico over a variety of bottom types. Juveniles often inhabit brackish water. Food consists of polychaetes, gastropods, cephalopods, shrimps, crabs, and ray-finned fishes. Feeding takes place mostly during the night. Maximum known size is 620 mm SL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Allen 1985; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Anderson 2002c.

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LUTJANIDAE 325

Lutjanus buccanella Blackfin snapper

(Cuvier, 1828)

Lutjanus buccanella has relatively large canine teeth in upper jaw and a black blotch at base of pectoral fin. Interorbital space is slightly convex. Maxilla extends nearly to anterior margin of pupil. Jaws have an outer row of enlarged teeth and an inner band of villiform teeth, with four canines in upper jaw, two of which are longer than half of pupil diameter. Vomerine teeth are in a V-shaped or crescentic patch, with a posteromedial extension. Preoperculum has serrated posterior and ventral margins, with serrae large at corner and on ventral margin. Gill rakers on first arch number 7 to 9 on upper limb and 17 to 19 on lower limb. Measurements are expressed as percent of SL: head length 37%– 42%, eye diameter 8%–12%, snout length 14%–15%, lower jaw length 15%–16%, pectoral fin length 31%–33%, pelvic fin length 22%–24%, body depth 34%– 40%. Pectoral fin has 16 to 18 rays. Dorsal fin is shallowly notched and has 10 spines and 14 rays. Anal fin is rounded and has 8 rays, with middle rays less than half of head length. Caudal fin is emarginate to moderately forked. Scales in horizontal series number 62 to 68, and tubed lateral line scales number 47 to 50 (usually 48 or 49). Scale rows on back rise obliquely above lateral line. Scales cover proximal parts

of membranes of rayed sections of dorsal and anal fins. Color is scarlet red on dorsal half of body and silvery on remainder of body, with a black spot at base and on axil of pectoral fin and a dark area along base of dorsal fin. Iris is yellow to orange, and fins are yellowish to orange. Juveniles to about 160 mm SL are yellow to greenish yellow along upper part of caudal peduncle, on rayed sections of dorsal and anal fins, and over entire caudal fin. This species occurs between 80 and 230 m in the western Atlantic from Massachusetts and Bermuda to southeastern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. Juveniles occur between 35 and 50 m. Adults are associated with sandy and rocky bottoms, drop-offs, and ledges. Food consists of cephalopods, crustaceans, tunicates, and ray-finned fishes. Spawning occurs over most of the year but most actively in April and September. Maximum known size is 660 mm SL. references: Böhlke and Chaplin 1968; Allen 1985; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Randall 1996; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Anderson 2002c.

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326 PERCIFORMES

Lutjanus campechanus Red snapper

(Poey, 1860)

Lutjanus campechanus has a pointed anal fin and lacks dark blotch on base of pectoral fin. Interorbital space is slightly convex. Jaws have an outer row of enlarged teeth, with four canines up to half of pupil diameter in length, and an inner band of villiform teeth. Vomerine tooth patch is V-shaped or crescentic, with a posteromedial extension. Tongue has two patches of granular teeth. Preoperculum has serrated posterior and ventral margins, with serrae enlarged at corner and on ventral margin. Gill rakers on first arch number 6 to 8 on upper limb and 14 to 16 on lower limb. Measurements are expressed as percent of SL: head length 38%– 44%, snout length 13%–15%, eye diameter 10%–11%, upper jaw length 15%–16%, pectoral fin length 25%–33%, body depth 40%– 43%. Pectoral fin has 15 to 18 (usually 17) rays. Dorsal fin is shallowly notched between spinous and rayed sections and has 10 spines and 14 (rarely 13 or 15) rays. Anal fin has 8 or 9 (usually 9) rays, and in specimens greater than 50 mm SL, it is pointed, with middle rays as long as or longer than half of head length. Caudal fin is truncate to lunate, with upper lobe slightly longer than lower lobe. Scales in horizontal series number 46 to 50, and tubed lateral line scales number 46 to 51 (usually 47 or 48). Scale rows on back rise obliquely above lateral line, and scales on an-

terior section of body below lateral line are larger than scales on posterior section of body below lateral line. Scales cover proximal parts of membranes of rayed sections of dorsal and anal fins. Color is scarlet to brick red dorsally and rosy ventrally. Iris of eye is red. Fins are red, and caudal fin has black posterior margin. Juveniles have black spot on lateral line. Specimens less than 300 to 350 mm SL have large black spot on upper flank below anterior dorsal fin rays. This species occurs between 10 and 190 m in the western Atlantic from Massachusetts to Yucatán, including the entire Gulf of Mexico. Juveniles inhabit shallow water on sand and mud bottoms. Adults are associated with rocky bottoms and reefs. Food consists of polychaetes, cephalopods, shrimps, crabs, and rayfinned fishes. In the northeastern Gulf of Mexico and Campeche Bank, spawning occurs from July through October. Maximum known age is 10 years, and maximum known size is 100 cm SL. references: Rivas 1966; Anderson 1967, 2002c; Hastings et al. 1976; Hoese and Moore 1977, 1998; Allen 1985; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999.

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LUTJANIDAE 327

Lutjanus cyanopterus Cubera snapper

(Cuvier, 1828)

Lutjanus cyanopterus has relatively large canine teeth in jaws and a rounded anal fin. Interorbital space is slightly convex. Maxilla extends to near middle of eye. Jaws have an outer row of enlarged teeth, including four large canines in upper jaw and four to six slightly larger canines in lower jaw, and an inner band of villiform teeth. Vomerine tooth patch is crescentic or triangular shaped, without a posteromedial extension. Preoperculum is serrate along posterior margin and entire along ventral margin, with serrae enlarged at corner. Gill rakers, including rudiments, on first arch number 5 to 7 on upper limb and 11 to 14 on lower limb. Measurements are expressed as percent of SL: head length 36%–37%, eye diameter 4%– 6%, snout length 14%–15%, lower jaw length 15%–16%, pectoral fin length about 27%, pelvic fin length 18%–19%, body depth 29%– 32%. Pectoral fin has 16 to 18 rays. Dorsal fin is slightly notched between spinous and rayed sections and has 10 spines and 14 rays. Anal fin has 7 or 8 rays, and middle rays are less than half of head length. Caudal fin is slightly emarginate to truncate. Scales in horizontal series number 56 to 59, and tubed lateral line scales

number 45 to 47. Scale rows on back rise obliquely above lateral line. Scales cover proximal parts of membranes of rayed sections of dorsal and anal fins. Color is pale to dark gray with a reddish tinge on upper half of body and pale on remainder of body. Dorsal and caudal fins are grayish, pelvic and anal fins are reddish, and pectoral fin is translucent or grayish. This species occurs to 40 m in the western Atlantic from Nova Scotia and Bermuda to northern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the coastal waters of the Gulf of Mexico. Adults are associated with rocky bottoms and reefs. Juveniles inhabit mangrove habitats. Food consists of shrimps, crabs, and ray-finned fishes. Maximum known size is about 160 cm TL.

references: Anderson 1967, 2002c; Böhlke and Chaplin 1968; Moe 1968; Hoese and Moore 1977, 1998; Allen 1985; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Randall 1996; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999.

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328 PERCIFORMES

Lutjanus griseus Gray snapper

(Linnaeus, 1758)

Lutjanus griseus has a rounded anal fin and a black margin along spinous section of dorsal fin. Snout is relatively long and acute. Interorbital space is slightly convex. Maxilla extends to anterior margin of pupil. Jaws have an outer row of enlarged teeth, with four canines in upper jaw, and an inner narrow band of villiform teeth. Vomerine tooth patch is V-shaped or crescentic, with a posteromedial extension. Tongue has an oval patch of small teeth. Preoperculum has serrated posterior margin, with serrae enlarged at corner. Gill rakers on first arch number 6 to 8 on upper limb and 12 to 14 on lower limb. Measurements are expressed as percent of SL: head length 36%– 40%, snout length 13%–14%, eye diameter 8%–10%, upper jaw length 14%–15%, pectoral fin length 24%–27%, body depth 31%–38%. Pectoral fin has 15 to 17 rays. Dorsal fin is slightly notched between spinous and rayed sections and has 10 spines and 13 or 14 (usually 14) rays. Anal fin has 7 or 8 (usually 8) rays, and middle rays are less than half of head length. Caudal fin is emarginate, with upper lobe slightly longer than lower lobe. Scales in horizontal series number 50 or 51, and tubed lateral line scales number 43 to 47. Scale rows on back run parallel with lateral line. Scales cover proximal parts of membranes of rayed sections of dorsal and anal fins. Color is gray, greenish gray, or dark olive, occasionally with a reddish tinge on dorsal half of body and grayish with a reddish tinge on remainder of body. Fins are

grayish or reddish, and spinous section of dorsal fin has black margin. Juveniles usually have a broad, oblique dark stripe from tip of snout through eye to base of spinous dorsal fin, and often have a blue line on cheek and pale bars on side. This species occurs in the Atlantic Ocean from the shoreline to 180 m. In the western Atlantic it occurs from Massachusetts and Bermuda to southeastern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout coastal waters in the Gulf of Mexico and is associated with a variety of habitats, including coral reefs, rocky substrates, estuaries, and mangrove regions. Food consists of gastropods, cephalopods, shrimps, crabs, and ray-finned fishes. Feeding generally takes place at night. Spawning occurs from May to September around times of the full moon. Maximum known age is 24 years, maximum known size is 920 mm SL, and maturity is reached at about 180 to 330 mm SL.

references: Longley and Hildebrand 1941 (as Lutianus griseus); Anderson 1967, 2002c; Böhlke and Chaplin 1968; Randall 1968a, 1996; Smith et al. 1975; Hastings et al. 1976; Hoese and Moore 1977, 1998; Allen 1985; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999.

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LUTJANIDAE 329

Lutjanus jocu Dog snapper

(Bloch and Schneider, 1801)

Lutjanus jocu has a pair of large canines in upper jaw and a white bar on side of head. Interorbital space is slightly convex. Maxilla extends to anterior margin of orbit. Jaws have an outer row of enlarged teeth, including four canines, two of which in upper jaw are about as long as orbit diameter, and a narrow inner row of villiform teeth. Vomerine tooth patch is Vshaped or crescentic, with an elongate posteromedial extension. Tongue has a single large oval patch of small teeth. Preoperculum has serrated posterior margin, with serrae largest at corner. Gill rakers on first arch number 6 to 8 on upper limb and 12 to 14 on lower limb. Measurements are expressed as percent of SL: head length 39%– 40%, eye diameter 6%– 9%, snout length 15%–16%, lower jaw length 15%–16%, pectoral fin length 29%–33%, pelvic fin length 19%–21%, body depth 36%– 43%. Pectoral fin has 16 or 17 (usually 17) rays. Dorsal fin is shallowly notched between spinous and rayed sections and has 10 spines and 13 or 14 (usually 14) rays. Anal fin is rounded and has 8 rays, with middle rays less than half of head length. Caudal fin is slightly emarginate to weakly forked. Lateral scales number 46 to 49, and tubed lateral line scales number 46 to 49. Scale rows on back rise obliquely above lateral line. Scales cover proximal parts of membranes of rayed sections of dorsal and anal fins. Color is olive brown with

a bronze tinge and occasionally narrow pale bars on upper half of body, and reddish with a copper tinge on remainder of body. Series of blue spots run from below eye to operculum, and a pale whitish triangular bar may be present between lower edge of eye and corner of mouth. Juveniles have a horizontal blue line below eye. Fins are brown to yellowish orange. This species occurs in the western Atlantic from Massachusetts to northern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico in coastal waters. There is a report that it was introduced to Bermuda, but this claim is unsubstantiated (Smith-Vaniz et al. 1999). Adults are associated with coral reefs, and juveniles are located in coastal waters, estuaries, and brackish water. Food consists of gastropods, cephalopods, shrimps, crabs, and ray-finned fishes. Spawning occurs in winter in the Greater Antilles. Maximum known size is 600 mm SL, and maturity is reached at about 300 to 400 mm SL.

references: Longley and Hildebrand 1941 (as Lutianus jocu); Anderson 1967, 2002c; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Allen 1985; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Smith 1997; Schaldach et al. 1997; Castro-Aguirre et al. 1999.

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330 PERCIFORMES

Lutjanus mahogoni Mahogany snapper

(Cuvier, 1828)

Lutjanus mahogoni has a dark spot on lateral line and a moderately developed preopercular notch and interopercular knob. Interorbital space is slightly convex. Maxilla extends to anterior margin of pupil, and lower jaw projects slightly beyond upper jaw. Jaws have an outer row of enlarged teeth, including four canines in upper jaw, two of which are about as long as half of pupil diameter, and an inner band of villiform teeth. Vomerine tooth patch is Vshaped or crescentic, with a short posteromedial process. Tongue has a long, narrow patch of small teeth. Preoperculum has finely serrated posterior margin, produced corner with coarse serrations, and entire ventral margin. Gill rakers on first arch number 7 or 8 on upper limb and 15 to 17 on lower limb. Measurements are expressed as percent of SL: head length 38%– 40%, eye diameter 8%–12%, snout length 13%–15%, lower jaw length 16%–17%, pectoral fin length 28%–31%, pelvic fin length 21%–23%, body depth 33%–36%. Pectoral fin has 14 or 15 rays. Dorsal fin is slightly notched between spinous and rayed sections and has 10 spines and 11 or 12 (usually 12) rays. Anal fin is rounded and has 8 rays, with middle rays less than half of head length. Caudal fin is moderately forked. Scales in horizontal series number 57 to 65, and tubed lateral line scales number 47 to 49. Scale rows

on back rise obliquely above lateral line. Scales cover proximal parts of membranes of rayed sections of dorsal and anal fins. Color is gray to dark olive on upper half and silvery on remainder of body, usually with a red tinge over entire body. A black spot, equal to eye diameter, is located on upper flank on lateral line below anterior dorsal fin rays. About one-quarter to one-third of black spot is located below lateral line. Fins are reddish to yellow, and caudal fin has a dusky posterior margin. This species occurs in the western Atlantic from North Carolina to Venezuela, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. Records of this species from the northern Gulf of Mexico are questionable (Boschung 1992). It is associated with clear shallow water over rocky bottoms near coral reefs, and with sandy bottoms and sea-grass beds, and often forms large schools. Food consists of cephalopods, shrimps, crabs, and small ray-finned fishes. Feeding takes place at night. Spawning occurs in summer in the northeastern Caribbean Sea. Maximum known size is 480 mm SL. references: Anderson 1967, 2002c; Böhlke and Chaplin 1968; Randall 1968a, 1996; Allen 1985; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997.

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LUTJANIDAE 331

Lutjanus purpureus (Poey, 1866)

Lutjanus purpureus has a pointed anal fin and relatively large scales. Interorbital space is slightly convex. Maxilla extends to anterior margin of pupil. Jaws have a row of enlarged teeth, including four canines in upper jaw, two of which are about half of pupil length, and an inner band of villiform teeth. Vomerine tooth patch is V-shaped or crescentic, with a relatively short posteromedial extension. Preoperculum is serrated along posterior and ventral margins, with serrae coarser at angle and on ventral margin than on posterior margin. Gill rakers on first arch number 6 or 7 on upper limb and 13 or 14 on lower limb. Measurements are expressed as percent of SL: head length 33%– 40%, eye diameter 5%–9%, snout length 13%–14%, upper jaw length 12%–15%, pectoral fin length 28%–34%, body depth 34%– 40%. Pectoral fin has 17 rays. Dorsal fin is slightly notched between spinous and rayed sections and has 10 spines and 14 rays. Anal fin is pointed in specimens longer than 50 mm SL and has 8 or 9 (usually 8) rays. Caudal fin is emarginate to lunate, with upper lobe slightly longer than lower lobe. Scales in horizontal series number 49 to 52, and tubed lateral line scales number 49 to 53. Cheek scale rows number five to seven (usually six). Scales between dorsal fin origin and lateral line number 9 to

11, and scales between anal fin origin and lateral line number 16 to 19. Scale rows on back rise obliquely above lateral line, and scales on anterior section of body above lateral line are smaller than those on anterior section of body below lateral line. Scales cover proximal parts of membranes of rayed sections of dorsal and anal fins. Color is deep red on dorsal half of body and rosy with a silvery sheen on remainder of body. Iris of eye is red, and fins are red. Small dark spot is occasionally present on upper section of pectoral fin base. Young specimens (250 mm SL) have a blackish spot about size of eye on upper flank below anterior section of dorsal fin rays. This species occurs between 70 and 160 m over rocky bottoms in the western Atlantic from North Carolina and Yucatán to central Brazil, including the Greater and Lesser Antilles. It has not been recorded from the Gulf of Mexico, but due to its confusion with L. campechanus in the northern section of its range, it is included herein. Food consists of cephalopods, shrimps, crabs, and ray-finned fishes. Maximum known size is 100 cm TL.

references: Allen 1985; Robins and Ray 1986; Cervigón 1993b; Anderson 2002c.

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332 PERCIFORMES

Lutjanus synagris Lane snapper

(Linnaeus, 1758)

Lutjanus synagris has yellow stripes and a black spot on side. Interorbital space is moderately convex. Maxilla extends to anterior margin of orbit. Jaws have a row of enlarged teeth, including four small canine teeth in upper jaw, and a narrow band of villiform teeth. Vomerine tooth patch is V-shaped or crescentic, with a short posteromedial extension. Preoperculum has finely serrated posterior margin, with serrae enlarged at corner. Gill rakers on first arch number 6 or 7 on upper limb and 12 to 15 on lower limb. Measurements are expressed as percent of SL: head length 36%–38%, eye diameter 7%–9%, snout length 13%–14%, length of lower jaw 14%–15%, pectoral fin length 30%–32%, pelvic fin length 22%–24%, body depth 36%–38%. Pectoral fin has 15 or 16 rays. Dorsal fin is slightly notched between spinous and rayed sections and has 10 spines and 12 or 13 (usually 12) rays. Anal fin is rounded and has 8 or 9 (usually 8) rays, with middle rays less than half of head length. Caudal fin is emarginate. Scales in horizontal series number 60 to 65, and tubed lateral line scales number 47 to 50. Scale rows on back rise obliquely above lateral line. Scales cover proximal parts of membranes of rayed sections of dorsal and anal fins. Color is pink to reddish with green tinge, darker vertical bars on dorsal half of body, and silvery with yellowish tinge on remainder of body. Series of six to eight

yellow or golden stripes occur on flanks, and three or four similarly colored stripes occur on head. Black spot equal to eye diameter or larger is usually located on flank above lateral line under anterior dorsal fin rays. In specimens longer than 60 mm SL, less than onefourth of black spot is located below lateral line. Fins are reddish to yellowish. This species occurs between the shoreline and 400 m in the western Atlantic from North Carolina and Bermuda to southeastern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the coastal waters of the Gulf of Mexico and is associated with a wide variety of habitats, but especially coral reefs and vegetated sandy areas. Food consists of polychaetes, gastropods, cephalopods, shrimps, crabs, and small ray-finned fishes. Feeding takes place at night. Maximum known age is 10 years. Maximum known size is 710 mm TL.

references: Longley and Hildebrand 1941; Anderson 1967, 2002c; Böhlke and Chaplin 1968; Randall 1968a, 1996; Smith et al. 1975; Hastings et al. 1976; Hoese and Moore 1977, 1998; Allen 1985; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999.

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LUTJANIDAE 333

Lutjanus vivanus Silk snapper

(Cuvier, 1828)

Lutjanus vivanus is bright red and has yellow eyes. Interorbital space is slightly convex. Maxilla extends to anterior margin of pupil. Jaws have a row of enlarged teeth, including four canines in upper jaw, two of which are about half of pupil length, and an inner band of villiform teeth. Vomerine tooth patch is V-shaped or crescentic, with a posteromedial extension. Tongue has oval patch of small teeth. Preoperculum is serrated along posterior and ventral margins, with serrae coarser at angle and on ventral margin than on posterior margin. Gill rakers on first arch number 6 to 9 on upper limb and 16 or 17 on lower limb. Measurements are expressed as percent of SL: head length 33%– 38%, eye diameter 6%–10%, snout length 13%–14%, upper jaw length 13%–14%, pectoral fin length 31%–33%, pelvic fin length 22%–25%, body depth 34%–37%. Pectoral fin has 16 to 18 rays. Dorsal fin has a shallow notch between spinous and rayed sections and has 10 spines and 13 or 14 (usually 14) rays. Anal fin is pointed, with middle rays equal to or longer than half of head length in specimens longer than 60 mm SL, and has 7 or 8 (usually 8) rays. Caudal fin is lunate to forked. Scales in horizontal series number 50 to 53, and tubed lateral line scales number 47 to 50. Scale rows on back rise obliquely above lateral line. Cheek scales number 7 or 8. Scales from dorsal fin origin to lateral line number 10 to 12, and scales

from anal fin origin to lateral line number 20 to 24. Bases of rayed sections of dorsal fin and anal fin are covered with scales. Color is pink to red on dorsal half of body and pinkish with a silvery sheen on remainder of body. Fine undulating yellow lines occur on flank. Iris of eye is bright yellow. Fins are reddish or pale yellow, and posterior margin of caudal fin is often deep red or dusky. Young specimens (250 mm SL) have a blackish spot on upper flank below anterior section of dorsal fin rays. This species occurs in the western Atlantic from North Carolina and Bermuda to northern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is common throughout the Gulf of Mexico. Depths of capture generally range from 90 to 140 m, but it is occasionally captured at depths greater than 200 m. It is associated with sandy, gravelly, and coralline bottoms. Food consists of gastropods, cephalopods, shrimps, crabs, tunicates, and ray-finned fishes. Maximum known size is 840 mm TL.

references: Ginsburg 1930; Longley and Hildebrand 1941; Rivas 1966; Anderson 1967, 2002c; Böhlke and Chaplin 1968; Hoese and Moore 1977, 1998; Allen 1985; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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334 PERCIFORMES

Ocyurus chrysurus (Bloch, 1791) Yellowtail snapper

Ocyurus chrysurus has symmetrical dorsal and ventral profiles and a deeply forked caudal fin. Interorbital space is strongly convex. Maxilla extends to anterior margin of orbit. Jaws have an outer row of enlarged teeth, including four to six small canines in upper jaw, and an inner narrow band of villiform teeth. Vomerine tooth patch is V-shaped, with a narrow posteromedial extension. Preoperculum is weakly serrated and has a shallow depression along posterior margin. Gill rakers on first arch number 9 to 11 on upper limb and 21 to 23 on lower limb. Measurements are expressed as percent of SL: head length about 29%, eye diameter 6%–11%, snout length 11%–13%, upper jaw length 10%–16%, pectoral fin length 27%– 31%, pelvic fin length 19%–22%, body depth 30%–34%. Pectoral fin has 15 or 16 (rarely 17) rays. Dorsal fin is slightly notched between spinous and rayed sections and has 10 (rarely 9) spines and 12 or 13 (rarely 14) rays. Anal fin is broadly rounded and has 8 or 9 (usually 9) rays. Upper caudal fin lobe is longer than lower lobe, and both lobes are pointed. Tubed lateral line scales number 46 to 49. Scale rows on back rise obliquely above lateral line. Scales cover proximal parts of membranes of rayed

sections of dorsal and anal fins. Color is blue to violet, with a scattering of unequal yellow spots on dorsal half of body, and whitish with narrow reddish and yellow stripes on remainder of body. Midlateral yellow band runs from snout to caudal fin base. Dorsal and caudal fins are yellow, and anal and pelvic fins are whitish. This species occurs in the tropical and warm temperate Atlantic between the shoreline and 165 m. In the western Atlantic it occurs from Massachusetts and Bermuda to southern Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. Adults are associated with coral reefs, and juveniles are associated with seaweed beds. Food consists of zooplankton, nekton, and benthic organisms, including polychaetes, gastropods, cephalopods, and rayfinned fishes. Maximum known age is 17 years. Maximum known size is 810 mm TL.

references: Longley and Hildebrand 1941; Anderson 1967, 2002c; Böhlke and Chaplin 1968; Randall 1968a, 1996; Bright and Cashman 1974; Smith et al. 1975; Hoese and Moore 1977, 1998; Allen 1985; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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LUTJANIDAE 335

Pristipomoides aquilonaris Wenchman

(Goode and Bean, 1896)

Pristipomoides aquilonaris has very large eyes and a relatively flat interorbital region. Maxilla extends to about center of eye. Jaws have an outer row of enlarged teeth and an inner band of villiform teeth. Vomerine tooth patch is V-shaped or triangular, without a posteromedial extension. Tongue lacks teeth. Preoperculum has serrated posterior and ventral margins, with serrae increasing in size ventrally. Operculum has 2 flat spines. Gill rakers on first arch number 7 to 9 on upper limb and 16 to 20 on lower limb. Measurements are expressed as percent of SL: head length 36%–38%, eye diameter 11%–13%, snout length 9%–11%, upper jaw length 15%–16%, pectoral fin length 30%–33%, pelvic fin length 23%–24%, body depth 31%– 41%. Pectoral fin has 15 or 16 (rarely 14 or 17) rays. Dorsal fin is unnotched between spinous and rayed sections and has 10 spines and 11 (rarely 10) rays. Anal fin is broadly rounded and has 8 (rarely 7) rays. Last ray of dorsal and anal fins is distinctly

longer than preceding rays. Caudal fin is deeply forked. Tubed lateral line scales number 48 to 52. Scale rows on back run parallel to lateral line. Scales are lacking from maxilla and dorsal and anal fins. Color is pink to reddish dorsally and pink to silvery white ventrally. Fins are translucent to pinkish, with outer sections of dorsal and caudal fins yellow. This species occurs between 24 and 488 m in the western Atlantic from North Carolina to southern Brazil, including the northern and southern Gulf of Mexico and the Greater and Lesser Antilles. It is associated with rough bottoms, including natural and artificial reefs. Food consists mostly of small ray-finned fishes. Maximum known size is 240 mm SL. references: Ginsburg 1952 (as Pristipomoides andersoni); Anderson 1966, 1967, 1972, 2002c; Allen 1985; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997.

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336 PERCIFORMES

Pristipomoides macrophthalmus (Müller and Troschel, 1848)

Pristipomoides macrophthalmus has very large eyes and a relatively flat interorbital region. Maxilla extends to anterior margin of pupil. Jaws have an outer row of enlarged teeth and an inner row of vomerine teeth. Vomerine tooth patch is V -shaped or triangular, without a posteromedial extension. Teeth are absent from tongue. Preoperculum has serrated posterior and ventral margins. Operculum has 2 flat spines. Gill rakers on first arch number 6 to 8 on upper limb and 13 to 17 (usually 14 to 16) on lower limb. Measurements are expressed as percent of SL: head length 36%– 39%, eye diameter 11%–12%, snout length 10%–11%, upper jaw length 15%–16%, pectoral fin length 30%–32%, pelvic fin length 22%–24%, body depth 32%–39%. Pectoral fin is long and has 15 or 16 rays. Dorsal fin is unnotched between spinous and rayed sections

and has 10 spines and 11 rays. Anal fin has slightly convex margin and 8 rays. Last ray of dorsal and anal fins is produced. Caudal fin is deeply forked. Tubed lateral line scales number 54 to 57. Scales are absent from maxilla and dorsal and anal fins. Color is pink with a silvery sheen on dorsal half of body and silvery on remainder of body. Fins are translucent to pink. This species occurs between 110 and 550 m in the western Atlantic from the Straits of Florida and Bermuda to Panama, including the Gulf of Mexico, the Bahamas, and the Greater Antilles. In the Gulf of Mexico it occurs off the Dry Tortugas and Veracruz. Maximum known size is 370 mm SL. references: Longley and Hildebrand 1941; Anderson 1967, 1972, 2002c; Allen 1985; Schaldach et al. 1997; Smith-Vaniz et al. 1999.

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LUTJANIDAE 337

Rhomboplites aurorubens Vermilion snapper

(Cuvier, 1829)

Rhomboplites aurorubens has very large eyes, a slightly projecting lower jaw, and no enlarged teeth in jaws. Interorbital space is strongly convex. Maxilla extends to anterior margin of orbit. Jaws have an outer row of slightly enlarged teeth and an inner band of villiform teeth. Vomerine tooth patch is rhomboid shaped, with an elongate medial extension. Tongue has ovate patch of granular teeth. Gill rakers on first arch number 8 to 10 on upper limb and 19 to 22 on lower limb. Measurements are expressed as percent of SL: head length 35%–37%, eye diameter 9%–11%, snout length 9%–11%, upper jaw length 12%–13%, pectoral fin length 29%–30%, pelvic fin length 22%–23%, body depth 31%–37%. Pectoral fin has 16 to 19 (usually 17 or 18) rays. Dorsal fin is very shallowly notched between spinous and rayed sections and has 12 (vary rarely 13) spines and 11 (rarely 10 or 12) rays. Anal fin is rounded and has 3 spines and 8 (rarely 9) rays. Caudal fin is lunate to forked. Scales in horizontal series number 66 to 71, and tubed lateral line scales number 46 to 52. Scale rows on back rise obliquely above lateral line. Scales cover proximal parts of membranes of rayed sections of dorsal and anal fins. Color is vermilion

on dorsal half of body and silvery with a reddish tinge on remainder of body. Faint oblique brown lines, following scale rows, run along upper flanks, and narrow horizontal yellow lines run along lower flank. Iris of eye is red. Dorsal fin is yellowish with vermilion blotches, caudal fins are yellowish, and anal and pelvic fins are whitish. This species occurs between 25 and 400 m in the western Atlantic from North Carolina and Bermuda to southern Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. Juveniles are found at the shallowest depths. It is associated with rocky bottoms near the edge of continental and insular shelves. Food consists of pelagic and benthic organisms, including polychaetes, shrimps, crabs, and ray-finned fishes. Maximum known age is 14 years. Maximum known size is 630 mm TL.

references: Longley and Hildebrand 1941; Anderson 1967, 2002c; Böhlke and Chaplin 1968; Hastings et al. 1976; Hoese and Moore 1977, 1998; Grimes et al. 1982; Allen 1985; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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338

SYMPHYSANODONTIDAE

Symphysanodontids are slender to moderately deep and moderately compressed, with a continuous dorsal fin, a bony elevation on posterodorsal surface of lower jaw, and two spines on operculum. Dorsal profile of head is moderately convex. Snout is short and blunt, and eye is moderate to large. Nostril is paired, with nares close set and about equal in size. Mouth is moderately large and moderately oblique. Upper jaw is moderately protrusible, and premaxilla has a broad midlateral process. Upper jaw is notched on either side of symphysis and bears small teeth lateral to notch. Dorsal section of maxilla is covered by very narrow suborbital bone. Lower jaw has symphyseal knob that fits into notch of upper jaw and bears teeth on bony elevation, with those at symphysis relatively large. Teeth are absent in vomer and usually absent in palatine. Teeth are absent on tongue. Preoperculum is serrated or smooth along posterior margin. Operculum has two flat, posteriorly directed spines, with lower one larger than upper one. Branchiostegal rays number 7. Pseudobranch is present on inner surface of operculum. Gill membranes are free of isthmus, and last full arch (fourth) has slit posteriorly. Gill rakers on first arch are slender and range from 9 to 14 on upper limb and 20 to 29 on lower limb. Pectoral fin is well developed and narrowly pointed, with a nearly vertical base and 15 to 18 rays, and upper rays are considerably longer than lower rays. Dorsal fin is unnotched and consists of nine spines and 10 rays. Pelvic fin is well developed but smaller than pectoral fin, inserts below pectoral fin base, and has one spine and 5 rays. Anal fin is equal to or shorter than rayed section of dorsal fin and has three spines and 7 or 8 rays. Caudal fin is deeply forked and has 17 principal rays, 15 of which are branched. Scales are ctenoid and cover body and most of head, including maxilla and lower jaw. Rayed sections of dorsal and anal fins lack scales, but scales form sheath at base of both fins. Axillary process on base of pelvic fin is well developed. Lateral line is complete, is gently curved beneath dorsal fin, and has 42 to 61 tubed scales. Vertebrae number 25: 10 precaudal and 15 caudal. Maximum known size is about 170 mm TL. Symphysanodontids occur in the western Atlantic, western and Central Pacific, and western Indian Oceans between 50 and 500 m in tropical seas on continental and insular shelves and slopes. They are thought to be planktivorous. There are six species in a single genus, two species in the western Central Atlantic, and one or both of these in the Gulf of Mexico. references: Anderson 1970, 2001, 2002a.

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SYMPHYSANODONTIDAE 339

Key to the Species of the Gulf of Mexico (Adapted from Anderson 1970, 2002a) 1a. Body depth 33% to 36% of SL; anal fin rays eight (rarely seven); gill rakers on first arch 39 to 42; pelvic fin length 26% to 28% of SL; lateral line scales 45 or 46 . . . . . . . . . . . . . . . . . . Symphysanodon octoactinus p. 341 1b. Body depth 22% to 28% of SL; anal fin rays seven; gill rakers on first arch 34 to 39; pelvic fin length 20% to greater than 87% of SL; lateral line scales 48 to 52 . . . . . . . . . . . . . . . . . . . . . . Symphysanodon berryi p. 340

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340 PERCIFORMES

Symphysanodon berryi

Anderson, 1970

Symphysanodon berryi is relatively slender and has strong serrations on posterior margin of preoperculum. Tip of snout extends slightly beyond upper jaw. Interorbital space is flat. Maxilla extends to about midlength of eye. Jaw teeth are small, conical, and arranged in narrow bands. Teeth are lacking in vomer and palatine. Gill rakers on first arch number 9 to 12 on upper limb and 24 to 28 on lower limb. Measurements are expressed as percent of SL: head length 31%–36%, snout length 6%–9%, eye diameter 10%–13%, upper jaw length 15%–17%, pectoral fin length 22%–25%, body depth 22%–28%. Pectoral fin has 16 to 18 rays. Dorsal fin has nine (rarely eight) spines and 10 (rarely 9 or 11) rays. First pelvic fin ray fails to reach vent in mature females but may extend beyond fork of caudal fin in mature male. Anal fin has 7 rays. Upper and lower caudal fin lobes are produced as filaments in mature males. Tubed lateral line scales number

48 to 52. Color in life is bright orange, with belly silvery orange. Iris of eye is mostly orange. In preservative, specimens are straw colored. This species occurs from 100 to 475 m in the western Atlantic from North Carolina, the Gulf of Mexico, the Bahamas, Cuba, the Dominican Republic, Puerto Rico, Tobago, and along the Caribbean coasts of Mexico, Belize, Honduras, Nicaragua, and Venezuela; and from the Central Atlantic off Ascension Island. There are two records of this species from the Gulf of Mexico, based on larvae collected off Louisiana (2652N, 8923W) and off Yucatán (2254N, 9136W). Most juvenile and adult specimens have been captured at depths greater than 200 m. Maximum known size is 150 mm SL.

references: Anderson 1970, 2002a; Cervigón 1993b.

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SYMPHYSANODONTIDAE 341

Symphysanodon octoactinus

Anderson, 1970

Symphysanodon octoactinus is relatively deep and has an entire or slightly serrated posterior preoperculum margin. Tip of snout extends slightly beyond upper jaw. Interorbital space is flat. Upper jaw extends to about midlength of eye. Jaw teeth are small, conical, and arranged in narrow bands. Teeth are lacking in vomer and are present or absent in palatine. Gill rakers on first arch number 12 to 14 on upper limb and 26 to 29 on lower limb. Measurements are expressed as percent of SL: head length 34%–36%, snout length 6%–9%, eye diameter 12%–15%, upper jaw length 14%–17%, pectoral fin length 28%–30%, body depth 33%–36%. Pectoral fin has 16 or 17 (usually

16) rays. Dorsal fin has 9 (rarely 10) spines and 10 (rarely 9) rays. Anal fin has 8 (occasionally 7) rays. Caudal fin lobes are not produced. Tubed lateral line scales number 45 or 46. In preservative, specimens are straw colored, with small dark spots on body. This species occurs between 155 and 404 m in the western Atlantic from the Bahamas, Cuba, Puerto Rico, Nicaragua, and Panama. It has not been captured in the Gulf of Mexico but may occur there. Maximum known size is 130 mm SL.

references: Anderson 1970, 2002a.

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342

LOBOTIDAE

Tripletails

Tripletails are oval to oblong in lateral profile, deep bodied, and compressed, with symmetrically shaped posterior dorsal and anal fin lobes and a rounded caudal fin. Dorsal profile behind head is strongly elevated and straight to slightly concave. Snout is short and moderately acute. Mouth is slightly oblique, with thick lips, lower jaw projecting beyond upper jaw, and upper jaw extending to center of eye. Nostril is paired, with nares round and close together. Eye is small and laterally located. Upper jaw is protrusible, and supramaxilla is absent. Jaws have an outer row of enlarged, conical, posteriorly directed teeth and an inner narrow band of villiform teeth. Teeth are absent in vomer and palatine. Preoperculum is angular and has strong serrae on posterior and ventral margins, with those at corner strongest and those on posterior margin curved dorsally. Operculum has two spines. Branchiostegal rays number 6. Pectoral fin is located on lower flank and is of moderate size. Dorsal fin is notched and consists of 11 to 13 stout spines and 13 to 16 rays. Pelvic fin is thoracic and well developed and consists of one spine and 5 rays. Anal fin has three spines and 8 to 11 rays. Moderate-sized cycloid scales with spines on posterior margin cover body and head except for preorbital region and jaws. Bases of dorsal and anal fins are covered with scales. Lateral line is complete and consists of 42 to 70 scales. Vertebrae number 24: 12 precaudal and 12 caudal. Gas bladder is present and lacks posterior extension. Maximum known size is 110 cm TL. Tripletails occur worldwide in tropical to warm temperate seas. They are benthic and associated with coastal waters and estuaries but occasionally are found offshore under flotsam. There are from five to eight species in two genera, and one species occurs in the western Central Atlantic and Gulf of Mexico. references: Jordan and Evermann 1896; Johnson 1984; Nelson 1994; Tominaga et al. 1996; Smith-Vaniz et al. 1999; Carpenter 2001b, 2002a.

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LOBOTIDAE 343

Lobotes surinamensis Tripletail

(Bloch, 1790)

Lobotes surinamensis has posterior lobes of dorsal and anal fins symmetrically expanded beyond caudal fin base and a relatively large third anal fin spine. Gill rakers on first arch number 5 to 7 on upper limb and 13 to 15 on lower limb. Measurements are expressed as percent of SL: head length 33%–39%, snout length about 8%, eye diameter 4%–7%, upper jaw length about 14%, body depth 48%– 54%. Pectoral fin has 16 rays. Dorsal fin consists of 11 or 12 stout spines and 15 or 16 rays. Anal fin has 11 or 12 rays. Lateral line scales number 43 to 48. Color is mottled tan to dark brown, with darker-colored blotches below dorsal fin, above anal fin, and on fins except for spinous section of dorsal fin. Caudal fin has pale margin in juveniles. This species occurs worldwide in tropical to warm temperate waters. In the western Atlan-

tic it occurs from Nova Scotia and Bermuda to northern Argentina, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. In the northern Gulf of Mexico it is present but uncommon in shallow water and bays during the summer months. Juveniles are associated with flotsam in open water and apparently mimic small flotsam. Adults are often observed floating on their sides at the surface. Maximum known size is 110 cm TL.

references: Hildebrand and Schroeder 1928; Baughman 1941, 1943; Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Hastings et al. 1976; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Carpenter 2002a.

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344

GERREIDAE

Mojarras

Mojarras are oblong or oval in lateral profile and moderately slender to deep bodied and compressed, with a very protrusible upper jaw, a continuous dorsal fin, and a forked caudal fin. Nostril is paired, and anterior and posterior nares are close set. Mouth is terminal, slightly oblique, and small. Supramaxilla is absent. Jaw teeth are villiform and in narrow bands. Vomer and palatine lack teeth. Preoperculum has entire to strongly serrated margin, and operculum lacks spine. Branchiostegal rays number 6. Gill membranes are free of isthmus. Gill rakers are short and broad. Pectoral fin inserts low on flank and is long and pointed. Dorsal fin is rather long and single but slightly to deeply incised between spinous and rayed sections, with 9 or occasionally 10 spines and 9 to 17 rays. Pelvic fin inserts below to somewhat posterior to pectoral fin base and has 1 spine and 5 rays. Scalelike axillary process is located behind base of pelvic fin. Anal fin is shorter than dorsal fin and has 2 or 3 or 5 or 6 spines and 6 to 18 rays. Caudal fin is forked to deeply forked. Body and much of the head are covered with cycloid or weakly ctenoid scales. Bases of dorsal and anal fins are bordered by scaly sheath. Gas bladder is present and often extends to base of anal fin spines. Vertebrae number 24: 10 precaudal and 14 caudal. Maximum known size is 35 cm TL. Mojarras occur worldwide in tropical to warm temperate seas to a depth of 70 m. Most species occur in coastal waters, but one species is confined to freshwater, several others occasionally enter freshwater, and many occur in brackish waters and estuaries. They are associated with sandy to muddy bottoms and occasionally with coral reefs. Most species form large schools. Food consists of small benthic invertebrates. There are 40 species in seven or eight genera, and 12 species in four genera occur in the western Central Atlantic and the Gulf of Mexico. references: Jordan and Evermann 1898; Johnson 1984; Woodland 1986, 2001; Nelson 1994; Tominaga et al. 1996.

Key to the Species of the Gulf of Mexico (Adapted from R. E. Matheson, pers. comm., 1983; Deckert and Greenfield 1987) 1a. Preopercular margin strongly serrated; second dorsal fin spine longer than distance between tip of snout and posterior margin of orbit; maxilla extending to or beyond anterior margin of pupil; body depth 42% to 59% SL . . . . . . 2 1b. Preopercular margin entire; second dorsal fin spine equal to or shorter than distance between tip of snout and poste-

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GERREIDAE 345

2a.

2b.

3a.

3b.

4a.

4b.

5a.

5b.

6a. 6b. 7a.

7b.

8a.

rior margin of orbit; maxilla usually not extending to anterior margin of pupil; body depth 30% to 44% SL . . . . . . 5 Lachrymal bone serrated (except entire in juveniles); body with bold black stripes; second anal fin spine longer than anal fin base; pharyngeal teeth molarlike . . . . . . . . . . . . . 3 Lachrymal bone entire; body without black stripes; second anal fin spine shorter than anal fin base; pharyngeal teeth pointed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Anal fin with three spines and seven rays (juveniles may have two spines and eight rays); gill rakers on lower limb of first arch 10 to 12 (usually 11 or 12) . . . . . . . . . . . . . . . . . . . . . . . . Eugerres brasilianus p. 356 Anal fin with three spines and eight rays (juveniles may have two spines and nine rays); gill rakers on lower limb of first arch 13 to 17 (usually 14 to 16) . . . . . . . . . . . . . . . . . . . . . . . . . . Eugerres plumieri p. 357 Gill rakers on lower limb of first arch 12 to 15 (usually 12 or 13); anal fin with three spines and eight rays (juveniles may have two spines and nine rays) . . . . . . . . . . . . . . . . . . . . . . . . . . Diapterus auratus p. 347 Gill rakers on lower limb of first arch 16 to 18 (usually 17); anal fin with two spines and nine rays . . . . . . . . . . . . . . . . . . . . . . . . Diapterus rhombeus p. 348 Body with up to seven or eight obscure, dark bars (specimens 50 mm SL), or body with six square, black spots (specimens 50 mm SL); gas bladder posteriorly bifurcated at level of first interhaemal bone, and passing for some distance on either side of bone; anterior interhaemal bone spine shaped . . . . . . . . . . . . . . Gerres cinereus p. 358 Body without dark bars (may have diagonal bars or mottling; specimens 50 mm SL); gas bladder not bifurcated at level of first interhaemal bone; anterior interhaemal bone spoon or cone shaped . . . . . . . . . . . . . . . . . . . . . . . 6 Anterior interhaemal bone spoon shaped; either with pectoral fins densely scaled or two anal fin spines . . . . . . . . . 7 Anterior interhaemal bone cone shaped; pectoral fins unscaled and anal fin with three spines. . . . . . . . . . . . . . . 8 Pectoral fins densely scaled; anal fin with three spines (for small specimens that have not developed scales on pectoral fin, second anal fin spine is greater than 15% of SL, and third anal fin element is jointed but unbranched) . . . . . . . . . . . . . . . . . . . . . . . Eucinostomus havana p. 352 Pectoral fins unscaled; anal fin with two spines (in small specimens, second anal fin spine is less than 15% of SL, and third anal fin element is jointed and branched) . . . . . . . . . . . . . . . . . . . . . . . . Eucinostomus lefroyi p. 354 Gill rakers on lower limb of first gill arch 9; tip of spinous dorsal fin with solid jet black blotch, which is separated

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346 PERCIFORMES

from remainder of fin by clear (white in fresh specimens) longitudinal band . . . . Eucinostomus melanopterus p. 355 8b. Gill rakers on lower limb of first gill arch 8; tip of spinous dorsal fin without jet black blotch above clear (white in fresh specimens) longitudinal band. . . . . . . . . . . . . . . . . . 9 9a. Premaxillary groove generally constricted or covered by scales in specimens between 40 and 60 mm SL, and closed by one or more scale rows in specimens greater than 60 mm SL; anal fin base 16% to 19% of SL. . . . . . . . . . 10 9b. Premaxillary groove usually unrestricted to slightly restricted by scales; anal fin base 13% to 16% of SL . . . 11 10a. Body depth 38% to 43% of SL in specimens 40 to 60 mm SL, and 40% to 46% of SL in specimens greater than 60 mm SL; pelvic fin length 21% to 26% of SL; last dorsal fin spine 7% to 10% of SL; light band crossing snout posterior to nostrils and V-shaped area between nostrils relatively indistinct in specimens greater than 80 mm SL . . . . . . . . . . . . . . . . . . . . . . . . . Eucinostomus gula p. 350 10b. Body depth 33% to 38% of SL in specimens 40 to 60 mm SL, and 35% to 40% of SL in specimens greater than 60 mm SL; pelvic fin length 18% to 22% of SL; last dorsal fin spine 6% to 8% of SL; light band crossing snout posterior to nostrils and V-shaped area between nostrils relatively distinct in specimens greater than 80 mm SL . . . . . . . . . . . . . . . . . . . . . Eucinostomus argenteus p. 349 11a. Lateral line scales 44 or 45 (modally 44, and rarely 43 or 46); body depth 33% to 37% of SL in specimens between 40 and 69 mm SL, and 34% to 38% of SL in specimens greater than 60 mm SL; least depth of caudal peduncle 10% to 13% of SL; pelvic fin length 18% to 26% of SL; snout pigment relatively uniform at all sizes, with indistinct band crossing snout posterior to nostrils and V-shaped area between nostrils . . . . . . . Eucinostomus harengulus p. 351 11b. Lateral line scales 46 to 48 (modally 47); body depth 29% to 33% of SL in specimens 40 to 60 mm SL, and 32% to 35% of SL in specimens greater than 60 mm SL; least depth of caudal peduncle 9% to 11% of SL; pelvic fin length 17% to 19% of SL; relatively distinct band across snout behind nostrils and V-shaped area between nostrils . . . . . . . . . . . . . . . . . . . . . . . . Eucinostomus jonesii p. 353

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GERREIDAE 347

Diapterus auratus Ranzani, 1842 Irish pompano

Diapterus auratus is rather deep bodied, with a strongly serrated preoperculum and a deeply notched dorsal fin. Snout is acute and beaklike. Maxilla extends to about midlength of eye. Lachrymal bone lacks serrations. Gill rakers on first arch are very short and number 12 to 15 (usually 12 or 13) on lower limb. Measurements are expressed as percent of SL: head length 32%–35%, snout length 8%–10%, eye diameter 10%–11%, last dorsal fin spine length 8%–10%, pelvic fin length 23%– 24%, second anal fin spine length 24%–27%, body depth 44%–51%, caudal peduncle depth 12%–13%. Pectoral fin has 16 rays. Dorsal fin has nine spines and 10 rays, with second spine as long as head. Anal fin has three spines and 8 rays, with second spine very stout. Juveniles may have two spines and 9 rays in anal fin. Body is covered with large cycloid scales, and top of head is covered with scales anteriorly to center of eye. Pored lateral line scales number 35 to 39 (usually 36 or 37). Color is silvery and slightly darker dorsally than ventrally, and oc-

casionally with four to six faint bars on side. Spinous section of dorsal fin has black edge, pelvic and anal fins are yellowish, and other fins are transparent. This species occurs in coastal waters in the western Atlantic from the east coast of Florida and the Bahamas to Brazil, including the Gulf of Mexico and the Antilles. In the Gulf of Mexico it is absent from north of Tampa Bay to the Mississippi River (R. E. Matheson, pers. comm., August 2002). Both adults and juveniles are associated with brackish and saltwater lagoons and protected bays. Food consists of benthic invertebrates. Maximum known size is 340 mm TL.

references: Hoese and Moore 1977 (as Diapterus olisthostomus), 1998; Robins and Ray 1986; Deckert and Greenfield 1987; Cervigón 1993a,b; Schaldach et al. 1997; Vega-Cendejas et al. 1997 (as D. olisthostomus); Castro-Aguirre et al. 1999; Gilmore and Greenfield 2002.

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348 PERCIFORMES

Diapterus rhombeus

(Cuvier, 1829)

Diapterus rhombeus is rather deep bodied, with a strongly serrated preoperculum, and a deeply notched dorsal fin. Snout is acute and beaklike. Maxilla extends to about middle of pupil. Lachrymal bone lacks serrations. Gill rakers on first arch are very short and number 16 to 18 on lower limb. Measurements are expressed as percent of SL: head length 33%– 37%, snout length 8%–10%, eye diameter 8%–12%, last dorsal fin spine length 12%– 14%, pelvic fin length 26%–29%, second anal fin spine length 25%–29%, body depth 47%– 54%, caudal peduncle depth 12%–13%. Pectoral fin has 16 rays. Dorsal fin has nine spines and 10 rays, with second spine as long as head. Anal fin has two spines and 9 rays, with second spine very stout. Body is covered with large cycloid scales, and head is covered with scales anteriorly to center of eye. Pored lateral line scales number 34 to 38 (usually 36 or 37).

Color is silvery and slightly darker dorsally than ventrally. Spinous section of dorsal fin has black edge, pelvic and anal fins are yellowish, and other fins are transparent. This species occurs in coastal waters in the western Atlantic from southeastern Florida, the southern Gulf of Mexico, and the Greater and Lesser Antilles to Bahia, Brazil. In the Gulf of Mexico it has been recorded from as far north as Laguna Madre de Tamaulipas. It is associated with mangrove-lined lagoons and sandyand muddy-bottomed regions in coastal areas. Food consists of benthic invertebrates. Maximum known size is 400 mm TL.

references: Deckert and Greenfield 1987; Cervigón 1993b; Schaldach et al. 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999; Gilmore and Greenfield 2002.

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GERREIDAE 349

Eucinostomus argenteus Spotfin mojarra

Baird and Girard, 1855

Eucinostomus argenteus is moderately slender, with maxillary groove constricted or crossed by scales, a V- or wedge-shaped unpigmented area between nostrils, and a light or unpigmented band crossing snout posterior to nostrils. Dorsal profile of head is slightly convex to nearly straight. Maxilla extends to near anterior margin of orbit. Margins of lachrymal and preopercular bones are entire. Opercular margin has a distinct notch above fleshy flap. Gill rakers on first arch are short and number eight (rarely nine) on lower limb. Measurements are expressed as percent of SL: head length 31%– 33%, upper jaw length 10%–11%, eye diameter 9%–11%, last dorsal fin spine length 6%– 8%, depressed anal fin length 23%–27%, second anal fin spine length 7%–14%, body depth 33%–38%, caudal peduncle depth 10%–12%. Pectoral fin has 14 to 16 rays. Dorsal fin is deeply notched, with nine spines and 10 rays. Anal fin has three spines and 7 rays, with third spine segmented but not branched in small

specimens. Lateral line scales number 44 to 46. Color is straw yellow to silver in preservative and silver in life. Juveniles have diagonal bars and mottling on upper sides, and these markings may be retained to some degree in adults. This species occurs along the continental shelf in the western Atlantic from New Jersey and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, Cuba, and the eastern Lesser Antilles. It occurs throughout the Gulf of Mexico. It is apparently most common at depths of 20 to 60 m. Maximum known size is 147 mm SL.

references: Böhlke and Chaplin 1968 (as E. gula, in part); Randall 1968a, 1996; Hoese and Moore 1977 (in part), 1998; Hastings 1979; Matheson 1983; Matheson and McEachran 1984; Boschung 1992; Schaldach et al. 1997; Smith 1997; VegaCendejas et al. 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Gilmore and Greenfield 2002.

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350 PERCIFORMES

Eucinostomus gula Silver jenny

(Quoy and Gaimard, 1824)

Eucinostomus gula is relatively deep bodied, with premaxillary groove constricted or crossed by scales, a moderately pigmented V-shaped pattern between nostrils, and a light band posterior to nostrils. Dorsal profile of head is slightly convex to nearly straight. Maxilla extends to near anterior margin of orbit. Margins of lachrymal and preopercular bones are entire. Opercular margin has a distinct notch above fleshy flap. Gill rakers on first arch are short and number eight (rarely nine) on lower limb. Measurements are expressed as percent of SL: head length 29%–35%, upper jaw length 9%–11%, eye diameter 8%–12%, last dorsal fin spine length 7%–10%, depressed anal fin length 24%–28%, second anal fin spine length 7%–14%, body depth 38%– 46%, caudal peduncle depth 11%–13%. Pectoral fin has 14 to 16 rays. Dorsal fin is deeply notched and has nine spines and 10 rays. Anal fin has three spines and 7 rays, with third spine segmented but not branched in small specimens. Lateral line scales number 42 to 44 (rarely 41 or 45).

Color is straw yellow to silver in preservative and silver in life. Juveniles have diagonal bars and mottling on upper sides that continue as blotches on lower flanks. Markings disappear in specimens greater than 70 mm SL. This species occurs in coastal waters in the western Atlantic from New Jersey and Bermuda to Argentina, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It occurs throughout the Gulf of Mexico. This species is often associated with grass beds. Juveniles may spend cooler months of the year in offshore waters between 9 and 24 m. Maximum known size is 147 mm SL.

references: Böhlke and Chaplin 1968 (as E. gula, in part); Hoese and Moore 1977 (in part), 1998; Matheson 1983; Matheson and McEachran 1984; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Vega-Cendejas et al. 1997; CastroAguirre et al. 1999; Smith-Vaniz et al. 1999; Gilmore and Greenfield 2002.

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GERREIDAE 351

Eucinostomus harengulus Tidewater mojarra

Goode and Bean, 1879

Eucinostomus harengulus is relatively slender, with premaxillary groove unconstricted by scales and a faint V-shaped pattern between nostrils but without a light band posterior to nostrils. Dorsal profile of head is slightly convex to nearly straight. Maxilla extends to near anterior margin of orbit. Margins of lachrymal and preopercular bones are entire. Opercular margin has a distinct notch above fleshy flap. Gill rakers on first arch are short and number eight (rarely nine) on lower limb. Measurements are expressed as percent of SL: head length 29%–36%, upper jaw length 9%–12%, eye diameter 8%–12%, last dorsal fin spine length 5%–9%, depressed anal fin length 21%–25%, second anal fin spine length 7%– 14%, body depth 33%–38%, caudal peduncle depth 10%–13%. Pectoral fin has 14 to 16 rays. Dorsal fin is deeply notched and has nine spines and 10 rays. Anal fin has three spines and 7 rays, with third spine segmented but not branched in small specimens. Lateral line scales number 44 or 45 (rarely 43 or 46). Color is straw yellow to silver in preservative and silver in life. Juveniles generally have diagonal bars and mottling on upper sides, with

anteriormost bar crossing nape above operculum, second bar forming a Y under spinous dorsal fin, third bar slanted posteriorly and located under anterior portion of rayed section of dorsal fin, fourth bar located at rear of rayed section of dorsal fin, and fifth bar at rear of caudal peduncle. This species occurs in coastal waters in the western Atlantic from New Jersey and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the northwestern Gulf of Mexico it has not been recorded from south of Laguna Madre, but it has been recorded from the west coast of Campeche. This species is often associated with protected inshore areas, including grass beds. It has been captured to a depth of 36 m but is most common in shallow water. Maximum known size is 135 mm SL. references: Böhlke and Chaplin 1968 (as E. argenteus, in part); Matheson 1983; Matheson and McEachran 1984; Boschung 1992; CastroAguirre et al. 1999; Smith-Vaniz et al. 1999; Gilmore and Greenfield 2002.

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352 PERCIFORMES

Eucinostomus havana Bigeye mojarra

(Nichols, 1912)

Eucinostomus havana is relatively deep bodied, with premaxillary groove unconstricted by scales, pectoral fins densely covered with scales, and a black blotch at tip of spinous section of dorsal fin. Dorsal profile of head is slightly convex to nearly straight. Maxilla extends to near anterior margin of orbit. Margins of lachrymal and preopercular bones are entire. Opercular margin has a distinct notch above fleshy flap. Eye length is about equal to snout length. Gill rakers on first arch are short and number four on upper limb and seven on lower limb. Measurements are expressed as percent of SL: head length 30%–32%, upper jaw length 9%–10%, eye diameter 11%–12%, last dorsal fin spine length 9%–10%, depressed anal fin length 21%–24%, second anal fin spine length 13%–16%, body depth 30%–37%, caudal peduncle depth 10%–11%. Pectoral fin has 15 or 16 rays. Dorsal fin is deeply notched and has nine spines and 10 rays. Anal fin has three

slender spines and 7 rays, with third spine segmented but not branched in small specimens. Lateral line scales number 46 or 48. Color is straw yellow to silver in preservative and slightly darker dorsally than ventrally. This species occurs in coastal waters in the western Atlantic from Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs at the Dry Tortugas and the Laguna de Términos, Veracruz. It is associated with mangrove-lined tidal creeks, tide pools, and open sandy beaches and has been captured to a depth of 3 m. Maximum known size is 180 mm TL. references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Robins and Ray 1986; Cervigón 1993a; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Gilmore and Greenfield 2002.

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GERREIDAE 353

Eucinostomus jonesii Slender mojarra

(Günther, 1879)

Eucinostomus jonesii is relatively slender, with premaxillary groove on snout unconstricted by scales, a light V-shaped pattern between nostrils, and a light band posterior to nostrils. Dorsal profile of head is slightly convex to nearly straight. Maxilla extends to near anterior margin of orbit. Margins of lachrymal and preopercular bones are entire. Opercular margin has a distinct notch above fleshy flap. Gill rakers on first arch are short and number eight (rarely nine) on lower limb. Measurements are expressed as percent of SL: head length 31%– 34%, upper jaw length 10%–12%, eye diameter 9%–12%, last dorsal fin spine length 4%– 7%, depressed anal fin length 21%–24%, second anal fin spine length 7%–14%, body depth 29%–35%, caudal peduncle depth 9%–11%. Pectoral fin has 14 to 16 rays. Dorsal fin is deeply notched and has nine spines and 10 rays. Anal fin has three spines and 7 rays, with third spine segmented but not branched in small specimens. Lateral line scales number 46 to 48. Color is straw yellow to silver in preservative and silver in life. Juveniles generally have diagonal bars and dark mottling on upper side, with anteriormost bar crossing nape above

operculum, second forming a Y under spinous dorsal fin, third slanting posteriorly and located under anterior portion of rayed section of dorsal fin, fourth located at rear of rayed section of dorsal fin, and fifth at rear of caudal peduncle. Bars are broader dorsally than ventrally. This species occurs in coastal waters in the western Atlantic from North Carolina and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it has been recorded from southwestern Florida, south Texas, Tamaulipas, Veracruz, and Campeche. It is associated with clear to slightly turbid water in both protected bays and open beaches over a variety of substrates. Adults are not common along the shore and thus are thought to spawn in deep water near the coast. Maximum known size is 159 mm SL. references: Böhlke and Chaplin 1968; Matheson 1983; Matheson and McEachran 1984; CastroAguirre et al. 1999; Smith-Vaniz et al. 1999; Gilmore and Greenfield 2002.

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354 PERCIFORMES

Eucinostomus lefroyi Mottled mojarra

(Goode, 1874)

Eucinostomus lefroyi is relatively slender, with premaxillary groove on snout unconstricted by scales, and two anal fin spines. Dorsal profile of head is slightly convex to nearly straight. Maxilla extends to near anterior margin of orbit. Margins of lachrymal and preopercular bones are entire. Opercular margin has a distinct notch above fleshy flap. Eye length is about equal to snout length. Gill rakers on first arch are short and number seven or eight on lower limb. Measurements are expressed as percent of SL: head length 32%–33%, upper jaw length 10%–11%, eye diameter 11%– 12%, last dorsal fin spine length 5%– 6%, depressed anal fin length 23%–24%, second anal fin spine length 8%–10%, body depth 29%– 30%, caudal peduncle 9%–10%. Pectoral fin has 16 rays. Dorsal fin is deeply notched and has nine spines and 10 rays. Anal fin has

8 rays. Lateral line scales number 44 to 46. Color is straw yellow to silver in preservative, with irregular brown diagonal marks above lateral line. This species occurs in coastal waters in the western Atlantic from Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is limited to the southern Gulf of Mexico. Adults are associated with sandy bottoms. Maximum known size is 230 mm TL.

references: Böhlke and Chaplin 1968 (as Ulaema lefroyi); Randall 1968a, 1996; Hoese and Moore 1977 (as U. lefroyi), 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997 (as U. lefroyi); Castro-Aguirre et al. 1999 (as U. lefroyi); Smith-Vaniz et al. 1999; Gilmore and Greenfield 2002.

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GERREIDAE 355

Eucinostomus melanopterus Flagfin mojarra

(Bleeker, 1863)

Eucinostomus melanopterus is moderately deep, with premaxillary groove unconstricted by scales and spinous section of dorsal fin dusky at base, white or clear above, and black tipped. Dorsal profile of head is slightly convex to nearly straight but slightly concave just anterior to eye. Maxilla extends to near anterior margin of orbit. Margins of lachrymal and preopercular bones are entire. Opercular margin has a distinct notch above fleshy flap. Gill rakers on first arch are short and number nine on lower limb. Measurements are expressed as percent of SL: head length 30%–34%, upper jaw length 10%–11%, eye diameter 8%– 10%, last dorsal fin spine length 7%– 8%, depressed anal fin length 24%–26%, second anal fin spine length 10%–13%, body depth 34%– 35%, caudal peduncle depth 10%–11%. Pectoral fin has 14 to 16 rays. Dorsal fin is deeply notched and has nine spines and 10 rays.

Anal fin has three spines and 7 rays, with third spine segmented but not branched in small specimens. Lateral line scales number 44 to 47. Color is straw yellow to silver in preservative and silver in life. Dorsal surface is slightly greenish to brownish. This species occurs in the tropical to warm temperate Atlantic. In the western Atlantic it occurs in coastal waters from Florida and Bermuda to Brazil, including the Gulf of Mexico and the Greater and Lesser Antilles. It is limited to the western and southern Gulf of Mexico. This species is often associated with lagoons and bays and occasionally with freshwater. Maximum known size is 180 mm TL. references: Matheson 1981; Robins and Ray 1986; Cervigón 1993b; Schaldach et al. 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999; Gilmore and Greenfield 2002.

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356 PERCIFORMES

Eugerres brasilianus

(Cuvier, 1830)

Eugerres brasilianus is rather deep, with preoperculum strongly serrated, lachrymal bone serrated, and dorsal fin deeply notched. Maxilla extends slightly beyond anterior margin of eye. Gill rakers on first arch are very short and number 10 to 12 (usually 11 or 12) on lower limb. Measurements are expressed as percent of SL: head length 29%– 42%, second anal fin spine length 17%–29%, body depth 37%–53%. Pectoral fin has 16 rays. Dorsal fin has nine spines and 10 rays, with second spine shorter than head length. Anal fin has three spines and 8 rays, with second spine very stout. Juveniles may have two spines and 9 rays in anal fin. Body is covered with large cycloid scales, and head is covered with scales anteriorly to near anterior margin of eye. Pored lateral line scales number 34 to 39 (usually 36 or 37). Color is silvery, with moderately distinct

dark stripes along scale rows. All fins except pectoral fin are dusky. Pectoral fin is transparent. In preservative, upper half of body is pale yellowish brown, and lower half of body is pale yellowish silver. Stripes are brown. This species occurs in the western Atlantic from the southern Gulf of Mexico to southern Brazil, including the Antilles. In the Gulf of Mexico it is known from Tamaulipas and Veracruz. There is also a single unconfirmed report from southern Texas (Baughman 1950). It is associated with inshore waters, including estuaries, bays, and lagoons. Maximum known size is 272 mm SL. references: Baughman 1950; Deckert and Greenfield 1987; Greenfield and Thomerson 1997; Castro-Aguirre et al. 1999; Gilmore and Greenfield 2002.

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GERREIDAE 357

Eugerres plumieri Striped mojarra

(Cuvier, 1830)

Eugerres plumieri is rather deep bodied, with lachrymal bone and preoperculum strongly serrated and dorsal fin deeply notched. Maxilla extends slightly beyond anterior margin of eye. Gill rakers on first arch are very short and number 13 to 17 (usually 15) on lower limb. Measurements are expressed as percent of SL: head length 29%–39%, snout length 8%–11%, eye diameter 8%–12%, last dorsal fin spine length 12%–14%, pelvic fin length 26%–30%, second anal fin spine length 21%– 32%, body depth 44%–53%, caudal peduncle depth 13%–14%. Pectoral fin has 16 or 17 rays. Dorsal fin has nine spines and 10 rays, with second spine as long as head. Anal fin has three spines and 8 rays, with second spine very stout. Juveniles may have two spines and 9 rays in anal fin. Body is covered with large cycloid scales, and head is covered with scales anteriorly to near anterior margin of eye. Pored lateral line scales number 32 to 38 (usually 34 to 36). Color is silvery, with greenish blue tinges

on back and distinct dark brown longitudinal stripes on sides, produced by spots on center of scales. All fins except pectoral fin are dusky, and pelvic and anal fins are occasionally dark orange. Pectoral fin is transparent. This species occurs in coastal waters in the western Atlantic from South Carolina to Brazil, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs along the west Florida coast north to about the middle of the peninsula and along the coasts of Tamaulipas, Veracruz, and Campeche. It is associated with mangrove-lined lagoons and muddy-bottomed coastal areas and enters brackish and freshwater. Maximum known size is 400 mm TL. references: Robins and Ray 1986 (as Diapterus plumieri); Cervigón 1993a,b; Schaldach et al. 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999; Gilmore and Greenfield 2002.

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358 PERCIFORMES

Gerres cinereus (Walbaum, 1792) Yellowfin mojarra

Gerres cinereus is rather deep bodied, with gas bladder bifurcating and passing on either side of first interhaemal radial and seven or eight dark bars on side. Snout is acute and beaklike. Maxilla extends to about midlength of eye. Gill rakers on first arch are very short and number five or six on upper limb and seven or eight on lower limb. Measurements are expressed as percent of SL: head length 33%–35%, upper jaw length 12%–13%, eye diameter 10%– 13%, last dorsal fin spine length 9%–10%, depressed anal fin length 22%–25%, second anal fin spine length 13%–18%, body depth 38%– 42%, caudal peduncle depth 11%–12%. Pectoral fin has 15 or 16 rays. Dorsal fin has nine spines and 10 rays. Anal fin has three spines and 7 rays, with second spine very stout. Pored lateral line scales number 42 to 46. Color is pale tan and slightly darker dorsally than ventrally, with seven or eight dark brown bars on sides. Pelvic fin is yellow.

This species occurs in the western Atlantic and eastern Pacific Oceans. In the western Atlantic it occurs in coastal waters from Florida and Bermuda to southeastern Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. It is rare in the northwestern Gulf of Mexico but common along the west coast of Florida and along the coasts of Tamaulipas, Veracruz, Tabasco, and Campeche. It is associated with open sandy beaches, sea-grass beds, mangrove habitats, and coral reefs. Food consists of benthic invertebrates. Maximum known size is 410 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Vega-Cendejas et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Gilmore and Greenfield 2002.

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359

HAEMULIDAE Grunts

Grunts are relatively oblong in profile, deep bodied, and compressed, with a long, continuous dorsal fin; maxilla covered by lachrymal bone when mouth is closed; enlarged chin pores; and scales on most of head. Dorsal profile of head is nearly straight to strongly convex and moderately to steeply elevated. Snout is moderately short to long. Nostril is paired, with posterior naris elongate and slightly larger than anterior naris. Mouth is small to moderately large, moderately oblique, and terminal to subterminal. Upper jaw is variably protrusible, and premaxilla has variably developed midlateral process. Supramaxilla is absent. Cardiform teeth occur in jaws (except for a single genus with molariform teeth), with those of outer row enlarged. Vomer and palatine typically lack teeth. Preoperculum is generally serrated along posterior and ventral margins. Operculum has 1 distinct spine. Branchiostegal rays number 7. Gill membranes are free of each other but attached to isthmus, and last full arch (fourth) has medial slit. Pectoral fin is well developed and narrowly pointed, with a nearly vertical base and upper rays considerably longer than lower rays. Dorsal fin is unnotched to slightly notched between spinous and rayed sections and consists of 9 to 15 strong spines and 12 to 26 rays. Pelvic fin is well developed but smaller than pectoral fin, inserts below pectoral fin base, and has 1 spine and 5 rays. Anal fin is equal to or shorter than rayed section of dorsal fin and has 3 spines and 6 to 13 rays. Caudal fin is truncate to deeply forked and consists of 17 principal rays and 18 to 27 procurrent rays. Scales are ctenoid and cover body, lachrymal bone, snout, cheek, preoperculum, and operculum. Axillary process on base of pelvic fin is well developed. Vertebrae number 26 or 27: 10 or 11 precaudal and 16 caudal. Gas bladder is present and is usually contained in body cavity. Maximum known length is about 120 cm TL. Grunts occur worldwide in tropical to warm temperate seas in coastal waters, brackish waters, and occasionally fresh waters. Many species are associated with coral reefs and form dense schools. Feeding generally takes place at dawn and dusk. Food consists of benthic invertebrates and small ray-finned fishes. The common name “grunt” refers to the sound these fishes produce by grinding their pharyngeal teeth and that is amplified by their gas bladders. There are about 150 species in approximately 17 genera, 23 species in 6 genera occur in the western Central Atlantic, and 18 species in 5 genera occur in the Gulf of Mexico. references: Johnson 1980, 1984; Nelson 1994; Tominaga et al. 1996; Smith 1997; Smith Vaniz et al. 1999; McKay 2001; Lindeman and Toxey 2002.

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360 PERCIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Courtenay 1961; Lindeman 1986) 1a. Preoperculum strongly serrated, with two enlarged spines at angle and serrae on ventral margin directed anteriorly; sides of body with eight broad vertical bands . . . . . . . . . . . . . . . . . . . . . . . . . . . Conodon nobilis p. 366 1b. Preoperculum finely serrated, with none of serrae directed anteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Rayed sections of dorsal and anal fins densely covered with scales to margins; mouth red in fresh specimens (except in Haemulon boschmae). . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2b. Rayed sections of dorsal and anal fins naked or with scales only along bases; mouth not red in fresh specimens . . . . 27 3a. Specimens 50 mm SL or longer . . . . . . . . . . . . . . . . . . . . 4 3b. Specimens between 20 and 50 mm SL (other than Haemulon album) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 4a. Dorsal fin spines 13 or 14 . . . . . . . . . . . . . . . . . . . . . . . . 5 4b. Dorsal fin spines 12 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 5a. Scales encircling caudal peduncle 22; gill rakers on first arch 24 to 28; dorsal fin rays 14 or 15; anal fin rays usually 9. . . . . . . . . . . . . . . . . Haemulon aurolineatum p. 368 5b. Scales encircling caudal peduncle 24 or more (usually 26); gill rakers on first arch 27 to 36; dorsal fin rays 13 or 14; anal fin rays 7 or 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6a. Scale rows below lateral line oblique; body with five dusky yellow stripes, lower two broadest and lowest on lower half of body; no black spot on caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . . Haemulon striatum p. 379 6b. Scale rows below lateral line mostly horizontal; body with four brown to bronze stripes on upper half of body; large black spot on caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . Haemulon boschmae p. 370 7a. Scales encircling caudal peduncle 23 or more; scales between lateral line and anal fin origin 13 or 14. . . . . . . . 8 7b. Scales encircling caudal peduncle 22 or fewer; scales between lateral line and anal fin origin 12 or 13 (usually 12) .............................................9 8a. Back below dorsal fin, upper half of caudal peduncle, and caudal fin black; scales encircling caudal peduncle 23 to 25 (usually 23) . . . . . . . . . . . . . Haemulon melanurum p. 375 8b. Back below dorsal fin, upper half of caudal peduncle, and caudal fin without concentration of black pigment; scales encircling caudal peduncle 25 to 27 (usually 26) . . . . . . . . . . . . . . . . . . . . . . . . . . Haemulon album p. 367 9a. Dorsal fin rays 13 (rarely 12 or 14) . . . . . . . . . . . . . . . . . . . . Haemulon chrysargyreum p. 372

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HAEMULIDAE 361

9b. Dorsal fin rays 14 to 18 . . . . . . . . . . . . . . . . . . . . . . . . . 10 10a. Body with dark stripes, formed by dark spots on scales, running obliquely along sides of body . . . . . . . . . . . . . . 11 10b. Body without stripes, formed by dark spots on scales, running obliquely along sides of body . . . . . . . . . . . . . . . . . 12 11a. Pectoral fin not scaled; pored lateral line scales 45 to 48 (usually 46); dorsal fin rays 15 or 16 . . . . . . . . . . . . . . . . . . . . . . . Haemulon bonariense p. 369 11b. Pectoral fin scaled for more than one-third of length; pored lateral line scales 47 to 49; dorsal fin rays 16 to 18 (usually 17 or 18) . . . . . . . . . . . . . . . . . . . . Haemulon parra p. 376 12a. Scales below lateral line larger than those above lateral line and in oblique rows; gill rakers 21 to 24 (usually 23) . . . . . . . . . . . . . . . . . . . . . Haemulon flavolineatum p. 373 12b. Scales above and below lateral line equal in size, or those above lateral line larger . . . . . . . . . . . . . . . . . . . . . . . . . 13 13a. Scales above lateral line larger than those below . . . . . . . . . . . . . . . . . . . . . . . . Haemulon plumierii p. 377 13b. Scales above and below lateral line equal in size. . . . . . . 14 14a. Gill rakers on first arch 23 to 25; anal fin rays 8 . . . . . . . . . . . . . . . . . . . . . Haemulon carbonarium p. 371 14b. Gill rakers on first arch 26 to 31 (rarely 25); anal fin rays usually 9. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 15a. Pectoral fin rays 17 or 18 (usually 18); pored lateral line scales 50 to 52 (usually 51) . . . . . . . . . . . . . . . . . . . . Haemulon macrostomum p. 374 15b. Pectoral fin rays 16 (rarely 17); pored lateral line scales 48 to 51 . . . . . . . . . . . . . . . . . . . . . . Haemulon sciurus p. 378 16a. Dorsal fin spines 13 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16b. Dorsal fin spines 12 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 17a. Distinct spot on distal two-thirds of caudal fin base; ventral lateral scale rows parallel; third dorsal stripe extending to junction between dorsal fin or to origin of second dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 17b. No distinct spot on caudal fin base; ventral lateral scale rows oblique; third dorsal stripe extending to posterior margin of pectoral fin . . . . . . . . Haemulon striatum p. 379 18a. Anal fin rays 7 to 9 (usually 8); dorsal fin rays 11 to 15 (usually 13); third dorsal stripe extending to first dorsal fin ray. . . . . . . . . . . . . . . . . . . Haemulon boschmae p. 370 18b. Anal fin rays 7 to 9 (usually 9); dorsal fin rays 14 to 16 (usually 15); third dorsal stripe extending to posterior margin of operculum . . . . . . . . Haemulon aurolineatum p. 368 19a. Dorsal fin rays 12 to 14 (usually 13); anal fin rays 9 or 10 (usually 9). . . . . . . . . . . . Haemulon chrysargyreum p. 372 19b. Dorsal fin rays either fewer than or more than 12 to 14; anal fin rays either more than or fewer than 9 or 10. . . . 20

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20a. Midlateral stripe not continuous with caudal spot or, if continuous, stripe not darkened or expanded slightly at junction of spot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 20b. Midlateral stripe continuous with caudal spot and darkening and expanded slightly at junction of spot . . . . . . . . . . . . . . . . . . . . . . Haemulon melanurum p. 375 21a. Caudal fin spot centered over or located anterior to posterior three-fourths of caudal fin base . . . . . . . . . . . . . . . . 22 21b. Caudal fin spot located over posterior three-fourths of caudal fin base or located posterior to caudal fin base . . . . . 25 22a. Scales below anterior section of lateral line either equal in size or no larger than 2 times size of scales above anterior section of lateral line; ventral lateral scale rows oblique to parallel; no melanophores concentrated beneath posterior opercular margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 22b. Scales below anterior section of lateral line 2 or 3 times as large as scales above anterior section of lateral line; ventral lateral scale rows strongly oblique; melanophores strongly concentrated beneath posterior opercular margin . . . . . . . . . . . . . . . . . . . . . Haemulon flavolineatum p. 373 23a. Third dorsal stripe extending to fifth dorsal fin spine; greatest height of caudal spot at or posterior to caudal fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 23b. Third dorsal stripe not extending beyond fourth dorsal fin spine (usually extending to posterior opercular margin); greatest height of caudal spot usually anterior to end of caudal fin base . . . . . . . . . . . Haemulon bonariense p. 369 24a. Anal fin rays 7 to 9 (usually 8); dorsal fin rays 15 or 16 (usually 15); third dorsal stripe extending to fifth to tenth dorsal fin spine; posterior margin of caudal fin slightly rounded; ventral lateral scale rows more or less parallel . . . . . . . . . . . . . . . . . . . . . Haemulon carbonarium p. 371 24b. Anal fin rays 8 or 9 (usually 9); dorsal fin rays 15 to 17 (usually 16); third dorsal stripe extending to first dorsal fin ray; posterior margin of caudal fin truncate; ventral lateral scale rows more or less oblique . . . . . . . . . . . . . . . . . . . . Haemulon macrostomum p. 374 25a. Dorsal fin rays 15 to 17 (usually 16); anal fin rays 8 to 10 (usually 9) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 25b. Dorsal fin rays 16 to 29 (usually 17 to 19); anal fin rays 8 or 9 (usually 8) . . . . . . . . . . . . . . . Haemulon parra p. 376 26a. Scales above anterior section of lateral line larger than those below anterior section of lateral line; midlateral stripe absent in fresh specimens longer than 35 mm SL and preserved specimens longer than 20 mm SL; five scale rows above lateral line . . . . . . . . . . . Haemulon plumierii p. 377 26b. Scales above anterior section of lateral line same size as those below anterior section of lateral line; midlateral stripe

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HAEMULIDAE 363

present in specimens between 15 and 35 mm SL and usually present in specimens longer than 35 mm SL; eight scale rows above lateral line . . . . Haemulon sciurus p. 378 27a. Body relatively short and deep; head relatively short; snout blunt; lips thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 27b. Body relatively slender; head relatively long; snout acute; lips thin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 28a. Gill rakers on lower limb of first arch 16 to 18; pored lateral line scales 50 to 53; scale rows between base of first dorsal fin spine and lateral line 6 or 7; no black bars or bands on anterior section of body . . . . . . . . . . . . . . . . . . . . Anisotremus surinamensis p. 364 28b. Gill rakers on lower limb of first arch 13 to 15; pored lateral line scales 56 to 60; scale rows between base of first dorsal fin spine and lateral line 10 or 11; diagonal black band on head and black bar just behind head . . . . . . . . . . . . . . . . . . . . . . . Arisotremus virginicus p. 365 29a. Anal fin with three spines and 12 or 13 rays . . . . . . . . . . . . . . . . . . . . . Orthopristis chrysoptera p. 380 29b. Anal fin with three spines and 6 or 7 rays . . . . . . . . . . . . . . . . . . . . . . . . . . Pomadasys crocro p. 381

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364 PERCIFORMES

Anisotremus surinamensis Black margate

(Bloch, 1791)

Anisotremus surinamensis lacks scales on dorsal and anal fin membranes and has a large black mantle on anterior part of body. Dorsal profile of head is moderately steep and nearly straight. Nares are close set, posterior margin of anterior naris is extended as flap, and posterior naris is small and oval shaped. Jaw teeth are villiform to setiform and arranged in bands that narrow to a single row posteriorly, with some teeth of outer series enlarged and sharply pointed. Preoperculum is finely serrated on posterior margin, with serrae enlarged near slightly produced angle. Gill rakers on first arch number 11 to 14 on upper limb and 16 to 18 on lower limb. Measurements are expressed as percent of SL: head length 39%– 42%, snout length 11%–12%, eye diameter 12%–15%, upper jaw length 11%–12%, body depth 45%– 48%. Pectoral fin has 18 (rarely 17) rays. Dorsal fin is distinctly incised between spinous and rayed sections and has 12 spines and 16 or 17 rays. Anal fin has 8 or 9 rays. Caudal fin is forked. Scale rows between first dorsal fin spine and lateral line number six or seven, and scales encircling caudal peduncle number 22. Lateral line scales number 50 to

53. Body is whitish, with a broad black band or mantle over middle third of body but most distinct below lateral line. Pectoral fin is dusky, and other fins are black. Juveniles are light, with a narrow stripe running from nape to end of dorsal fin and another running from posterior margin of eye to large, round caudal fin spot. This species occurs in shallow water in the western Atlantic from Florida to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it is rare north of southern Florida and southern Texas, although it occurs on the Flower Garden Banks off East Texas. It is associated with coral reefs and rocky areas. Food consists of sea urchins, crustaceans, and small ray-finned fishes. Maximum known size is 600 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992 (undocumented from Alabama); Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Pattengill 1998; Castro-Aguirre et al. 1999; Lindeman and Toxey 2002.

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HAEMULIDAE 365

Arisotremus virginicus Porkfish

(Linnaeus, 1758)

Arisotremus virginicus has either a large and intense caudal spot or alternating dark and light stripes and lacks scales on dorsal and anal fin membranes. Dorsal profile of head is very steep and nearly straight. Nares are close set, posterior margin of anterior naris is extended as a flap, and posterior naris is small and oval shaped. Jaw teeth are villiform to setiform and arranged in bands that narrow to a single row posteriorly, with some teeth of outer series enlarged and sharp pointed. Preoperculum is finely serrated on posterior margin, with serrae enlarged near slightly produced angle. Gill rakers on first arch number 11 on upper limb and 13 to 15 on lower limb. Measurements are expressed as percent of SL: head length 32%– 33%, snout length 10%–13%, eye diameter 9%–12%, upper jaw length 9%–11%, body depth 48%–53%. Pectoral fin has 17 or 18 rays. Dorsal fin is slightly incised between spinous and rayed sections and has 12 spines and 16 or 17 rays. Anal fin has 9 rays. Caudal fin is forked. Scale rows between first dorsal fin spine and lateral line number 10 or 11, and scales encircling caudal peduncle number 24 to 26. Lateral line scales number 56 to 60. Body is patterned with broad silvery white stripes alternating with narrow yellow stripes. Head has an

oblique black band running from corner of mouth through eye to forehead and a nearly vertical band passing from first dorsal fin base to upper section of pectoral fin base. Spinous section of dorsal fin and pelvic fin is dusky to black, and other fins are yellow. Juveniles are yellowish, with two dark stripes on body and caudal spot in addition to head stripes of adults. This species occurs in shallow water in the western Atlantic from Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it is rare north of southern Florida and southern Texas. It is associated with coral reefs and rocky areas. Food consists of ophiurans, mollusks, annelids, and crustaceans, and feeding takes place at night. Juveniles clean other fishes of parasites. Maximum known size is 380 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Lindeman and Toxey 2002.

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366 PERCIFORMES

Conodon nobilis Barred grunt

(Linnaeus, 1758)

Conodon nobilis is elliptical in profile, with preoperculum strongly serrated and eight dark bands on body. Dorsal profile of head is slightly convex. Nares are moderately separated, posterior margin of anterior naris is expanded into a flap, and posterior naris is small and oval. Jaw teeth are villiform and arranged in bands that narrow to a single row posteriorly, with some on side of symphysis moderately enlarged and sharp pointed. Preoperculum has serrations resembling flat spines along margin, with serrations increasing to large spines at corner and those on ventral margin oriented anteroventrally. Gill rakers on first arch number 6 to 8 on upper limb and 14 to 16 on lower limb. Measurements are expressed as percent of SL: head length 30%–33%, snout length 9%– 10%, eye diameter 6%– 8%, upper jaw length 11%–12%, body depth 31%–33%. Pectoral

fin has 16 rays. Dorsal fin is deeply incised and has 12 spines and 12 or 13 rays. Anal fin has 7 rays. Caudal fin is emarginate. Scales occur on membranes between dorsal and anal fin rays but not on rays. Lateral line scales number 50 to 53. Color is light tan, with eight dark brown bars or saddles extending to lower flank. This species occurs to 100 m in the western Atlantic from eastern Florida to Brazil, including the Gulf of Mexico and the Antilles. In the Gulf of Mexico it occurs west of the Florida Panhandle to Yucatán. It is associated with soft substrates and turbid water, including estuaries. Maximum known size is 300 mm TL. references: Dawson 1962b; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; CastroAguirre et al. 1999; Lindeman and Toxey 2002.

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HAEMULIDAE 367

Haemulon album Margate

Cuvier, 1830

Haemulon album has a dense covering of scales on rayed sections of dorsal and anal fins and an obscure dark spot at the base of each scale. Dorsal profile of head is almost straight. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis and are enlarged, slender, slightly recurved, and in a single row on lateral aspects of jaws. Preoperculum is finely serrated, with serrae increasing in size toward angle in adults. Gill rakers on first arch number 21 to 23. Measurements are expressed as percent of SL: head length 35%–39%, snout length 14%–17%, upper jaw length 15%–16%, pre-dorsal-fin length 44%– 48%, body depth 37%– 42%. Pectoral fin has 18 or 19 rays. Dorsal fin has 12 spines and 16 or 17 rays, and anal fin has 8 (rarely 5 or 7) rays. Scales above pectoral fin base are not elongated vertically, and scales above and below lateral line are about the same size. Scale rows between first dorsal fin spine and lateral line number 14, and those between lateral line and first anal fin spine number 7.

Scales encircling caudal peduncle number 25 or 26 (rarely 27). Lateral line scales number 50 to 52 (rarely 49). Color is pearly gray, with rayed sections of dorsal and caudal fins dusky to black. Occasionally black stripes are present on pearly gray or olive ground color. Inside of mouth is red. Coloration of juvenile specimens is unknown. This species occurs between 20 and 60 m in the western Atlantic from the Florida Keys and Bermuda to Brazil, including the northeastern Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it occurs at the Dry Tortugas and from southwest Florida. It is associated with clear water around coral reefs, rocky bottoms, wrecks, and sea-grass beds. Food consists of crustaceans and small ray-finned fishes. Maximum known size is about 750 mm TL.

references: Longley and Hildebrand 1941; Courtenay 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Lindeman 1986; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Smith-Vaniz et al. 1999; Lindeman and Toxey 2002.

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368 PERCIFORMES

Haemulon aurolineatum Tomtate

Cuvier, 1830

Haemulon aurolineatum has a dense covering of scales on rayed sections of dorsal and anal fins and a large black blotch on caudal peduncle. Dorsal profile of head is evenly convex. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis, and are enlarged, slender, slightly recurved, and in a single row on lateral aspects of jaws. Preoperculum is finely serrated in adults. Gill rakers on first arch number 24 to 28. Measurements are expressed as percent of SL: head length 32%–38%, snout length 11%–17%, eye diameter 8%–9%, upper jaw length 15%–23%, body depth 27%–38%. Pectoral fin has 17 or 18 (rarely 16) rays. Dorsal fin has 13 spines and 14 to 16 rays, and anal fin has 9 (rarely 7 or 8) rays. Scale rows between first dorsal fin spine and lateral line number 11 to 13, and those between lateral line and first anal fin spine number 6. Scales encircling caudal peduncle number 22. Scales above pectoral fin base are not elongated vertically. Color is whitish to pale tan, with a yellow to bronze stripe running from opercular flap to

caudal peduncle blotch and a yellow stripe running from nape to end of dorsal fin. Inside of mouth is red. In juveniles, lateral stripe is separated from caudal fin spot except for presence of small melanophores between stripe and spot in small specimens. Caudal spot is oval in small specimens but becomes constricted and dumbbell shaped in large juveniles. This species occurs between the shoreline and about 30 m in the western Atlantic from Chesapeake Bay and Bermuda to Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is associated with sandy and rough bottoms, including artificial reefs. Maximum known size is about 250 mm TL.

references: Longley and Hildebrand 1941; Courtenay 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Castro-Aguirre and Márquez-Espinoza 1981; Lindeman 1986; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Murdy et al. 1997; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Lindeman and Toxey 2002.

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HAEMULIDAE 369

Haemulon bonariense Black grunt

Cuvier, 1830

Haemulon bonariense has a dense covering of scales on rayed sections of dorsal and anal fins and dark spots on scales forming oblique stripes. Dorsal profile of head is nearly straight. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis, and are enlarged, slender, slightly recurved, and in a single row on lateral aspects of jaws. Preoperculum is finely serrated, with serrae increasing in size toward angle in adults. Gill rakers on first arch number 18 to 24. Measurements are expressed as percent of SL: head length 35%– 42%, snout length 12%–15%, upper jaw length 13%–15%, pre-dorsal-fin length 41%– 46%, body depth 33%– 41%. Pectoral fin has 16 or 17 rays. Dorsal fin has 12 spines and 15 or 16 rays (rarely 17), and anal fin has 8 or 9 rays. Pectoral fin lacks scales. Scales above pectoral fin base are not elongated vertically, and scales above and below lateral line are about the same size. Scale rows between first dorsal fin spine and lateral line number 11 or 12 (rarely 10), and those between

lateral line and first anal fin spine number 6. Scales encircling caudal peduncle number 21 or 22 (rarely 19). Lateral line scales number 45 to 48 (rarely 44 to 49). Color is pale or silvery, with many wavy dark lines on head and body. Inside of mouth is red. Juveniles have three dark stripes on body and a large, elongated spot on base of caudal fin, and lateral stripe is widely separated from spot. Majority of caudal fin spot is posterior to caudal fin base, and part of spot anterior to base is expanded dorsoventrally. This species occurs in the western Atlantic from the Gulf of Mexico to Brazil, including the Greater Antilles. In the Gulf of Mexico it is known from the Dry Tortugas, Veracruz, and the Laguna de Términos, Campeche. It is associated with clear water over coral reefs, grass beds, algal beds, or mud. Maximum known size is about 400 mm TL. references: Longley and Hildebrand 1941; Courtenay 1961; Randall 1968a, 1996; Robins and Ray 1986; Cervigón 1993b; Castro-Aguirre et al. 1999; Schaldach et al. 1999c; Lindeman and Toxey 2002.

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370 PERCIFORMES

Haemulon boschmae

(Metzelaar, 1919)

Haemulon boschmae has a dense covering of scales on rayed sections of dorsal and anal fins, brown stripes on body, and a large spot spanning caudal peduncle and caudal fin base. Dorsal profile of head is slightly convex. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis, and are enlarged, slender, slightly recurved, and in one or two rows on lateral aspects of jaws. Preoperculum is finely serrated, with serrae increasing in size toward slightly produced angle in adults. Gill rakers on first arch number 29 to 36. Measurements are expressed as percent of SL: head length 29%– 31%, snout length 6%–7%, eye diameter 7%– 9%, upper jaw length 10%–11%, pre-dorsalfin length 34%–35%, body depth 26%–30%. Pectoral fin has 17 to 19 rays. Dorsal fin has 13 (rarely 14) spines and 13 or 14 rays, and anal

fin has 7 to 9 rays. Horizontal scale rows below lateral line are more or less parallel to long axis of body. Scales encircling caudal peduncle number 23 to 27. Lateral line scales number 49 to 54. Color is yellowish to greenish silver dorsally and silvery ventrally, with four brown to bronze stripes on sides and a large black spot on caudal peduncle and base of caudal fin. This species occurs between the shoreline and about 100 m in the western Atlantic from the southern Gulf of Mexico and from Colombia to French Guiana. In the Gulf of Mexico it has been reported from Isla de Lobos, Veracruz. Food consists of mollusks and crustaceans. Maximum known size is about 190 mm TL.

references: Randall 1968a, 1996; Castro-Aguirre and Márquez-Espinoza 1981; Lindeman 1986; Lindeman and Toxey 2002.

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HAEMULIDAE 371

Haemulon carbonarium Caesar grunt

Poey, 1860

Haemulon carbonarium has a dense covering of scales on rayed sections of dorsal and anal fins and bronze or coppery stripes on body. Dorsal profile of head is evenly convex. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis, and are enlarged, slender, slightly recurved, and in a single row on lateral aspects of jaws. Preoperculum is finely serrated, with serrae increasing in size toward angle in adults. Gill rakers on first arch number 23 to 25. Measurements are expressed as percent of SL: head length 35%–38%, snout length 14%–17%, upper jaw length 14%–17%, pre-dorsal-fin length 40%– 46%, body depth 34%– 40%. Pectoral fin has 16 or 17 rays. Dorsal fin has 12 spines and 15 or 16 rays, and anal fin has 8 (rarely 7 or 9) rays. Scales above pectoral fin base are not elongated vertically, and scales above and below lateral line are about the same size. Scale rows between first dorsal fin spine and lateral line number 13, and those between lateral line and first anal fin spine num-

ber 6. Scales encircling caudal peduncle number 22. Lateral line scales number 49 or 50 (rarely 52). Color is pale or silvery, with many straight stripes. Head is dark, with bronze stripes posteriorly. Fins are dusky to blackish. Inside of mouth is red. Juveniles have three dark stripes on body and a large rectangular spot with rounded edges on caudal peduncle, and lateral stripe is separated from spot. This species occurs between the shoreline and about 25 m in the western Atlantic from southern Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it occurs off the Dry Tortugas. It is associated with coral reefs. Feeding takes place at night, and food consists of polychaetes, gastropods, crabs, and asteroids. Maximum known size is greater than 425 mm TL. references: Longley and Hildebrand 1941; Courtenay 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Lindeman 1986; Robins and Ray 1986; Cervigón 1993b; Smith 1997; SmithVaniz et al. 1999; Lindeman and Toxey 2002.

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372 PERCIFORMES

Haemulon chrysargyreum Smallmouth grunt

Günther, 1859

Haemulon chrysargyreum has a dense covering of scales on rayed sections of dorsal and anal fins and yellowish stripes on body. Dorsal profile of head is evenly convex. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis, and are enlarged, slender, slightly recurved, and in a single row along lateral aspect of jaws. Preoperculum is finely serrated, with serrae larger toward slightly extended corner in adults. Gill rakers on first arch number 29 to 34. Measurements are expressed as percent of SL: head length 34%–38%, snout length 10%–13%, upper jaw length 10%– 13%, pre-dorsal-fin length 38%– 42%, body depth 27%–33%. Pectoral fin has 16 or 17 (rarely 15) rays. Dorsal fin has 12 spines and 12 or 13 (rarely 14) rays, and anal fin has 9 or 10 rays. Scales above pectoral fin base are not elongated vertically, and longitudinal scale rows below lateral line are parallel to body axis. Scale rows between first dorsal fin spine and lateral line number 11 or 12, and those between lateral line and first anal fin spine number 7. Scales encircling caudal peduncle num-

ber 21 or 22 (rarely 20). Lateral line scales number 49 to 51. Color is whitish, with bright yellow stripes. Lower side of head has yellow spots, and fins are yellow. Inside of mouth is red. Juveniles have three dark stripes and a spot on base of caudal fin, with lateral stripe separated from oval spot. This species occurs in shallow water in the western Atlantic from southern Florida and the Bahamas to Brazil, including the Gulf of Mexico and the Antilles. In the Gulf of Mexico it is known from the Dry Tortugas and Veracruz. Juveniles are associated with hard bottoms, coral reefs, or Thalassia grass beds, and adults are associated with coral reefs. Feeding takes place at night, and food consists of zooplankton such as copepods, amphipods, ostracods, shrimps, and crab larvae. Maximum known size is about 230 mm TL.

references: Longley and Hildebrand 1941 (as Brachygenys chrysargyreus); Courtenay 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Lindeman 1986; Robins and Ray 1986; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; Lindeman and Toxey 2002.

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HAEMULIDAE 373

Haemulon flavolineatum French grunt

(Desmarest, 1823)

Haemulon flavolineatum has a dense covering of scales on rayed sections of dorsal and anal fins and many straight and oblique stripes on body. Dorsal profile of head is nearly straight. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis, and are enlarged, slender, slightly recurved, and in a single row on lateral aspects of jaws. Preoperculum is finely serrated, with serrae increasing in size toward slightly produced angle in adults. Operculum lacks spine. Gill rakers on first arch number 20 to 25. Measurements are expressed as percent of SL: head length 35%–39%, snout length 10%–15%, upper jaw length 14%– 16%, pre-dorsal-fin length 40%– 46%, body depth 33%–38%. Pectoral fin has 16 or 17 rays. Dorsal fin has 12 spines and 14 or 15 rays, and anal fin has 8 (rarely 7) rays. Scales above pectoral fin base are not elongated vertically, and scales below lateral line are larger than those above. Scale rows between first dorsal fin spine and lateral line number 9 (rarely 10), and those between lateral line and first anal fin spine number 6. Scales encircling caudal peduncle number 22. Lateral line scales num-

ber 47 to 50. Color is pale or silvery, with many oblique yellowish bronze stripes. Head is dark, with bronze stripes posteriorly. Fins are dusky to blackish. Inside of mouth is red. Juveniles have three dark stripes on body and a rectangular spot with rounded edges on caudal peduncle, and lateral stripe is separated from spot. This species occurs in shallow water in the western Atlantic from South Carolina and Bermuda to Brazil, including the eastern and southern Gulf of Mexico, the Bahamas, and the Antilles. In the eastern Gulf of Mexico it is known from the Dry Tortugas. It is associated with reefs and sea-grass beds and appears in large schools over coral reefs. Feeding on small crustaceans takes place at night. Maximum known size is about 300 mm TL.

references: Longley and Hildebrand 1941; Courtenay 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Castro-Aguirre and MárquezEspinoza 1981; Lindeman 1986; Robins and Ray 1986; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; CastroAguirre et al. 1999; Smith-Vaniz et al. 1999; Lindeman and Toxey 2002.

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374 PERCIFORMES

Haemulon macrostomum Spanish grunt

Günther, 1859

Haemulon macrostomum has a dense covering of scales on rayed sections of dorsal and anal fins and three dark stripes on body. Dorsal profile of head is nearly straight. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis, and are enlarged, slender, slightly recurved, and in a single row on lateral aspects of jaws. Preoperculum is finely serrated, with serrae increasing in size toward slightly produced angle in adults. Gill rakers on first arch number 25 to 30. Measurements are expressed as percent of SL: head length 36%– 41%, snout length 13%–19%, upper jaw length 16%– 22%, pre-dorsal-fin length 42%– 48%, body depth 36%– 42%. Pectoral fin has 17 or 18 rays. Dorsal fin has 12 spines and 15 or 16 (rarely 17) rays, and anal fin has 9 (rarely 8) rays. Scales above pectoral fin base are not elongated vertically, and scales above and below lateral line are about the same size. Scale rows between first dorsal fin spine and lateral line number 12 (rarely 11), and those between lateral line and first anal fin spine number 7.

Scales encircling caudal peduncle number 22 (rarely 21). Lateral line scales number 50 to 52. Color is white to bronze, with three dark stripes on back and area below dorsal fin; first stripe is greenish yellow. Pectoral fin is greenish yellow. Inside of mouth is pale red. Juveniles have three dark stripes on body and a wedge-shaped dark spot on caudal peduncle, with the lateral stripe separated from spot. This species occurs in shallow, clear water in the western Atlantic from southern Florida to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it is known from the Dry Tortugas and has twice been reported from Texas, including a visual observation, from the Flower Garden Banks off eastern Texas. This species is associated with coral reefs, and juveniles appear to be associated only with reefs. Maximum known size is about 430 mm TL. references: Longley and Hildebrand 1941; Courtenay 1961; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Lindeman 1986; Robins and Ray 1986; Cervigón 1993b; Smith 1997; Pattengill 1998; Lindeman and Toxey 2002.

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HAEMULIDAE 375

Haemulon melanurum Cottonwick

(Linnaeus, 1758)

Haemulon melanurum has a dense covering of scales on rayed sections of dorsal and anal fins, a broad dark stripe running along dorsal margin of body to lower tip of tail, and a second stripe diverging from first at caudal peduncle and running to upper tip of tail. Dorsal profile of head is slightly arched. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis, and are enlarged, slender, and slightly recurved along margin of jaws. Preoperculum is finely serrated, with serrae increasing in size toward slightly produced angle in adults. Gill rakers on first arch number 21 to 23. Measurements are expressed as percent of SL: head length 35%–39%, snout length 12%–15%, upper jaw length 16%–18%, pre-dorsal-fin length 38%– 43%, body depth 33%–37%. Pectoral fin has 16 to 18 rays. Dorsal fin has 12 spines and 15 or 16 (rarely 17) rays, and anal fin has 8 (rarely 7 or 9) rays. Scales above pectoral fin base are not elongated vertically, and scales above and below lateral line are about the same size. Scale rows between first dorsal fin spine and lateral line number 13, and those between lateral line and first anal fin spine number 6. Scales encircling caudal peduncle number 23 to 25 (rarely 22). Lateral line scales number 49 to

51. Color is pale or silvery, with many straight stripes. Head is silvery white. Sides of body are marked by narrow yellow stripes. Inside of mouth is red. Juveniles have three dark stripes on body and an elongate spot on base of caudal fin, and lateral stripe is continuous with spot. Elongate caudal spot may appear as two partially divided spots. This species occurs between the shore and about 30 m in the western Atlantic from North Carolina and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it has been reported from Veracruz, and there are several reports of this species from offshore reefs and banks, including the Dry Tortugas and the Flower Garden Banks off eastern Texas. It is associated with clear water along outer reefs and sea-grass beds. Maximum known size is about 435 mm TL.

references: Longley and Hildebrand 1941; Courtenay 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Bright and Cashman 1974; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Lindeman 1986; Robins and Ray 1986; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; Pattengill 1998; Smith-Vaniz et al. 1999; Lindeman and Toxey 2002.

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376 PERCIFORMES

Haemulon parra (Desmarest, 1823) Sailors choice

Haemulon parra has a dense covering of scales on rayed sections of dorsal and anal fins and on pectoral fins, and bronze to gray stripes running along scale rows. Dorsal profile of head is nearly straight. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis, and are enlarged, slender, slightly recurved, and in a single row on lateral aspects of jaws. Preoperculum is finely serrated, with serrae increasing in size toward slightly produced angle in adults. Gill rakers on first arch number 21 to 24. Measurements are expressed as percent of SL: head length 35%– 41%, snout length 13%–16%, upper jaw length 14%–16%, pre-dorsal-fin length 41%– 49%, body depth 36%– 40%. Pectoral fin has 17 (rarely 16) rays. Dorsal fin has 12 spines and 16 to 18 (rarely 19) rays, and anal fin has 8 (rarely 9) rays. Scales above pectoral fin base are not elongated vertically, and scales above and below lateral line are about the same size. Scale rows between first dorsal fin spine and lateral line number 12, and those between lateral line and first anal fin spine number 6. Scales encircling caudal peduncle num-

ber 21 or 22. Lateral line scales number 47 to 50. Color is pearly gray to silvery, with spots on scales forming oblique stripes. Dorsal, pelvic, anal, and caudal fins are chalky. Inside of mouth is red. Juveniles have dark midlateral stripe on body and round dark spot on caudal peduncle, with the lateral stripe separated from or connected to spot by a few melanophores. This species occurs in shallow water in the western Atlantic from Florida to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. It is common in the eastern Gulf of Mexico from the Dry Tortugas and also occurs in Veracruz, but it is rare to absent in the northwestern Gulf. Juveniles are associated with grass beds. Adults school in the more open areas of coral reefs. Feeding takes place at night, and food consists of polychaetes, mollusks, and small ray-finned fishes. Maximum known size is about 400 mm TL. references: Longley and Hildebrand 1941; Courtenay 1961; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Robins and Ray 1986; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; Lindeman and Toxey 2002.

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HAEMULIDAE 377

Haemulon plumierii White grunt

(Lacepède, 1801)

Haemulon plumierii has a dense covering of scales on rayed sections of dorsal and anal fins and blue and yellow stripes on head. Dorsal profile of head is nearly straight. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis, and are enlarged, slender, and slightly recurved in lateral aspects of jaws. Preoperculum is finely serrated, with serrae increasing in size toward slightly produced angle in adults. Gill rakers on first arch number 21 to 28. Measurements are expressed as percent of SL: head length 37%– 41%, snout length 15%–20%, upper jaw length 17%–21%, pre-dorsal-fin length 39%– 48%, body depth 37%– 42%. Pectoral fin has 17 (rarely 16) rays. Dorsal fin has 12 spines and 15 to 17 rays, and anal fin has 8 or 9 rays. Scales above pectoral fin base are not elongated vertically but are larger than those below lateral line. Scale rows between first dorsal fin spine and lateral line number 11, and those between lateral line and first anal fin spine number 5. Scales encircling caudal peduncle number 22 (rarely 23). Lateral line scales number 48 to 51. Color is bluish gray dorsally and paler laterally and ventrally, with

many alternating blue and yellow stripes on head. Scales on body are whitish with blue centers. Some specimens have a broad black stripe behind pectoral fin extending about half of the distance to caudal fin and a black spot beneath edge of preoperculum. Inside of mouth is bright red. Juveniles have several stripes on body and a round spot on caudal peduncle, and lateral stripe is continuous with spot. This species occurs in shallow water in the western Atlantic from Maryland to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. It is found throughout the Gulf of Mexico within its depth range. Adults school during the day near patch reefs. Feeding takes place during the night, and food consists of polychaetes, gastropods, and crustaceans. Maximum known size is about 450 mm TL.

references: Longley and Hildebrand 1941; Courtenay 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hastings et al. 1976; Hoese and Moore 1977, 1998; Hastings 1979; Lindeman 1986; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999; Lindeman and Toxey 2002.

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378 PERCIFORMES

Haemulon sciurus Bluestriped grunt

(Shaw, 1803)

Haemulon sciurus has a dense covering of scales on rayed sections of dorsal and anal fins and alternating blue and yellow stripes on head and body. Dorsal profile of head is nearly straight. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis, and are enlarged, slender, and slightly recurved on lateral aspects of jaws. Preoperculum is finely serrated. Gill rakers on first arch number 26 to 31. Measurements are expressed as percent of SL: head length 36%– 40%, snout length 13%– 16%, upper jaw length 16%–19%, pre-dorsalfin length 42%– 46%, body depth 36%–39%. Pectoral fin has 16 or 17 (rarely 15) rays. Dorsal fin has 12 spines and 16 or 17 rays (rarely 15), and anal fin has 9 or 10 (rarely 8) rays. Scales above pectoral fin base are not elongated vertically and are about the same size above and below lateral line. Scale rows between first dorsal fin spine and lateral line number 12, and those between lateral line and first anal fin spine number 8 (rarely 7). Scales encircling caudal peduncle number 22. Lateral line scales number 48 to 51. Color is bronze dorsally, yel-

low on sides, and cream on belly, with blue spots running continuously from head to caudal peduncle. Dorsal and caudal fins are dusky, and other fins are yellow. Inside of mouth is red. Juveniles have three stripes on body and a round spot on caudal peduncle, and lateral stripe is continuous with oval spot. This species occurs in shallow water in the western Atlantic from South Carolina and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. It is abundant in the eastern Gulf of Mexico at the Dry Tortugas, but it is rare to absent in the northwestern Gulf and uncommon in the southern Gulf. It is associated with coral reefs, mangroves, and sea grasses. Juveniles are most common in shallow water. Food consists of crustaceans and small ray-finned fishes. Maximum known size is about 450 mm TL.

references: Longley and Hildebrand 1941; Courtenay 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Lindeman 1986; Robins and Ray 1986; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; CastroAguirre et al. 1999; Smith-Vaniz et al. 1999; Lindeman and Toxey 2002.

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HAEMULIDAE 379

Haemulon striatum Striped grunt

(Linnaeus, 1758)

Haemulon striatum has a dense covering of scales on rayed sections of dorsal and anal fins and five dark brown stripes on body. Dorsal profile of head is slightly convex. Nares are close set, anterior naris is oval and possesses a flap along posterior margin, and posterior naris is small and oval. Jaw teeth are villiform and arranged in a patch on either side of symphysis, and are enlarged, slender, and slightly recurved on lateral aspects of jaw. Preoperculum is finely serrated, with serrae increasing in size toward slightly produced angle in adults. Gill rakers on first arch number 27 to 36. Measurements are expressed as percent of SL: head length 28%–32%, snout length 8%– 11%, upper jaw length 11%–14%, pre-dorsalfin length 33%–38%, body depth 26%–32%. Pectoral fin has 17 to 19 rays. Dorsal fin has 13 spines and 12 to 14 (rarely 15) rays, and anal fin has 7 or 8 (rarely 9) rays. Scales above pectoral fin base are not elongated vertically, and scales above and below lateral line are about the same size. Scale rows between first dorsal fin spine and lateral line number 12 to 14,

and those between lateral line and first anal fin number 7 (rarely 6). Scales encircling caudal peduncle number 24 to 27. Lateral line scales number 50 to 56. Color is pale tan to silvery, with five yellow-edged stripes on body, and front of head is olive brown. Inside of mouth is red. Juvenile specimens have three midlateral stripes and no caudal spot. This species occurs between 12 and 100 m in the western Atlantic from Florida to Brazil, including the Gulf of Mexico, the western Bahamas, and the Antilles. It is rare in the northern Gulf of Mexico and has been reported only once from the eastern Gulf, at the Dry Tortugas. Food consists of zooplankton. Maximum known size is about 280 mm TL.

references: Jordan and Thompson 1905; Longley and Hildebrand 1941 (as Bathystoma striatum); Courtenay 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Lindeman 1986; Robins and Ray 1986; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; VegaCendejas et al. 1997; Lindeman and Toxey 2002.

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380 PERCIFORMES

Orthopristis chrysoptera (Linnaeus, 1766) Pigfish

Orthopristis chrysoptera is ovate or elliptical in profile, with dorsal and anal fin rays free of scales. Dorsal profile of head is slightly convex. Nares are close set, posterior margin of anterior naris is expanded into an elongate flap, and posterior naris is small with a fringed flap on posterior margin. Jaw teeth are villiform or setiform and arranged in a narrow band, with outer teeth slightly larger. Preoperculum is finely serrated along posterior margin, and corner is not produced and is free of serrae. Gill rakers on first arch number 8 on upper limb and 12 or 13 on lower limb. Measurements are expressed as percent of SL: head 31%–33%, snout length 12%–15%, eye diameter 6%–7%, upper jaw length 9%–10%, body depth 35%–37%. Pectoral fin has 17 to 19 rays. Dorsal fin is very shallowly notched between last two spines and has 12 or 13 spines and 15 or 16 rays. Anal fin has 12 or 13 rays. Caudal fin is forked. Scale rows between first dorsal fin spine and lateral line number 10,

and those between lateral line and first anal fin spine number 19. Lateral line scales number 55 to 58. Color is light bluish gray dorsally and shades to silvery ventrally, with blue and orange spots on body scales forming oblique stripes. Head has orange spots, and fins are yellowish bronze. This species occurs in the western Atlantic from New Jersey and Bermuda to the southern Gulf of Mexico and the northern coast of Yucatán, including the coast of Cuba. It occurs throughout the Gulf of Mexico in shallow water over muddy and sandy bottoms, including bays and estuaries. Food consists of benthic crustaceans and ray-finned fishes. Maximum known size is 460 mm TL. references: Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997 (as O. poeyi); Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Lindeman and Toxey 2002.

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HAEMULIDAE 381

Pomadasys crocro Burro grunt

(Cuvier, 1830)

Pomadasys crocro is elliptical in profile, with dorsal and anal fin rays free of scales. Dorsal profile of head is slightly convex. Anterior naris has an elongate flap on posterior margin, and posterior naris is small and elliptical. Jaw teeth are setiform and widely spaced, with outer teeth slightly larger. Preoperculum is coarsely serrated, with serrae increasing in size to moderately produced angle. Gill rakers on first arch number five or six on upper limb and seven to nine on lower limb. Measurements are expressed as percent of SL: head 36%–37%, eye diameter 9%–10%, upper jaw length 11%– 13%, body depth 35%–38%. Pectoral fin has 16 rays. Dorsal fin is very shallowly notched between last two spines and has 13 spines and 12 or 13 rays. Anal fin has 7 rays. Caudal fin is emarginate. Scale rows between first dorsal fin spine and lateral line number 5 or 6,

and those between lateral line and first anal fin spine number 16. Lateral line scales number 53 to 55. Color is olivaceous dorsally and silvery ventrally. Pelvic and anal fins are yellowish. This species occurs in the western Atlantic from the northeastern and southern Gulf of Mexico and the Greater and Lesser Antilles to Brazil in sheltered inshore waters, including bays and estuaries. It also occurs in brackish waters. It is rare to absent in the northwestern Gulf of Mexico. Food consists of crustaceans and ray-finned fishes. Maximum known size is 330 mm TL. references: Hoese 1965; Hoese and Moore 1977, 1998; Pezold and Edwards 1983; Robins and Ray 1986; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Lindeman and Toxey 2002.

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382

INERMIIDAE

Bonnetmouths

Bonnetmouths are elongate and fusiform, with dorsal fin subdivided by deep notch or separated into two fins and caudal fin deeply forked. Dorsal profile of head is nearly straight to slightly convex. Snout is about equal to eye diameter. Nostril is paired, and nares are located near dorsal margin of head. Mouth is terminal, relatively small, and moderately oblique, with lower jaw projecting slightly beyond upper jaw. Upper jaw is very protrusible. Maxilla is completely covered by lachrymal bone when mouth is closed. Teeth are lacking in jaws, vomer, and palatine. Chin has two enlarged pores. Preoperculum is coarsely serrated along posterior margin, and corner is produced and acutely rounded. Operculum has a broad, flat spine on posterior margin. Gill membranes are narrowly united at isthmus. Gill rakers on first arch are long, slender, and very numerous. Pseudobranch is well developed. Pectoral fin inserts relatively high on flank and is pointed. Dorsal fin has 9 or 10, or 14 to 17, slender spines in anterior section and 2 slender spines and 9 or 10 rays in posterior section. Pelvic fin inserts slightly posterior to pectoral fin base and has 1 spine and 5 rays. Anal fin is located beneath posterior section of dorsal fin and has 2 or 3 slender spines and 9 to 11 rays. Caudal fin is deeply forked, and lobes fold in scissorlike fashion. Body and much of the head are covered with medium-sized ctenoid scales. Maxilla lacks scales. Sheath of scales runs along base of posterior section of dorsal fin. Axillary process is located on base of pelvic fin. Lateral line extends onto caudal fin, nearly to margin. Vertebrae number 26: 12 or 13 precaudal and 13 or 14 caudal. Maximum known size is 250 mm TL. Bonnetmouths occur in the tropical western Atlantic over continental and insular shelves to a depth of 300 m. They are pelagic schooling fishes that feed on zooplankton and small ray-finned fishes in clear water. Prey organisms are caught individually by means of the greatly protrusible jaws. Bonnetmouths often school around coral reefs. There are two species in separate genera, both occur in the western Central Atlantic, and at least one occurs in the Gulf of Mexico. references: Jordan and Evermann 1896; Johnson 1980, 1984; Fritzsche and Fuiman 1982; Robins and Ray 1986; Nelson 1994; Randall 1996; Smith 1997.

Key to the Species of the Gulf of Mexico (Adapted from Böhlke and Chaplin 1968) 1a. Sections of dorsal fin widely separated, with interspace between sections scaled . . . . . . . . . . . . . . . . . . . Emmelichthyops atlanticus p. 383 1b. Sections of dorsal fin not separated or barely separated, with interspace unscaled . . . . . . . . . Inermia vittata p. 384

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INERMIIDAE 383

Emmelichthyops atlanticus Bonnetmouth

Schultz, 1945

Emmelichthyops atlanticus has scaled interspace between dorsal fins and a deeply forked caudal fin. Upper jaw extends to anterior margin of eye. Gill rakers on first arch number 8 on upper limb and about 20 on lower limb. Measurements are expressed as percent of SL: head length 25%–27%, snout length 6%– 8%, eye diameter 7%–9%, upper jaw length 10%– 12%, pectoral fin length 15%–18%, pelvic fin length 13%–15%, body depth 16%–18%. Pectoral fin has 18 or 19 rays. First dorsal fin has 9 or 10 spines, with last several fitting into groove when depressed. Second dorsal fin has 2 spines and 9 or 10 rays. Anal fin has 2 spines and 9 to 11 rays. Caudal peduncle is broad. Body and most of head are covered with ctenoid scales. Axillary process on pelvic fin base is about half the length of pelvic fin. Lateral line scales number about 75 to end of caudal pe-

duncle. Color is metallic, yellowish gray dorsally and whitish on side and ventrally, with a bluish tinge anteriorly and four brown stripes on upper side. In preservative, specimens are pale, with several brown stripes on upper body. Juveniles have three brown stripes. This species occurs to a depth of 91 m in the western Atlantic from the Florida Keys and the Bahamas to northern South America, including the Virgin Islands, Barbados, and Tobago. There is a single record in the Gulf of Mexico based on a sighting at the West Flower Garden Bank (Pattengill 1998). This species is associated with coral reefs. Maximum known size is 125 mm TL. references: Schultz 1945; Böhlke and Chaplin 1968; Robins and Ray 1986; Randall 1996; Smith 1997; Pattengill 1998; Orrell 2002b.

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384 PERCIFORMES

Inermia vittata Boga

Poey, 1860

Inermia vittata has a deeply notched or marginally subdivided dorsal fin and a deeply forked caudal fin. Upper jaw extends to anterior margin of eye. Gill rakers on first arch are very long and slender and number 8 on upper limb and 23 to 25 on lower limb. Measurements are expressed as percent of SL: head length 25%–27%, snout length 6%–7%, eye diameter 7%– 8%, upper jaw length 8%–10%, pectoral fin length 17%–20%, pelvic fin length 12%–14%, body depth 19%–22%. Pectoral fin has 19 or 20 rays. First dorsal fin has 14 to 17 spines, with last several fitting into groove when depressed. Second dorsal fin has 2 spines and 9 or 10 rays. Anal fin has 2 spines and 9 rays. Caudal peduncle is broad and has lateral keel. Body and most of head are covered with ctenoid scales. Axillary process above pelvic fin is about half the length of pelvic fin. Lateral

line scales number about 80 to end of caudal peduncle. Color is metallic blue to green dorsally and bluish white ventrally, with a broad greenish stripe extending from eye to base of tail and three similar stripes on upper flank. Snout is yellowish. This species occurs to a depth of 91 m in the western Atlantic from North Carolina and Bermuda to northern South America, including the northern Gulf of Mexico, the Bahamas, and the Lesser Antilles. In the Gulf of Mexico it has been captured off Pascagoula, MS, and off southwestern Florida (2505N, 8339W). Maximum known size is 230 mm TL.

references: Böhlke and Chaplin 1968; Robins and Ray 1986; Randall 1996; Smith 1997; SmithVaniz et al. 1999; Orrell 2002b.

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385

SPARIDAE

Porgies

Porgies are oblong to ovate in lateral profile and usually deep bodied and moderately compressed to compressed, with a large head and a small mouth. Dorsal profile of head is usually steep. Mouth is small and horizontal to slightly oblique, and upper jaw is slightly protrusible. Posterior end of maxilla is covered by lachrymal and second suborbital bones when mouth is closed. Nostril is paired, and nares are located closer to eye than to tip of snout. Jaw teeth are usually conical or incisor-like in front of jaws and rounded and molariform on side of jaws. Vomer and palatine lack teeth. Preoperculum lacks serrations. Operculum has 2 flat spines. Gill membrane is free of isthmus, and last full arch (fourth) has slit posteriorly. Gill rakers on first arch number 7 to 20 on lower limb. Pseudobranch is well developed. Pectoral fin is well developed and narrowly pointed, with a nearly vertical base. Dorsal fin is little to slightly notched and consists of 10 to 13 spines and 9 to 17 rays. Pelvic fin inserts below to slightly behind pectoral fin base and has 1 spine and 5 rays. Anal fin is considerably shorter than dorsal fin and has 3 spines and 7 to 15 rays. Caudal fin is weakly to strongly forked and consists of 17 principal rays. Scales are weakly ctenoid and cover body and head except for snout and suborbital region. Rayed sections of dorsal and anal fins lack scales. Lateral line is complete and does not extend onto caudal fin. Axillary process at base of pelvic fin is well developed. Gas bladder is present and usually has a posterior extension into caudal myomeres. Vertebrae number 24: 10 precaudal and 14 caudal. Maximum known size is 75 cm TL. Porgies occur worldwide in tropical to warm temperate seas on continental and insular shelves. Some species enter estuaries, and a few occur in freshwater. Many species are hermaphroditic; some of these are first sexually mature as males and later change to females (protandry), others change from females to males (protogyny), and a few have functional ovaries and testes at the same time. Eggs and larvae are pelagic. Eggs range from 0.8 to 1.2 mm in diameter. Larvae have spines on preopercular, interopercular, subopercular, opercular, cleithral, postcleithral, supracleithral, posttemporal, and tabular bones. Food consists largely of benthic invertebrates, mostly polychaetes, mollusks, crustaceans, and sea urchins. There are about 100 species in 29 genera, 19 species in 6 genera in the western Central Atlantic, and 16 species in 6 genera in the Gulf of Mexico. references: Johnson 1980, 1984; Fritzsche and Fuiman 1982; Smith and Smith 1986; Nelson 1994; Tominaga et al. 1996; Smith 1997; Smith-Vaniz et al. 1999; Carpenter 2001c, 2002b.

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386 PERCIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Randall and Caldwell 1966) 1a. Teeth in anterior part of jaws strongly flattened, not conical .............................................2 1b. Teeth in anterior part of jaws slender and conical. . . . . . . 7 2a. Teeth in anterior part of jaws narrow and in close-set bands, with those in outer row enlarged and lanceolate; body without dark spots, stripes, or bars . . . . . . . . . . . . . . . . . . . . . . . . Stenotomus caprinus p. 404 2b. Teeth in anterior part of jaws broad and incisor-like; body with dark spots, stripes, or bars . . . . . . . . . . . . . . . . . . . . 3 3a. Caudal peduncle with large black blotch; without procumbent (lying flat and in forward direction) spine preceding dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3b. Caudal peduncle without large black blotch; with procumbent spine preceding dorsal fin (may be hidden by scales and integument . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4a. Anterior section of both jaws with eight well-developed incisor-like teeth; in adults black blotch often continuous over dorsal midline of caudal peduncle but generally not reaching ventral margin . . . . . . Diplodus argenteus p. 400 4b. Anterior section of both jaws with six well-developed incisor-like teeth; in adults black blotch sometimes forming complete ring around caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . Diplodus holbrookii p. 401 5a. Incisor-like teeth in anterior section of jaws without or with only shallow notch; lateral molariform teeth in three rows in upper jaw and in two rows in lower jaw . . . . . . . 6 5b. Incisor-like teeth in anterior section of jaws deeply notched; lateral molariform teeth in two and one-half rows in both jaws . . . . . . . . . . . . . . . . . . . . Lagodon rhomboides p. 402 6a. Dorsal fin usually with 12 spines; nape and body with five or six dark bars, slightly narrower than pale interspaces; origin of lateral line without dark spot . . . . . . . . . . . . . . . . . Archosargus probatocephalus p. 389 6b. Dorsal fin with 13 spines; nape and body with golden yellow stripes; origin of lateral line with dark spot . . . . . . . . . . . . . . . . . . . . Archosargus rhomboidalis p. 390 7a. Color pinkish to reddish; posterior naris oval; anal fin with 8 rays; suborbital space shallow. . . . . Pagrus pagrus p. 403 7b. Color bluish, copper, or occasionally yellowish, with bright silvery reflections; posterior naris elongate and slitlike; anal fin with 10 or 11 rays; suborbital space deep. . . 8 8a. Lateral line scales 43 to 49; pectoral fin rays 15 or 16 (rarely 14); no enlarged canine teeth in front of upper jaw and no molariform teeth on side of upper jaw . . . . . . . . . 9 8b. Lateral line scales 50 to 57; pectoral fin rays 14 or 15 (rarely 13 or 16); one or two enlarged canine teeth in front

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SPARIDAE 387

of upper jaw (except Calamus nodosus) and group of small molariform teeth anterior and medial to third row of teeth on side of upper jaw . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 9a. Blackish spot, darker than other markings on body and about 4 or 5 lateral line scales in diameter, on anterior section of lateral line; generally with either alternating wavy light and dark lines on head below eye or with dark V-shaped marking on base of caudal fin . . . . . . . . . . . . . 10 9b. Blackish spot present or absent on anterior section of lateral line and, if present, not darker than other markings on body; without wavy lines on head below eye or V-shaped marking on caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10a. Pectoral fin rays usually 16; gill rakers on first arch modally 10; dark V-shaped marking usually present on base of caudal fin; without alternating light and dark wavy lines on head below eye . . . . . . . . . . . Calamus arctifrons p. 391 10b. Pectoral fin rays usually 14 or 15; gill rakers on first arch modally 12; without dark V-shaped marking on base of caudal fin; alternating light and dark wavy lines on head below eye. . . . . . . . . . . . . . . Calamus campechanus p. 394 11a. Pectoral fin rays usually 16; upper base of pectoral fin without prominent small dark spot; pectoral fin usually 36% to 42% of SL in specimens greater than 80 mm SL; body depth 44% to 54% of SL . . . . . Calamus leucosteus p. 395 11b. Pectoral fin rays usually 15; upper base of pectoral fin with prominent small dark spot; pectoral fin usually 30% to 36% of SL in specimens greater than 80 mm SL; body depth 39% to 50% of SL . . . . . . . . Calamus penna p. 397 12a. Pectoral fin rays 15 (rarely 14 or 16); adults without curved canine teeth; dorsal profile of head below eye forming 43 to 55 angle with horizontal line from tip of snout; without light and dark wavy lines on head below eye . . . . . . . . . . . . . . . . . . . . . . . . . Calamus bajonado p. 392 12b. Pectoral fin rays 14 (rarely 13 or 15); third or fourth canine from symphysis enlarged and outcurved in adults (except in C. nodosus); dorsal profile of head below eye forming 57 to 65 angle with horizontal line from tip of snout; usually with light and dark wavy lines on head below eye . . . . . 13 13a. Anal fin with 10 (rarely 9 or 11) rays; unscaled portion of cheek and snout with irregular dark and light horizontal lines; broad pale blue horizontal band above gill opening . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13b. Anal fin with 11 (rarely 10) rays; unscaled portion of cheek and snout dark with pale spots; no blue horizontal band above gill opening . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14a. Dorsal profile of head above eye forming 32 to 40 angle with horizontal line; longest dorsal fin spine 12% to 15% of SL in specimens greater than 100 mm SL . . . . . . . . . . . . . . . . . . . . . . . . . Calamus pennatula p. 398

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388 PERCIFORMES

14b. Dorsal profile of head above eye forming 43 to 69 angle with horizontal line; longest dorsal fin spine usually 14% to 16% of SL in specimens greater than 100 mm SL . . . . . . . . . . . . . . . . . . . . . . . . . Calamus proridens p. 399 15a. Third canine from symphysis in upper jaw enlarged in specimens greater than 160 to 180 mm SL, and strongly outcurved in adults; body depth 44% to 50% of SL; dorsal profile of head slightly convex and section above eye forming 35 to 45 angle with horizontal line . . . . . . . . . . . . . . . . . . . . . . . . . . Calamus calamus p. 393 15b. Anterior canine teeth in upper jaw about equal in size and none outcurved; body depth 47% to 56% of SL; dorsal profile of head from snout to above center of eye nearly straight and section above eye forming 40 to 54 angle with horizontal line . . . . . . . . . . . Calamus nodosus p. 396

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SPARIDAE 389

Archosargus probatocephalus (Walbaum, 1792) Sheepshead

Archosargus probatocephalus has a small procumbent spine preceding dorsal fin, a very stout and long second anal spine, and dark bars on body. Dorsal profile of head is steep and angular at level of eyes. Posterior naris is long and slitlike. Four anterior teeth in jaws are broad and incisor-like and are not notched or only slightly notched. Lateral teeth in jaws are molariform and in three rows in upper jaw and two rows in lower jaw. Upper and lower lips possess arched grooves running most of length of jaws. Gill rakers on first arch number eight or nine. Measurements are expressed as percent of SL: head length 31%–32%, snout length 13%–14%, eye diameter 7%–9%, suborbital depth 6%– 8%, pectoral fin length 33%–37%, second anal fin spine length 17%– 20%, body depth 44%–53%, caudal peduncle depth 11%–13%. Pectoral fin has 15 to 17 rays, dorsal fin has 10 to 13 (usually 12) spines and 10 to 13 (usually 11 or 12) rays, and anal fin has 9 to 11 rays. Lateral line scales number 45 to 49. Color is gray to olive brown dorsally,

with five or six (rarely four to seven) dark bars on body. Bars are normally slightly narrower than pale interspaces. Juveniles are more brightly colored than adults. This species occurs along coasts and in estuaries and brackish water in the western Atlantic from Nova Scotia to Brazil, including the northern and southern Gulf of Mexico. It is divided into three subspecies: Archosargus p. probatocephalus occurs from Nova Scotia to Cedar Key, FL, in the northeastern Gulf of Mexico; A. p. oviceps occurs from St. Marks, FL, in the northeastern Gulf of Mexico, to Campeche Bank; and A. p. aries occurs from Belize to Rio de Janeiro. Food consists of mollusks and crustaceans. Maximum known size is 910 mm TL. references: Hastings et al. 1976; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Carpenter 2002b.

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390 PERCIFORMES

Archosargus rhomboidalis Sea bream

(Linnaeus, 1758)

Archosargus rhomboidalis has a small procumbent spine preceding dorsal fin, a very stout and long second anal fin spine, and a striped body. Dorsal profile of head is slightly convex to straight. Posterior naris is long and slitlike. Four anterior teeth in jaws are broad and incisor-like and are not notched or only slightly notched. Lateral teeth in jaws are molariform and arranged in three rows in upper jaw and two rows in lower jaw. Upper and lower lips possess arched grooves running most of length of jaws. Gill rakers on first arch number eight or nine on lower limb. Measurements are expressed as percent of SL: head length 27%– 32%, snout length 10%–13%, eye diameter 6%–9%, suborbital depth 6%–7%, pectoral fin length 34%– 42%, second anal fin spine length 14%–18%, body depth 46%–51%, caudal peduncle depth about 12%. Pectoral fin has 14 or 15 rays, dorsal fin has 13 or 14 (rarely

12) spines and 10 or 11 rays, and anal fin has 10 or 11 rays. Lateral line scales number 46 to 53. Color is silvery olivaceous, with golden yellow stripes and blackish spot about size of eye near origin of lateral line. This species occurs along the coast and in estuaries and brackish water in the western Atlantic from New Jersey to Brazil, including the northeastern and southern Gulf of Mexico and the Antilles. It is rare to absent along the Florida gulf coast (R. E. Matheson, pers. comm., August 2002). It is associated with muddy bottoms, mangrove habitats, and coral reefs that are near mangroves. Food consists of benthic mollusks, crustaceans, and plant material. Maximum known size is 320 mm TL. references: Randall 1968a, 1994; Robins and Ray 1986; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Carpenter 2002b.

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SPARIDAE 391

Calamus arctifrons Grass porgy

Goode and Bean, 1882

Calamus arctifrons has a small procumbent spine preceding dorsal fin, a relatively short pectoral fin, and a distinct dark spot on anterior section of lateral line. Dorsal profile of head is smoothly convex, with portion below eye forming 50 to 57 angle with horizontal line from tip of snout above upper lip to midbase of caudal fin. Posterior naris is long and slitlike. Anterior jaw teeth are conical and about equal in size. Teeth on sides of jaws are in three rows in upper jaw and two rows in lower jaw, and molariform teeth are lacking medial to rows in upper jaw. Lower anterior section of maxilla possesses bony tubercle with free semicircular margin. Tubercle on prefrontal bone, just behind posterior naris, is poorly developed. Gill rakers on first arch are short and number 10 to 12. Measurements are expressed as percent of SL: head length 29%–33%, snout length 17%–20%, eye diameter 8%–9%, suborbital depth 13%–15%, pectoral fin length 30%–32%, length of long-

est dorsal fin spine 13%–15%, body depth 40%– 43%, caudal peduncle depth 10%–11%. Pectoral fin has 15 or 16 rays, dorsal fin is slightly notched and has 11 to 13 spines and 11 to 13 rays, and anal fin has 10 (rarely 11) rays. Lateral line scales number 43 to 49. Color is light brown, with a spot slightly smaller than eye on anterior section of lateral line, five vertical rows of dark blotches on side, and often a dark V-shaped marking near base of caudal fin. Juveniles have more distinct blotches and dark bar on head through eye. This species occurs in the western Atlantic from southern Florida and the Florida Keys to Louisiana in coastal waters. It is associated with sea-grass beds. Maximum known size is 250 mm TL.

references: Randall and Caldwell 1966; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Smith 1997; Carpenter 2002b.

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392 PERCIFORMES

Calamus bajonado Jolthead porgy

(Bloch and Schneider, 1801)

Calamus bajonado has two enlarged canine teeth in upper jaw, a small procumbent spine preceding dorsal fin, and a relatively long pectoral fin. Dorsal profile of head is smoothly convex, with portion below eye forming 43 to 55 angle with horizontal line from tip of snout above upper lip to midbase of caudal fin. Posterior naris is long and slitlike. Anterior jaw teeth are conical, with second and third tooth from symphysis enlarged but not outcurved in specimens greater than 135 mm TL. Teeth on sides of jaws are molariform and in three rows in upper jaw and two rows in lower jaw, with additional irregular row of molariform teeth medial to three rows in upper jaw. Lower anterior section of maxilla possesses bony tubercle with free semicircular margin. Tubercle on prefrontal bone, just behind posterior naris, is poorly developed. Gill rakers on first arch are short and number 11 to 14. Measurements are expressed as percent of SL: head length 31%– 34%, snout length 17%–20%, eye diameter 9%–10%, suborbital depth 10%–14%, pectoral fin length 32%– 40%, length of longest dorsal fin spine 11%–14%, body depth 40%– 48%, caudal peduncle depth 9%–10%. Pec-

toral fin has 14 to 16 rays, dorsal fin is slightly notched and has 12 spines and 11 or 12 rays, and anal fin has 10 or 11 rays. Lateral line scales number 50 to 57. Color is silvery to light brown, with edges of scales darker than centers. Naked section of head is brown, and occasionally with dark brown bar from eye to cheek. Juveniles may have dark vertical bars on body and crossbars on caudal fin lobes. This species occurs in coastal waters to 45 m in the western Atlantic from Rhode Island and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is rare to absent along the Florida gulf coast and in the western Gulf of Mexico off Tamaulipas (R. E. Matheson, pers. comm., August 2002). Food consists of sea urchins, mollusks, and crabs. Maximum known size is 680 mm TL.

references: Randall and Caldwell 1966; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Randall 1996; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Carpenter 2002b.

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SPARIDAE 393

Calamus calamus Saucereye porgy

(Valenciennes, 1830)

Calamus calamus has a small procumbent spine preceding dorsal fin, one or two enlarged canine teeth in upper jaw, and a blue line below eye. Dorsal profile of head is slightly convex, with portion below eye forming 60 to 65 angle with horizontal line from tip of snout above upper lip to midbase of caudal fin. Posterior naris is long and slitlike. Anterior jaw teeth are conical, with third and occasionally fourth teeth from symphysis enlarged in specimens 160 to 180 mm SL and larger, and outcurved in specimens greater than 220 mm SL. Teeth on sides of jaws are molariform and in three rows in upper jaw and two rows in lower jaw, and an irregular row of molariform teeth is present medial to three rows in upper jaw. Lower anterior section of maxilla possesses bony tubercle with free semicircular margin. Tubercle on prefrontal bone, just behind posterior naris, is well developed. Gill rakers on first arch are short and number 11 to 13. Measurements are expressed as percent of SL: head length 29%–31%, snout length 16%– 18%, eye diameter 10%–11%, suborbital depth 11%–13%, pectoral fin length 34%– 39%, length of longest dorsal fin spine 12%– 15%, body depth 44%–50%, caudal peduncle

depth 9%–10%. Pectoral fin has 14 rays, dorsal fin is slightly notched and has 12 spines and 12 rays, and anal fin has 10 or 11 rays. Lateral line scales number 51 to 55. Color is silvery to brassy, occasionally with dark blotches, and with blue line below eye and bluish blotch on upper base of pectoral fin. Scaleless section of head is purplish brown and often with light spots or stripes. This species occurs in coastal waters to 75 m in the western Atlantic from North Carolina and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. It is apparently rare to absent along the Florida gulf coast (R. E. Matheson, pers. comm., August 2002) and in the western Gulf of Mexico. Adults are associated with coral reefs, and juveniles are associated with turtle grass (Thalassia) beds. Food consists of polychaetes, sea urchins, brittle stars, crabs, and hermit crabs. Maximum known size is 410 mm TL. references: Randall and Caldwell 1966; Böhlke and Chaplin 1968; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Smith 1997; Hoese and Moore 1998; Smith-Vaniz et al. 1999; Carpenter 2002b.

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394 PERCIFORMES

Calamus campechanus

Randall and Caldwell, 1966

Calamus campechanus has a small procumbent spine preceding dorsal fin, a relatively short pectoral fin, and a distinct dark spot on anterior section of lateral line. Dorsal profile of head is strongly convex, with portion below eye forming 60 to 68 angle with horizontal line from tip of snout above upper lip to midbase of caudal fin. Posterior naris is long and slitlike. Anterior jaw teeth are conical and about equal in size. Teeth on sides of jaws are in three rows in upper jaw and two rows in lower jaw, and molariform teeth are lacking medial to rows in upper jaw. Lower anterior section of maxilla possesses bony tubercle with free semicircular margin. Tubercle on prefrontal bone, just behind posterior naris, is poorly developed. Gill rakers on first arch are short and number 10 to 12. Measurements are expressed as percent of SL: head length 29%–34%, snout length 17%–22%, eye diameter 7%–9%, suborbital depth 15%–18%, pectoral fin length 29%–33%, length of longest dorsal fin spine 12%–16%, body depth 41%– 46%, caudal pe-

duncle depth 10%–11%. Pectoral fin has 14 to 16 rays, dorsal fin is slightly notched and has 12 or 13 spines and 11 or 12 rays, and anal fin has 9 or 10 rays. Lateral line scales number 45 to 49. Color is light brown or yellowish brown and somewhat darker dorsally than ventrally, with alternating light and dark wavy lines on head below eye, a spot slightly smaller than eye on anterior section of lateral line, and five indistinct vertical rows of dark blotches on sides. Juveniles have more distinct blotches and dark bar on head through eye. This species occurs in coastal waters to a depth of 11 to 18 m in the western Atlantic from Campeche Bay in the Gulf of Mexico and the northeastern Yucatán Peninsula. Maximum known size is 250 mm TL. references: Randall and Caldwell 1966; Hoese and Moore 1997, 1998; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Carpenter 2002b.

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SPARIDAE 395

Calamus leucosteus Whitebone porgy

Jordan and Gilbert, 1885

Calamus leucosteus has a small procumbent spine preceding dorsal fin, a relatively long pectoral fin, and an unpatterned pectoral fin base. Dorsal profile of head is evenly convex, with portion below eye forming 60 to 64 angle with horizontal line from tip of snout above upper lip to midbase of caudal fin. Posterior naris is long and slitlike. Anterior jaw teeth are conical and about equal in size. Teeth on sides of jaws are in three rows in upper jaw and two rows in lower jaw, and molariform teeth are lacking medial to rows in upper jaw. Lower anterior section of maxilla possesses bony tubercle with free semicircular margin. Tubercle on prefrontal bone, just behind posterior naris, is poorly developed. Gill rakers on first arch are short and number 10 to 12. Measurements are expressed as percent of SL: head length 31%–33%, snout length 18%–19%, eye diameter 10%–11%, suborbital depth 12%–13%,

pectoral fin length 29%– 42%, length of longest dorsal fin spine 10%–18%, body depth 44%–54%, caudal peduncle depth 10%–11%. Pectoral fin has 15 to 17 rays, dorsal fin is slightly notched and has 11 to 13 spines and 11 to 13 rays, and anal fin has 9 to 11 rays. Lateral line scales number 44 to 49 (usually 47 or 48). Color is bluish silver, with small irregular blotches. Snout is purplish gray, and blue lines occur above and below eye. This species occurs between 10 and 100 m in the western Atlantic from North Carolina to northern Yucatán, including the Gulf of Mexico. It is associated with soft bottoms. Maximum known size is 460 mm TL. references: Randall and Caldwell 1966; Hoese and Moore 1977, 1998; Boschung 1992; Smith 1997; Castro-Aguirre et al. 1999; Carpenter 2002b.

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396 PERCIFORMES

Calamus nodosus Knobbed porgy

Randall and Caldwell, 1966

Calamus nodosus has a small procumbent spine preceding dorsal fin, suborbital region lightly spotted on a contrasting dark background, and a large blue spot on axil of pectoral fin. Dorsal profile of head is nearly straight, with portion below eye forming 57 to 65 angle with horizontal line from tip of snout above upper lip to midbase of caudal fin. Posterior naris is long and slitlike. Anterior jaw teeth are conical and about equal in size. Teeth on sides of jaws are molariform and in three rows in upper and lower jaws, and an irregular row of molariform teeth is present medial to rows in upper jaw. Lower anterior section of maxilla possesses bony tubercle with free semicircular margin. Tubercle on prefrontal bone, just behind posterior naris, is well developed. Gill rakers on first arch are short and number 10 to 12. Measurements are expressed as percent of SL: head length 30%– 34%, snout length 19%–23%, eye diameter 7%–11%, suborbital depth 14%–18%, pec-

toral fin length 36%–38%, length of longest dorsal fin spine 12%–14%, body depth 47%– 56%, caudal peduncle depth 9%–10%. Pectoral fin has 14 or 15 rays, dorsal fin is slightly notched and has 12 or 13 spines and 11 to 13 rays, and anal fin has 10 or 11 rays. Lateral line scales number 52 to 57 (usually 53 to 55). Color is silvery with a rosy sheen. Snout and cheek are purplish gray, with many bronze spots. This species occurs between 9 and 89 m in the western Atlantic from North Carolina to northern Yucatán, including the Gulf of Mexico. It occurs throughout the Gulf of Mexico. This species is associated with hard bottoms. Maximum known size is 540 mm TL. references: Randall and Caldwell 1966; Bright and Cashman 1974; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Carpenter 2002b.

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SPARIDAE 397

Calamus penna (Valenciennes, 1830) Sheepshead porgy

Calamus penna has a small procumbent spine preceding dorsal fin, a relatively long pectoral fin, and a small prominent black spot on upper part of pectoral fin base. Dorsal profile of head is evenly convex, with portion below eye forming 55 to 62 angle with horizontal line from tip of snout above upper lip to midbase of caudal fin. Posterior naris is long and slitlike. Upper lip of adults is divided in half by lengthwise groove. Anterior jaw teeth are conical and about equal in size. Teeth on sides of jaws are in three rows in upper jaw and two rows in lower jaw, and molariform teeth are lacking medial to rows in upper jaw. Lower anterior section of maxilla possesses bony tubercle without free margin or rarely with short, free, straight to slightly curved margin. Tubercle on prefrontal bone, just behind posterior naris, is poorly developed. Gill rakers on first arch are short and number 10 to 13. Measurements are expressed as percent of SL: head length 30%–33%, snout length 16%–19%, eye diameter 6%–11%, suborbital depth 8%–14%, pectoral fin length 29%–39%, length of longest dorsal fin spine

10%–14%, body depth 39%–50%, caudal peduncle depth 9%–10%. Pectoral fin has 14 to 16 rays, dorsal fin is slightly notched and has 11 to 13 spines and 11 to 13 rays, and anal fin has 9 to 11 rays. Lateral line scales number 45 to 49. Color is silvery, with iridescent reflections and diffuse longitudinal grayish stripes. Cheek is silvery and tinged with brown, and a brown vertical bar passes from lower rim of eye to corner of mouth. Juveniles have dark gray bars. This species occurs between 3 and 87 m in the western Atlantic from Florida and the Bahamas to Brazil, including the eastern Gulf of Mexico and the Antilles. It is rare along the Florida gulf coast (R. E. Matheson, pers. comm., August 2002). It is associated with hard and semihard bottoms. Maximum known size is 460 mm TL. references: Randall and Caldwell 1966; Böhlke and Chaplin 1968; Matsuura 1983f; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Carpenter 2002b.

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398 PERCIFORMES

Calamus pennatula

Guichenot, 1868

Calamus pennatula has a small procumbent spine preceding dorsal fin, an enlarged canine tooth in upper jaw, and a moderately steep dorsal profile of head. Dorsal profile of head is evenly convex, with portion below eye forming 57 to 64 angle with horizontal line from tip of snout above upper lip to midbase of caudal fin. Posterior naris is long and slitlike. Anterior jaw teeth are conical, with fourth tooth from symphysis enlarged (in specimens greater than 100 mm SL) and strongly outcurved (in specimens greater than 170 mm SL). Teeth on sides of jaws are molariform and in three rows in upper and lower jaws, and an irregular row of molariform teeth is present medial to rows in upper jaw. Lower anterior section of maxilla possesses bony tubercle with free semicircular margin. Tubercle on prefrontal bone, just behind posterior naris, is poorly developed. Gill rakers on first arch are short and number 9 to 13. Measurements are expressed as percent of SL: head length 30%–34%, snout length 19%–23%, eye diameter 7%–11%, suborbital depth 14%–18%, pectoral fin length 33%–

42%, length of longest dorsal fin spine 12%– 15%, body depth 42%–53%, caudal peduncle depth 8%– 8%. Pectoral fin has 13 to 15 rays, dorsal fin is slightly notched and has 12 spines and 11 to 13 rays, and anal fin has 9 to 11 rays. Lateral line scales number 51 to 56 (usually 53 to 55). Color is silvery gray to tan, often with a dusky spot at center of each scale on upper part of body. Suborbital region is patterned with alternating narrow blue and yellow stripes, a rectangular blue blotch is located behind eye, and an iridescent blue area and a small orange spot are located on upper part of pectoral fin base. This species occurs between 5 and 84 m in the western Atlantic from northern Yucatán to Brazil, including the southern Gulf of Mexico, the Bahamas, and the Antilles. Maximum known size is 370 mm TL. references: Randall and Caldwell 1966; Böhlke and Chaplin 1968; Matsuura 1983f; Cervigón 1993a,b; Randall 1996; Smith 1997; Carpenter 2002b.

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SPARIDAE 399

Calamus proridens Littlehead porgy

Jordan and Gilbert, 1884

Calamus proridens has a small procumbent spine preceding dorsal fin, an elongate dark blotch over upper end of gill opening, and a very steep dorsal profile of upper part of head. Dorsal profile of head is evenly convex, with portion below eye forming 57 to 64 angle with horizontal line from tip of snout above upper lip to midbase of caudal fin. Posterior naris is long and slitlike. Anterior jaw teeth are conical, with fourth tooth from symphysis enlarged (in specimens greater than 100 mm SL) and strongly outcurved (in specimens greater than 148 mm SL). Teeth on sides of jaws are molariform and in three rows in upper and lower jaws, and an irregular row of molariform teeth is present medial to rows in upper jaw. Lower anterior section of maxilla possesses bony tubercle with free semicircular margin. Tubercle on prefrontal bone, just behind posterior naris, is poorly developed. Gill rakers on first arch are short and number 11 to 13 (usually 11). Measurements are expressed as percent of SL: head length 29%–32%, snout length 16%–20%, eye diameter 7%– 8%, sub-

orbital depth 10%–16%, pectoral fin length 36%– 40%, length of longest dorsal fin spine 14%–16%, body depth 46%–51%, caudal peduncle depth 9%–10%. Pectoral fin has 13 to 15 (usually 14) rays, dorsal fin is slightly notched and has 12 or 13 spines and 11 or 12 rays, and anal fin has 10 or 11 rays. Lateral line scales number 52 to 57. Color is tan to light brown, with a dark bluish spot at center of each scale on upper part of body forming a narrow line along each scale row. Suborbital region is patterned with wavy dark blue lines, with area between lines often brassy. This species occurs between 11 and 60 m in the western Atlantic from northeastern Florida to northern Yucatán, including the Gulf of Mexico, Cuba, and Hispaniola. It occurs in the eastern and southern Gulf of Mexico off the Yucatán Peninsula. Maximum known size is 460 mm TL.

references: Randall and Caldwell 1966; Robins and Ray 1986; Smith 1997; Carpenter 2002b.

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400 PERCIFORMES

Diplodus argenteus Silver pinfish

(Valenciennes, 1830)

Diplodus argenteus is oval in profile, with a moderate-sized black spot on caudal peduncle. Dorsal profile of head is nearly straight to posterior margin of eye. Posterior naris is rounded. Maxilla barely extends to anterior margin of eye. Anterior eight teeth in both jaws are broad, incisor-like, and unnotched. Teeth in sides of jaws are molariform and arranged in three rows. Gill rakers on first arch are short and number 17 to 20. Measurements are expressed as percent of SL: head length 31%–36%, snout length 9%–11%, eye diameter 10%–12%, suborbital depth about 7%, pectoral fin length 27%–33%, length of second anal fin spine 16%–17%, body depth 45%– 49%, caudal peduncle depth 11%–14%. Pectoral fin has 15 to 17 rays. Dorsal fin is not notched and has 12 spines and 12 to 14 rays. Anal fin has 11 to

14 rays. Color is silvery, with a pale yellow stripe along each scale row and about nine narrow dusky bars on side. Black spot or saddle on caudal peduncle is largely limited to area above lateral line. Fins are yellow. This species occurs in shallow coastal waters in the western Atlantic from southern Florida and the Bahamas to Argentina, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs off southwestern Florida. It is associated with clear water over rocky or coral bottoms. Food consists of algae, mollusks, and crabs. Maximum known size is 280 mm TL.

references: Robins and Ray 1986; Cervigón 1993b; Smith 1997; Carpenter 2002b.

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SPARIDAE 401

Diplodus holbrookii (Bean, 1878) Spottail pinfish

Diplodus holbrookii is oval in profile and compressed, with a large black spot on caudal peduncle. Dorsal profile of head is nearly straight to posterior margin of eye. Posterior naris is rounded. Maxilla barely extends to anterior margin of eye. Anterior six teeth in both jaws are incisor-like, broad, and unnotched. Teeth in sides of jaws are molariform and arranged in three rows. Gill rakers on first arch are short and number 17 to 21. Measurements are expressed as percent of SL: head length 31%– 32%, snout length 10%–11%, eye diameter 9%–10%, suborbital depth 4%–5%, pectoral fin length 29%–31%, second anal fin spine length 14%–17%, body depth 46%– 48%, caudal peduncle depth 11%–12%. Pectoral fin has 15 to 17 rays. Dorsal fin is unnotched and has 11 or 12 spines and 12 to 16 rays. Anal fin

has 12 to 15 rays. Color is bluish brown dorsally and silvery on side. Black spot on caudal peduncle extends nearly to ventral midline or forms complete ring. Juveniles have narrow dark bars on body. This species occurs in coastal waters to 28 m in the western Atlantic from Chesapeake Bay to the northwestern Gulf of Mexico. It is benthic and associated with submerged vegetation but is rare in brackish water. Food consists of small benthic invertebrates. Spawning takes place in winter. Maximum known size is 460 mm TL. references: Caldwell 1955b; Hastings et al. 1976; Hoese and Moore 1977, 1998; Hastings 1979; Robins and Ray 1986; Boschung 1992; Smith 1997; Carpenter 2002b.

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402 PERCIFORMES

Lagodon rhomboides Pinfish

(Linnaeus, 1766)

Lagodon rhomboides is oval in profile, with a small procumbent spine preceding dorsal fin and a dark spot centered on lateral line. Dorsal profile of head is slightly convex to anterior margin of eye. Posterior naris is oval shaped. Four anterior teeth in jaws are incisor-like, notched, and forwardly directed. Lateral teeth in jaws are molariform and in two and onehalf rows. Gill rakers on first arch are of moderate length and number 6 to 9 on upper limb and 10 to 15 on lower limb. Measurements are expressed as percent of SL: head length 30%– 33%, snout length 10%–11%, eye diameter 7%– 8%, suborbital depth 4%–5%, pectoral fin length 32%–36%, second anal fin spine length 10%–13%, body depth 36%– 43%, caudal peduncle depth 9%–11%. Pectoral fin has 16 rays, dorsal fin has 11 to 13 spines and 10 to 12 rays, and anal fin has 10 or 11 rays. Lateral line scales number 65 to 70. Color is

silvery, with yellow stripes broader than interspaces, blackish spot near origin of lateral line, and six broad vertical bars on side. Pectoral and caudal fins are yellow, and anal fin is yellow with a broad blue margin. This species occurs in shallow water in the western Atlantic from Massachusetts and Bermuda to the southern Gulf of Mexico, Quintana Roo, and Cuba. It is benthic and associated with seaweeds and occasionally mangroves and rocky bottoms. It also enters brackish and fresh waters. Food consists of benthic invertebrates and ray-finned fishes. Maximum known size is 350 mm TL. references: Caldwell 1957; Hastings et al. 1976; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Carpenter 2002b.

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SPARIDAE 403

Pagrus pagrus (Linnaeus, 1758) Red porgy

Pagrus pagrus has a shallow suborbital space and small spots on body. Dorsal profile of head is strongly convex. Posterior naris is oblong and larger than anterior naris. Anterior two teeth in upper jaw and four teeth in lower jaw are caninelike. Teeth on sides of jaws are molariform and arranged in two rows. Gill rakers on first arch are short and number 6 or 7 on upper limb and 9 or 10 on lower limb. Measurements are expressed as percent of SL: head length 31%–34%, snout length 12%–13%, eye diameter 9%–10%, suborbital depth 10%– 12%, pectoral fin length 33%–35%, length of longest dorsal fin spine 11%–14%, body depth 39%– 40%, caudal peduncle depth about 9%. Pectoral fin has 15 or 16 rays, dorsal fin is slightly notched and has 12 or 13 spines and 9

to 11 rays, and anal fin has 7 or 8 rays. Lateral line scales number 51 to 59. Color is silvery red, with very small blue or yellow spots. Pectoral, dorsal, and caudal fins are pink. This species occurs in the Atlantic Ocean in tropical to warm temperate seas. In the western Atlantic it occurs between 10 and 80 m from New York to Argentina, including the entire Gulf of Mexico. It is associated with rocky and hard sand bottoms. Maximum known size is 910 mm TL. references: Manooch et al. 1976; Hoese and Moore 1977, 1998; Manooch and Hassler 1978; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Carpenter 2002b.

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404 PERCIFORMES

Stenotomus caprinus Longspine porgy

Jordan and Gilbert, 1882

Stenotomus caprinus is oval in profile, compressed, and deep bodied, with a pointed snout; a procumbent spine preceding dorsal fin; and the second, third, and fourth dorsal fin spines greatly elongated. Dorsal profile of head is nearly straight to eye, slightly convex over eye, and slightly convex on upper nape. Posterior naris is elongate and slitlike. Maxilla fails to reach anterior margin of eye. Anterior jaw teeth are arranged in two close-set bands, and those of outer band are narrow and have spatulate crowns. Teeth in sides of jaws are molariform and arranged in two rows. Gill rakers on first arch are short and number 15 to 18. Measurements are expressed as percent of SL: head length 30%–34%, snout length 13%–16%, eye diameter 9%–10%, suborbital depth 9%– 11%, pectoral fin length 37%– 41%, second

anal fin spine length 11%–14%, body depth 52%– 62%, caudal peduncle depth 10%–12%. Pectoral fin has 15 or 16 rays, dorsal fin is unnotched and has 11 or 12 spines and 11 to 13 rays, and anal fin has 11 or 12 (usually 11) rays. Lateral line scales number 46 to 50. Color is silvery and slightly olivaceous on dorsal surface. Juveniles have faint dark bars. This species occurs between 5 and 120 m in the western Atlantic from the east coast of Florida to northern Yucatán, including the entire Gulf of Mexico. It is associated with mud bottoms. Maximum known size is 300 mm TL. references: Caldwell 1955a; Hastings et al. 1976; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Carpenter 2002b.

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405

POLYNEMIDAE

Threadfins

Threadfins are oblong to elongate, and moderately compressed, with an inferior mouth, lower pectoral fin rays separate and threadlike, and two widely separated dorsal fins. Snout is conical and projects beyond mouth. Nostril is paired, and nares are close set. Eye is laterally located and has well-developed adipose eyelid. Mouth is large, premaxilla is slightly protractile, and maxilla is expanded posteriorly and extends far beyond eye. Supramaxilla is absent. Jaw and palatine teeth are cardiform and arranged in bands. Vomerine teeth are villiform when present. Preopercular margin is strongly angular or lobate and serrated along dorsal margin. Opercular margin is strongly rounded or angular. Gill membranes are narrowly joined at isthmus, and branchiostegal rays number 7. Pectoral fin is low on flank and divided into two sections, with 12 to 19 principal rays and 3 to 16 lower, elongated, and unbranched rays that are free of membranes. First dorsal fin is triangular and has seven or eight weak spines. Second dorsal fin is triangular and has one spine and 9 to 18 rays. Pelvic fin is subabdominal (located posterior to pectoral fin base under anterior section of abdomen) and has one spine and 5 rays. Anal fin has two or three spines and 9 to 30 rays. Caudal fin is deeply forked. Body, head, and fin bases are covered with weakly ctenoid scales. Median fins are covered with small ctenoid scales. Lateral line is complete to margin of caudal fin. Large axillary process is located on pelvic fin base. Gas bladder is present and is contained in body cavity. Vertebrae number 24 or 25. Maximum known size is 460 cm TL. Threadfins occur worldwide in tropical to warm temperate shallow waters. They are associated with sandy and muddy bottoms, often occur in turbid waters, and are common in brackish and occasionally fresh waters with connections to the sea. Food consists of crustaceans and small ray-finned fishes. There are about 33 species in seven genera, 3 species in the same genus in the western Central Atlantic, and 2 or 3 of these in the Gulf of Mexico. references: Smith 1986; Nelson 1994; Feltes 2001, 2002.

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406 PERCIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Feltes 2002) 1a. Eight filamentous lower pectoral fin rays . . . . . . . . . . . . . . . . . . . . . Polydactylus octonemus p. 407 1b. Seven filamentous lower pectoral fin rays . . . . . . . . . . . . . 2 2a. Posterior margin of maxilla rounded; anal fin base 21% to 23% of SL; lateral line scales 67 to 73 . . . . . . . . . . . . . . . . . . . . . . . Polydactylus oligodon p. 408 2b. Posterior margin of maxilla truncate or concave; anal fin base 17% to 19% of SL; lateral line scales 54 to 63 . . . . . . . . . . . . . . . . . . . . . . Polydactylus virginicus p. 409

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POLYNEMIDAE 407

Polydactylus octonemus Atlantic threadfin

(Girard, 1858)

Polydactylus octonemus has 8 detached rays in lower section of pectoral fin. Maxilla extends well beyond eye and has truncate posterior margin. Teeth in upper jaw are exposed when mouth is closed. Vomer has villiform teeth. Margin of preoperculum is serrated, with a large, flat spine and a membranous flap at corner. Opercular margin is broadly rounded and entire. Gill rakers on first arch number 34 to 38. Measurements are expressed as percent of SL: head length 29%–31%, snout length 5%–7%, eye diameter 5%– 6%, upper jaw length 12%–13%, upper section of pectoral fin length 23%–27%, anal fin base 18%–21%, body depth 28%–30%, caudal peduncle depth 13%–15%. Pectoral fin has 14 to 16 rays in upper section and 8 free, filamentous rays in lower section. First dorsal fin has eight spines, and second dorsal fin has one spine and 11 or

13 rays. Anal fin has three spines and 12 to 14 rays. Lateral line scales number 56 to 64. Color is dull silvery, darker dorsally, and grading to whitish ventrally. Pectoral fin is black, and dorsal and anal fins are dusky yellow and black distally. This species occurs between 5 and 22 m in the western Atlantic from New York to the southern Gulf of Mexico. It occurs throughout Gulf of Mexico. It is associated with sandy and muddy bottoms in shallow water. Maximum known size is 230 mm TL. references: Hoese and Moore 1977, 1998; Martin and Drewry 1978; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Dentzau and Chittenden 1990; Boschung 1992; Schaldach et al. 1997; Smith 1997; Feltes 2002.

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408 PERCIFORMES

Polydactylus oligodon Littlescale threadfin

(Günther, 1860)

Polydactylus oligodon has 7 detached rays in lower section of pectoral fin. Upper lip is absent, and lower lip is fully developed. Teeth in upper jaw are exposed when mouth is closed. Vomer has villiform teeth. Margin of preoperculum is serrated, with a large, flat spine and a membranous flap at corner. Opercular margin is broadly rounded and entire. Gill rakers on first arch number 22 to 30. Measurements are expressed as percent of SL: head length 31%– 33%, snout length 6%–7%, eye diameter 8%– 9%, upper jaw length 14%–15%, upper section of pectoral fin length 23%–25%, anal fin base 21%–23%, body depth 26%–29%, caudal peduncle depth 13%–14%. Pectoral fin has 15 or 16 rays in upper section and 7 in lower section. First dorsal fin has eight spines, and

second dorsal fin has one spine and 11 or 12 rays. Anal fin has three spines and 13 to 15 rays. Lateral line scales number 67 to 73. Color is dull silvery dorsally and lighter ventrally, with edges of dorsal and caudal fins blackish near tip, and borders of anal fin pale. This species occurs in the western Atlantic from southern Florida and the Bahamas to northern Brazil, including Jamaica and Trinidad. There is a questionable record of this species from the Laguna de Términos, Campeche, in the Gulf of Mexico. This species is associated with sandy and muddy bottoms in shallow water. Maximum known size is 460 mm TL. references: Randall 1966a, 1996; Böhlke and Chaplin 1968; Smith 1997; Feltes 2002.

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POLYNEMIDAE 409

Polydactylus virginicus Barbu

(Linnaeus, 1758)

Polydactylus virginicus has 7 detached rays in lower section of pectoral fin. Upper lip is absent, and lower lip is fully developed. Teeth in upper jaw are exposed when mouth is closed. Vomer has villiform teeth. Margin of preoperculum is serrated, with a large, flat spine and a membranous flap at corner. Opercular margin is broadly rounded and entire. Gill rakers on first arch number 26 to 30, with 11 or 13 on upper limb. Measurements are expressed as percent of SL: head length 30%– 31%, snout length 5%–7%, eye diameter 7%– 8%, upper jaw length 13%–14%, pectoral fin length 22%–26%, anal fin base 17%–19%, body depth 27%–33%, caudal peduncle depth 13%–15%. Pectoral fin has 14 to 16 rays in upper section and 7 in lower section. First dorsal fin has eight spines, and second dorsal fin

has one spine and 11 or 12 rays. Anal fin has three spines and 11 to 14 rays. Lateral line scales number 54 to 63. Color is dull silvery, with a yellowish cast. First dorsal fin is blackish near tip, and paired fins are often blackish near centers. This species occurs in the western Atlantic from New Jersey (rare) and Bermuda to Uruguay, including the southern Gulf of Mexico, the Florida Keys, and the Antilles. In the Gulf of Mexico it is known from Progreso, Yucatán. This species is associated with sandy and muddy bottoms in shallow water. Maximum known size is 330 mm TL.

references: Kanazawa 1952; Randall 1996; Smith 1997; Smith-Vaniz et al. 1999; Feltes 2002.

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410

SCIAENIDAE

Drums

Drums are oblong to moderately elongate and moderately compressed, with conspicuous pores on snout and chin, and lateral line extending to margin of caudal fin. Head usually has bony ridges on top of cranium and large cavernous canals that are part of lateral line system. Snout is rounded to slightly pointed. Nostril is paired, with nares generally closer to anterior margin of orbit than to tip of snout. Eye is small to moderate in size. Mouth is terminal, subterminal, or slightly superior; small to rather large; and nearly horizontal to moderately oblique. Jaw teeth are usually small, conical, and arranged in bands. Occasionally canine teeth occur at tips of jaws. Vomer and palatine lack teeth. Snout possesses three to seven rostral pores along anterior margin, and two to five marginal pores along edge of rostral fold. Chin has two to six mental pores, one pore at midline, and remaining pores paired and located along each side. Barbels may be associated with pores on chin and may also occur on suboperculum. Preopercular margin is entire or weakly to strongly serrate. Operculum has two broad spines along upper posterior margin that are connected by ridge. Gill membranes are free of isthmus, and last full arch (fourth) has slit posteriorly. Branchiostegal rays number 7. Pectoral fin inserts on lower half of flank, has a nearly vertical base, and has 15 to 25 rays. Dorsal fin is long and single, with a deep notch between spinous and rayed sections in most genera but separated into separate spinous and rayed sections in one genus. Spinous section of dorsal fin has 7 to 13 (usually 10) spines, and rayed section has 1 to 4 spines and 18 to 46 rays. Pelvic fin inserts below or slightly posterior to pectoral fin insertion and consists of 1 spine and 5 rays. Anal fin has 1 or 2 spines and 6 to 20 rays. Caudal fin is slightly emarginate, truncate, rounded, or pointed and consists of 17 principal rays. Ctenoid or cycloid scales cover body and part of head. In some species, scales are ctenoid on body and cycloid on head and breast. Small intercalated scales are often associated with lateral line. Scaly sheaths cover bases of dorsal and anal fins in some species. Vertebrae range from 22 to 29 but usually number 25. Gas bladder is present, is highly modified in some species, and is associated with muscles that cause it to vibrate and produce drumming or croaking sounds. Drums occur worldwide in tropical to warm temperate seas between the shoreline and about 600 m. Most species live in coastal marine habitats, many species are associated seasonally or yearround with estuaries, and some species are limited to freshwater. Many species utilize estuaries as nursery grounds. Most species are associated with sandy and muddy bottoms in the vicinity of river mouths; others are found on coral reefs and in surf zones. Food consists of benthic invertebrates and ray-finned fishes. Eggs

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SCIAENIDAE 411

and larvae are pelagic. Maximum known size is 200 cm TL. There are about 270 species in 70 genera, 55 species in 22 genera in the western Central Atlantic, and 25 to 29 species in 14 or 15 genera in the Gulf of Mexico. references: Chao 1986, 1995, 2002; Johnson 1984; Heemstra 1986g; Sasaki 1989; Nelson 1994; Smith 1997.

Key to the Species of the Gulf of Mexico (Adapted from Chao 2002; R. E. Matheson, pers. comm., August 2002) 1a. 1b. 2a. 2b. 3a. 3b. 4a.

4b.

5a.

5b.

6a.

6b.

7a.

Chin with barbels. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Chin without barbels . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Single barbel at tip of lower jaw . . . . . . . . . . . . . . . . . . . . 3 Two or more barbels on lower jaw. . . . . . . . . . . . . . . . . . 6 Anal fin with two spines; body oblong and compressed . . . . . . . . . . . . . . . . . . . . . . . . . . Umbrina coroides p. 446 Anal fin with one spine; body elongate and oval in cross section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Scales on breast not uniform in size, anterior scales smaller than scales along lateral line; pectoral fin not reaching beyond tip of pelvic fin; adults with three or more gill rakers on lower limb of first gill arch . . . . . . . . . . . . . . . . . . . . . . . Menticirrhus littoralis p. 432 Scales on breast uniform in size, anterior scales same size as those of lateral line; pectoral fin reaching beyond tip of pelvic fin; adults with tuberculate or no gill rakers on lower limb of first gill arch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Anal fin rays 6 to 8 (usually 7); depressed spinous dorsal fin seldom extends beyond base of second or third dorsal fin ray; longest dorsal fin spine 16% to 24% of SL . . . . . . . . . . . . . . . . . . . . . Menticirrhus americanus p. 431 Anal fin rays 7 to 9 (usually 8); depressed spinous dorsal fin often extends beyond base of second or third dorsal fin ray; longest dorsal fin spine 25% to 39% of SL . . . . . . . . . . . . . . . . . . . . . . . Menticirrhus saxatilis p. 433 Three pairs of barbels arranged in tuft at tip of lower jaw; more than 10 pairs of barbels on rami of lower jaw; eye diameter about 11% of head length; second anal fin spine less than half the length of first anal fin ray . . . . . . . . . . . . . . . . . . . . Paralonchurus brasiliensis p. 437 No tuft of barbels on tip of lower jaw; fewer than 10 barbels on rami of lower jaw; eye diameter about 20% of head length; second anal fin spine longer than two-thirds the length of first anal fin ray . . . . . . . . . . . . . . . . . . . . . . . . . 7 Preopercular margin strongly serrate; scales in horizontal series 64 to 72; body silvery with many oblique stripes .............................................8

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412 PERCIFORMES

7b. Preopercular margin entire; scales in horizontal series 41 to 45; body grayish to dark or with black bars . . . . . . . . . . . . . . . . . . . . . . . . . . . Pogonias cromis p. 442 8a. Dark spots on scales on side of body usually arranged in parallel or wavy lines, most distinct below rayed section of dorsal fin; scales between dorsal fin origin and lateral line number six or seven; dorsal fin rays usually 26 or 27 . . . . . . . . . . . . . . . . . . . . . . Micropogonias furnieri p. 434 8b. Dark spots on scales on side of body not arranged in parallel or wavy lines; scales between dorsal fin origin and lateral line number eight or nine; dorsal fin rays usually 28 or 29 . . . . . . . . . . . . . . . Micropogonias undulatus p. 435 9a. Mouth strongly oblique; lower jaw projecting beyond upper jaw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9b. Mouth not strongly oblique; tip of lower jaw usually not projecting beyond upper jaw . . . . . . . . . . . . . . . . . . . . . 16 10a. Body depth greater than 29% of TL; jaw teeth minute and in one or two rows; first gill arch with more than 28 rakers; about 50 lateral scale rows. . . . . . . . . . . . . . . . . . . . 11 10b. Body depth less than 25% of TL; jaw teeth conical, arranged in narrow bands, usually with large canine tooth at tip of upper jaw; first gill arch with fewer than 20 rakers; more than 80 lateral scale rows . . . . . . . . . . . . . . . . 12 11a. Body uniformly dark gray dorsally and silvery ventrally; gill rakers on first arch longer than eye diameter and number 28 to 33 . . . . . . . . . . . . . . . . Larimus breviceps p. 428 11b. Body dark, with seven to nine bars on side; gill rakers on first arch equal to eye diameter and number 34 to 36 . . . . . . . . . . . . . . . . . . . . . . . . . . Larimus fasciatus p. 429 12a. Body and rayed section of dorsal fin lack large dark spots, but oblique rows of small dark spots may be present . . . 13 12b. Body and rayed section of dorsal fin covered with large dark spots. . . . . . . . . . . . . . . . Cynoscion nebulosus p. 421 13a. Side of body with small dark spots forming distinct oblique lines above lateral line; usually 11 or more gill rakers on lower limb of first arch. . . . . . . . . Cynoscion regalis p. 423 13b. Side of body without distinct dark spots but occasionally with faint streaks above lateral line; usually 11 or fewer gill rakers on lower limb of first arch . . . . . . . . . . . . . . . . . . 14 14a. Anal fin with 10 to 12 (usually 11) rays . . . . . . . . . . . . . . . . . . . . . . . . Cynoscion arenarius p. 419 14b. Anal fin with 8 to 10 (usually 9) rays . . . . . . . . . . . . . . . 15 15a. Basal three-fourths or more of rayed section of dorsal fin covered with small scales; dorsal fin rays 23 to 27; gill rakers on first arch 9 to 13 . . . . Cynoscion jamaicensis p. 420 15b. Basal two-thirds or less of rayed section of dorsal fin covered with small scales; dorsal fin rays 26 to 31; gill rakers on first arch 11 to 14. . . . . . . . . . Cynoscion nothus p. 422 16a. More than 36 dorsal fin rays . . . . . . . . . . . . . . . . . . . . . 17

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SCIAENIDAE 413

16b. Fewer than 36 dorsal fin rays (fewer than 30 in all species except Leiostomus xanthurus) . . . . . . . . . . . . . . . . . . . . 21 17a. More than 44 dorsal fin rays; bold, black oblique bars through eye, from nape to breast and from spinous dorsal fin to or onto base of caudal fin . . . . . . . . . . . . . . . . . . . 18 17b. Fewer than 42 dorsal fin rays; bars not as above (17a), but stripes may be present . . . . . . . . . . . . . . . . . . . . . . . . . . 19 18a. Body with two narrow stripes on either side of third oblique bar; second dorsal fin, anal fin, and caudal fin dark brown with white spots; pectoral fin dark brown; 45 to 47 dorsal fin rays . . . . . . . . . . . . . . . . . . . . Equetus punctatus p. 426 18b. Body without narrow stripes on either side of third oblique bar; second dorsal fin and anal fin pale without white spots; caudal fin with black stripe but without white spots; pectoral fin pale; 47 to 55 dorsal fin rays . . . . . . . . . . . . . . . . . . . . . . . . . Equetus lanceolatus p. 424 19a. Body with broad oblique bar running from base of spinous section of dorsal fin to pelvic fin; one longitudinal stripe along midline of side, reaching to near tip of caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . Pareques iwamotoi p. 440 19b. Body without broad oblique bar; usually with several longitudinal stripes on side, although these may be obscured by dark background pigment . . . . . . . . . . . . . . . . . . . . . 20 20a. Adults and larger juveniles with bold lateral pigment pattern of dark stripes on light background, some dark stripes as wide as pupil; juveniles with straight dark bar across top of head between orbits . . . Pareques acuminatus p. 438 20b. Adults and larger juveniles with relatively obscure dark lateral stripes on dark background, dark stripes narrower than pupil; juveniles with anteriorly directed V-shaped bar connecting orbits on top of head . . . . . . . . . . . . . . . . . . . . . . . . . Pareques umbrosus p. 441 21a. More than 11 anal fin rays; 29 or more dorsal fin rays; dark shoulder spot present . . . . . . Leiostomus xanthurus p. 430 21b. Fewer than 11 anal fin rays; 29 or fewer dorsal fin rays; no dark shoulder spot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22a. Seven to 9 gill rakers on lower limb of first arch; one or more black ocellated spots on rear portion of body, commonly with a single spot larger than pupil at upper base of caudal fin . . . . . . . . . . . . . . . . . . Sciaenops ocellata p. 443 22b. Thirteen to 23 gill rakers on lower limb of first arch; pigmentation not as above (22a) . . . . . . . . . . . . . . . . . . . . . 23 23a. Large black blotch at base of pectoral fin; pair of enlarged canines near tip of lower jaw . . . . . . . . . . . . . . . . . . . . . . . . Odontoscion dentex p. 436 23b. No large black blotch at base of pectoral fin (a faint dark spot is present near origin of pectoral fin in Corvula sanctaeluciae); without pair of enlarged canines near tip of lower jaw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

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414 PERCIFORMES

24a. Twenty-two or 23 gill rakers on lower limb of first arch; conspicuous cavernous canals on top of head; profile from spinous dorsal fin to snout strongly concave. . . . . . . . . . 25 24b. Thirteen to 18 gill rakers on lower limb of first arch; without cavernous canals on top of head; profile from spinous dorsal fin to snout nearly straight to slightly concave . . . 26 25a. Head extremely cavernous, spongy; dorsal extreme of upper lip on a level with or above lower margin of eye; snout usually not projecting beyond upper lip; gill rakers on first arch 10 to 13 on upper limb and 22 or 23 on lower limb (total 32 to 36); eye diameter 18% to 22% of head length . . . . . . . . . . . . . . . . . . . . . . . . . Stellifer lanceolatus p. 445 25b. Head cavernous but not spongy; dorsal extreme of upper lip on a level with suborbital section of head (below level of eye); snout projecting slightly beyond upper lip; gill rakers on first arch 10 to 12 on upper limb and 19 to 22 on lower limb (total 29 to 34); eye diameter 16% to 18% of head length. . . . . . . . . . . . . . . . . Stellifer colonensis p. 444 26a. Preopercular margin with distinct strong spines at angle; length of second anal fin spine nearly as long as first anal fin ray. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 26b. Preopercular margin with weak serrations, no spines at angle; length of second anal fin spine less than two-thirds the length of first anal fin ray . . . . . . . . . . . . . . . . . . . . . 28 27a. Usually with 23 to 25 (rarely 21 or 22) dorsal fin rays; second anal fin spine very strong, with length about equal to that of first anal fin ray . . . . . . . . Bairdiella ronchus p. 416 27b. Nineteen to 23 dorsal fin rays; second anal fin spine not as strong as and somewhat shorter than first anal fin ray . . . . . . . . . . . . . . . . . . . . . . . . Bairdiella chrysoura p. 415 28a. Mouth slightly inferior; distinct stripes on side; dorsal fin rays 25 to 29; anal fin rays 7 or 8; second anal fin spine 33% to 40% of head length; gill rakers on first gill arch 18 to 22 . . . . . . . . . . . . . . . . . . . . . . Corvula batabana p. 417 28b. Mouth terminal; faint stripes on side; dorsal fin rays 22 to 24; anal fin rays 9 (rarely 8); second anal fin spine 28% to 31% of head length; gill rakers on first gill arch 23 to 25 . . . . . . . . . . . . . . . . . . . . . . . Corvula sanctaeluciae p. 418

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SCIAENIDAE 415

Bairdiella chrysoura (Lacepède, 1802) Silver perch

Bairdiella chrysoura is oblong in profile, with a terminal, slightly oblique mouth and a truncate to double emarginate caudal fin. Dorsal profile is evenly convex. Posterior naris is oval shaped. Maxilla extends to about center of pupil. Snout has three rostral and five marginal pores, and chin has six mental pores, with median pair often in pit. Jaw teeth are small, conical, and arranged in a band in upper jaw and in a single series in lower jaw. Preopercular margin has several spines at angle. Gill rakers on first arch are moderately long and slender and number 22 to 25. Measurements are expressed as percent of SL: head length 31%–36%, snout length 8%–9%, interorbital distance 6%–7%, eye diameter 7%–9%, upper jaw length 13%– 15%, pectoral fin length 18%–22%, body depth 30%–33%. Pectoral fin has 15 to 17 rays. Dorsal fin is deeply notched and has 10 or 11 spines in anterior section and 1 spine and 19 to 23 rays in posterior section. Anal fin has 2 spines, with second spine stout and about two-thirds the length of first ray, and 8 to 10 rays. Body is covered with ctenoid scales, and

most of head is covered with cycloid scales. Basal parts of rayed sections of dorsal and anal fins are scaled. Gas bladder has yoke-shaped anterior chamber and carrot-shaped posterior chamber but lacks appendages. Color is silvery and olivaceous dorsally, with obscure dark stripes along scale rows on side, and bright silvery to yellowish ventrally. Dorsal, pelvic, anal, and caudal fins are pale yellow. This species occurs in coastal waters in the western Atlantic from Cape Cod to southern Florida and throughout the Gulf of Mexico. It is associated with both sandy and muddy bottoms and utilizes estuaries and occasionally fresh waters as nursery grounds. Food consists of crustaceans, polychaetes, and, occasionally, small ray-finned fishes. Maximum known size is 300 mm TL. references: Hoese and Moore 1977, 1998; Chao 1978, 2002; Johnson 1978; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999.

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416 PERCIFORMES

Bairdiella ronchus (Cuvier, 1830)

Bairdiella ronchus is oblong to slightly elongate in profile, with a strongly oblique, slightly subterminal mouth; a truncate to slightly rounded caudal fin; and a body patterned with faint dark streaks. Dorsal profile is nearly straight. Posterior naris is oval shaped. Maxilla extends to about center of pupil. Snout has three rostral pores and five marginal pores, and chin has five mental pores. Jaw teeth are small, conical, and arranged in bands in both jaws, with outer row in upper jaw and inner row in lower jaw enlarged. Preopercular margin has several strong spines at angle. Gill rakers on first arch are moderately long and slender and number 21 to 27 (usually 24 or 25). Measurements are expressed as percent of SL: head length 29%–32%, snout length 7%– 8%, interorbital distance 6%–7%, eye diameter 6%–7%, upper jaw length 12%–13%, pectoral fin length 20%–22%, body depth 28%–30%, caudal peduncle depth 9%–10%. Pectoral fin has 16 or 17 rays. Dorsal fin is deeply notched and has 10 (rarely 11) spines in anterior section and 1 spine and 21 to 26 (usually 23 to 25) rays in posterior section. Anal fin has 2 spines, with

second spine stout and about same length as first anal ray, and 7 to 9 (usually 8) rays. Body and top of head are covered with ctenoid scales, and cheeks are covered with cycloid scales. Basal parts of rayed sections of dorsal and anal fins are scaled. Gas bladder has yoke-shaped anterior chamber and carrot-shaped posterior chamber but lacks appendages. Color is grayish dorsally and silvery ventrally. Streaks on flanks are oblique above lateral line and longitudinal below lateral line. Dorsal and caudal fins are grayish with black margin, and anal fin is covered with minute dark specks. This species occurs between 16 and 40 m in the western Atlantic from the western and southern Gulf of Mexico to southeastern Brazil, including the Greater and Lesser Antilles. It is associated with muddy and sandy bottoms and brackish estuaries. Food consists of crustaceans and small ray-finned fishes. Maximum known size is 350 mm TL. references: Darnell 1962; Chao 1978, 2002; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999.

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SCIAENIDAE 417

Corvula batabana Blue croaker

(Poey, 1860)

Corvula batabana is oblong in profile, with a moderately oblique, subterminal mouth; a convex caudal fin; and a body patterned with stripes below lateral line. Dorsal profile is nearly straight to dorsal fin origin. Posterior naris is oval shaped. Maxilla extends to about center of pupil. Snout has three rostral and five marginal pores, and chin has five mental pores. Jaw teeth are small, conical, and arranged in bands, with those in outer row of upper jaw and those of inner row of lower jaw slightly enlarged. Preopercular margin is finely serrated but lacks spines. Gill rakers on first arch are moderately long and slender and number 18 to 22. Measurements are expressed as percent of SL: head length 29%–36%, snout length 7%– 9%, interorbital width 7%– 8%, eye diameter 5%– 6%, upper jaw length 13%–14%, pectoral fin length 17%–20%, second anal fin spine length 11%–14%, body depth 32%– 37%, caudal peduncle depth 11%–13%. Pectoral fin has 15 to 17 rays. Dorsal fin is deeply notched and has 10 or 11 spines in anterior

section and 1 spine and 25 to 29 rays in posterior section. Anal fin has 2 spines, with second spine relatively short, and 7 or 8 rays. Body and most of head are covered with ctenoid scales. Basal parts of rayed sections of dorsal and anal fins are scaled. Gas bladder has yoke-shaped anterior chamber and carrot-shaped posterior chamber but lacks appendages. Color is bluish gray, with scattered dark spots above lateral line and stripes below. This species occurs between 10 and 30 m in the western Atlantic from both coasts of southern Florida, the Greater Antilles, and the Bay of Campeche. In the Gulf of Mexico it is known from the southwest coast of Florida and the Bay of Campeche. It is associated with clear, high saline water over vegetated mudflats and coral reef areas. Food consists of crustaceans. Maximum known size is 250 mm TL. references: Robins and Tabb 1965; Chao 1978, 2002; Robins and Ray 1986; Smith 1997; VegaCendejas et al. 1997; Castro-Aguirre et al. 1999.

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418 PERCIFORMES

Corvula sanctaeluciae Striped croaker

Jordan, 1890

Corvula sanctaeluciae is oblong to slightly elongate in profile, with a moderately oblique, terminal mouth; a truncate caudal fin; and a body marked with faint streaks. Dorsal profile is slightly convex. Posterior naris is oval shaped. Maxilla extends to about center of pupil. Snout has three rostral and five marginal pores, and chin has five mental pores. Jaw teeth are small, conical, and arranged in bands in upper jaw, with those in outer row enlarged, and those in lower jaw arranged in a single row. Preopercular margin is slightly serrated and lacks spines. Gill rakers on first arch are moderately long and slender and number 23 to 26. Measurements are expressed as percent of SL: head length 32%–34%, snout length 7%– 8%, interorbital distance 6%– 8%, eye diameter 7%–9%, upper jaw length 13%–16%, pectoral fin length 19%–21%, body depth 29%– 33%, caudal peduncle depth 10%–11%. Pectoral fin has 15 or 16 rays. Dorsal fin is deeply notched and has 10 or 11 spines in anterior section and 1 spine and 21 to 24 rays in posterior section. Anal fin has 2 spines, with sec-

ond spine moderately stout and less than twothirds the length of first ray, and 9 (rarely 8) rays. Body and most of head are covered with ctenoid scales. Basal parts of rayed sections of dorsal and anal fins are scaled. Gas bladder has yoke-shaped anterior chamber and carrotshaped posterior chamber but lacks appendages. Color is gray to grayish blue dorsally and silvery ventrally, with a faint dark spot at pectoral fin origin and oblique streaks above and longitudinal streaks below lateral line. Fins are pale yellow, with dark spots. This species occurs in coastal waters in the western Atlantic from southeastern Florida, Campeche Bank, and the Greater and Lesser Antilles, and from Costa Rica to Guyana. It is associated with muddy and sandy bottoms, but juveniles are also found in rocky areas. Food consists of shrimps. Maximum known size is 260 mm TL. references: Hildebrand 1955; Chao 1978, 2002; Robins and Ray 1986; Cervigón 1993b; Smith 1997; Vega-Cendejas et al. 1997.

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SCIAENIDAE 419

Cynoscion arenarius Sand seatrout

Ginsburg, 1930

Cynoscion arenarius is elongate, with a large, oblique mouth; large canine teeth in tip of upper jaw; and an unpatterned body. Dorsal profile of head is nearly straight. Snout is acute, with lower jaw slightly projecting and upper jaw extending to posterior margin of eye. Posterior naris is slitlike and larger than anterior naris. Teeth in upper jaw are conical and arranged in a narrow band, with outer row slightly enlarged. Teeth in lower jaw are slightly enlarged, are in a single row except at tip of jaw, and increase in size posteriorly. Snout has two marginal pores, and chin lacks pores and barbels. Preopercular margin is entire. Gill rakers on first arch are long and slender and number 11 to 15. Measurements are expressed as percent of SL: head length 31%–33%, snout length 8%–9%, eye diameter 6%–7%, interorbital width 6%–7%, upper jaw length 14%– 15%, pectoral fin length 16%–19%, body depth 21%–23%. Pectoral fin has 16 or 17 rays. Dorsal fin has 9 or 10 spines in anterior section and 1 spine and 25 to 29 rays in posterior section. Anal fin has 2 weak spines and 10 to 12 rays. Caudal fin is double emarginate. Ctenoid scales cover body, and cycloid scales cover much of head. Anterior portion of rayed

section of dorsal fin is covered with scales over basal one-third of fin. Gas bladder is single chambered, with a pair of prominent, straight, hornlike diverticula anteriorly. Color is yellowish brown dorsally and silvery ventrally, with a faint dark area at base of pectoral fin. Fins are pale yellow. This species occurs in coastal waters in the western Atlantic in the northern and southern Gulf of Mexico and possibly along the east coast of Florida (R. E. Matheson, pers. comm., August 2002). It is associated with sandy bottoms, including surf areas. Estuaries serve as nursery grounds. Food consists of crustaceans and ray-finned fishes. In the northern Gulf of Mexico maturity is reached between 140 and 180 mm TL at the end of the first year. Spawning occurs between March and September but is most intense between March and May and between August and September. Maximum known size is 450 mm TL. references: Hoese and Moore 1977, 1998; Chao 1978, 2002; Shlossman and Chittenden 1981; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999.

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420 PERCIFORMES

Cynoscion jamaicensis

(Vaillant and Bocourt, 1883)

Cynoscion jamaicensis is elongate, with a large, oblique mouth; large canine teeth in tip of upper jaw; and faint streaks along scale rows. Dorsal profile of head is nearly straight. Snout is acute, with lower jaw slightly projecting and upper jaw extending to near posterior margin of eye. Posterior naris is slitlike and larger than anterior naris. Teeth in upper jaw are conical and arranged in a narrow band, with outer row slightly enlarged. Teeth in lower jaw are slightly enlarged, are in a single row except at tip of jaw, and increase in size posteriorly. Snout has two marginal pores, and chin lacks pores and barbels. Preopercular margin is entire. Gill rakers on first arch are long and slender and number 9 to 13. Measurements are expressed as percent of SL: head length 32%–33%, snout length about 8%, eye diameter 7%– 8%, interorbital width 6%–7%, upper jaw length 14%– 15%, pectoral fin length 19%–21%, body depth 25%–26%. Pectoral fin has 16 or 17 rays. Dorsal fin has 10 spines in anterior section and 1 spine and 23 to 27 rays in posterior section. Anal fin has 2 weak spines and 8 to 10 rays. Caudal fin is truncate to emarginate. Scales cover body and much of head and are

ctenoid on body and cycloid on snout. Basal one-half to three-fourths of anterior portion of rayed section of dorsal fin is covered with scales. Gas bladder is single chambered, with a pair of prominent, straight, hornlike diverticula anteriorly. Color is grayish brown dorsally and silvery ventrally, with faint streaks above lateral line. Pectoral fin is dark basally and along upper rays, and dorsal and caudal fins have dark margins. This species occurs along the coastline to 60 m in the western Atlantic from Panama, Puerto Rico, Jamaica, and the Lesser Antilles to Brazil. It has been reported from Veracruz and the Laguna de Términos in the Gulf of Mexico; however, Castro-Aguirre et al. (1999) state that all Gulf of Mexico records of C. jamaicensis are most likely misidentifications of C. nothus. It is associated with sandy and muddy bottoms. Juveniles inhabit estuaries. Food consists of crustaceans and ray-finned fishes. Maximum known size is 500 mm TL. references: Chao 1978, 2002; Castro-Aguirre and Márquez-Espinoza 1981; Uyeno and Sato 1983a; Smith 1997.

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SCIAENIDAE 421

Cynoscion nebulosus Spotted seatrout

(Cuvier, 1830)

Cynoscion nebulosus is elongate, with a large, oblique mouth; large canine teeth in tip of upper jaw; and numerous black spots on back and upper sides. Dorsal profile of head is nearly straight. Snout is acute, with lower jaw slightly projecting and upper jaw extending to posterior margin of eye. Posterior naris is slitlike and larger than anterior naris. Teeth in upper jaw are conical and arranged in a narrow band, with outer row slightly enlarged in upper jaw, and inner row distinctly enlarged and increasing in size posteriorly in lower jaw. Snout has two marginal pores, and chin lacks pores and barbels. Preopercular margin is entire. Gill rakers on first arch are short and slender and number 9 to 12. Measurements are expressed as percent of SL: head length 30%– 33%, snout length 8%–9%, eye diameter 5%– 7%, interorbital width 5%– 6%, upper jaw length 13%–14%, pectoral fin length 12%– 15%, body depth 21%–26%. Pectoral fin has 15 to 18 rays. Dorsal fin has 9 or 10 spines in anterior section and 1 spine and 25 to 28 rays in posterior section. Anal fin has 2 spines and 10 or 11 rays. Caudal fin is truncate to emar-

ginate. Scales cover body and much of head and are ctenoid on body and cycloid on head. Rayed section of dorsal fin has two or three rows of scales on base. Gas bladder is single chambered, with a pair of prominent, straight, hornlike diverticula anteriorly. Color is dark gray, with bluish reflections dorsally and numerous black spots, and silvery or whitish ventrally. Spinous section of dorsal fin is dusky, and other fins are pale to yellowish. Rayed section of dorsal fin and caudal fin is spotted with black. This species occurs in coastal waters in the western Atlantic from New York to the southern Gulf of Mexico. It is associated with sandy bottoms, sea-grass beds, and estuaries. Estuaries serve as nursery grounds. Food consists of crustaceans and ray-finned fishes. Maximum known size is 700 mm TL. references: Hoese and Moore 1977, 1998; Chao 1978, 2002; Johnson 1978; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999.

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422 PERCIFORMES

Cynoscion nothus Silver seatrout

(Holbrook, 1848)

Cynoscion nothus is elongate, with a large, oblique mouth; large canine teeth in tip of upper jaw; and an unpatterned body. Dorsal profile of head is nearly straight. Snout is acute, with lower jaw slightly projecting and upper jaw nearly extending to posterior margin of eye. Posterior naris is slitlike and larger than anterior naris. Teeth in upper jaw are conical and arranged in a narrow band, with outer row slightly enlarged in upper jaw. Teeth in lower jaw are slightly larger than those in upper jaw, arranged in a single row except at tip of jaw, and widely spaced. Snout has two marginal pores, and chin lacks pores and barbels. Preopercular margin is entire. Gill rakers on first arch are long and slender and number 11 to 15. Measurements are expressed as percent of SL: head length 30%–31%, snout length 7%– 8%, eye diameter 7%– 8%, interorbital width 6%–7%, upper jaw length 13%–14%, pectoral fin length about 17%, body depth 23%– 26%. Pectoral fin has 15 to 17 rays. Dorsal fin has 10 spines in anterior section and 1 spine and 26 to 31 (usually 28 or 29) rays in posterior section. Anal fin has 2 spines, with second spine slender, and 8 to 11 (usually 9) rays. Caudal fin is rhomboidal to truncate in adults. Scales cover body and much of head and are

ctenoid on body and cycloid on head. Basal two-thirds of anterior portion of rayed section of dorsal fin is covered with scales. Gas bladder is single chambered, with a pair of prominent, straight, hornlike diverticula anteriorly. Color is grayish dorsally and silvery ventrally, occasionally with faint irregular rows of spots on upper side. Dorsal fin is dusky, and other fins are pale. This species occurs in coastal waters in the western Atlantic from Maryland to the Gulf of Mexico. It is found throughout the Gulf of Mexico over sandy bottoms along beaches. Food consists of crustaceans and ray-finned fishes. In the northern Gulf of Mexico maturity is reached between 140 and 170 mm SL at the end of the first year. Spawning occurs from early May through late October but is most intense in May and from August through September. Maximum known size is 360 mm TL. Records of C. jamaicensis from the Gulf of Mexico may be this species. references: Hoese and Moore 1977, 1998; Chao 1978, 2002; Johnson 1978; Shlossman and Chittenden 1981; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; CastroAguirre et al. 1999.

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SCIAENIDAE 423

Cynoscion regalis Weakfish

(Bloch and Schneider, 1801)

Cynoscion regalis is elongate and moderately compressed, with a large mouth, large canine teeth in tip of upper jaw, and back and upper sides marked with oblique streaks. Dorsal profile of head is nearly straight. Snout is acute, with lower jaw slightly projecting and upper jaw extending to posterior margin of eye. Posterior naris is slitlike and larger than anterior naris. Mouth is strongly oblique. Teeth in upper jaw are conical and arranged in a narrow band, with outer row slightly enlarged. Teeth in lower jaw are arranged in a band, with outer row distinctly enlarged from center to end of jaw. Snout has two marginal pores, and chin lacks pores and barbels. Preopercular margin is entire. Gill rakers on first arch are moderately long and slender and number 14 to 20. Measurements are expressed as percent of SL: head length 29%–32%, snout length 7%– 8%, eye diameter 6%–7%, interorbital width 5%– 6%, upper jaw length 13%–14%, pectoral fin length 13%–15%, body depth 24%–25%. Pectoral fin has 18 or 19 rays. Dorsal fin has 10 spines in anterior section and 1 spine and 24 to 29 (usually 26 to 28) rays in posterior section. Anal fin has 2 spines, with second spine

slender, and 10 to 13 (usually 12) rays. Caudal fin is truncate to emarginate. Scales cover body and much of head and are ctenoid on body and cycloid on head. Basal half of rayed section of dorsal fin is covered with scales. Gas bladder is single chambered, with a pair of prominent, straight, hornlike diverticula anteriorly. Color is greenish dorsally and silvery ventrally, with a small, irregular black spot on each scale of upper half of body forming oblique streaks. Pelvic and anal fins are bright yellow, and other fins are pale. This species occurs in shallow coastal water in the western Atlantic from Nova Scotia to southwestern Florida. It is rare along the southwestern coast of Florida. It is associated with sandy and muddy bottoms, including surf areas. Estuaries serve as nursery grounds. Spawning occurs in inlets and in estuaries with deep water. Food consists of crustaceans and ray-finned fishes. Maximum known size is 900 mm TL. references: Hildebrand and Schroeder 1928; Chao 1978, 2002; Johnson 1978; Robins and Ray 1986; Smith 1997.

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424 PERCIFORMES

Equetus lanceolatus Jackknife-fish

(Linnaeus, 1758)

Equetus lanceolatus is oblong and moderately compressed, with back elevated and strongly tapering to narrow caudal peduncle, and broad oblique bars on body. Dorsal profile of head is nearly straight, and snout is blunt and broadly rounded. Mouth is small, inferior, and oblique. Posterior naris is oval and slightly larger than anterior naris. Jaw teeth are villiform and arranged in broad bands. Snout has three marginal and five rostral pores. Chin has one small medial pore and two pairs of large lateral mental pores. Preopercular margin is membranous. First gill arch has 5 or 6 short gill rakers on upper limb and 10 to 13 on lower limb. Measurements are expressed as percent of SL: head length 26%–27%, snout length 7%–9%, eye diameter 6%–7%, upper jaw length 9%–10%, length of longest dorsal fin spine 55%– 65%, body depth 34%–36%. Pectoral fin has 14 to 16 rays. Dorsal fin has 12 or 13 spines in anterior section and 1 spine and 47 to 55 rays

in posterior section. Anal fin has 2 spines, with second spine rather stout, and 6 rays. Caudal fin is lanceolate. Body and much of head are covered with ctenoid scales. Most of posterior sections of dorsal and anal fins are covered with small ctenoid scales. Gas bladder is simple and carrot shaped, without well-developed anterior horns or diverticula. Color is tan, with three broad black bands. Upper band runs from tip of dorsal fin to tip of caudal fin. Middle band runs from nape to base of pelvic fin and is white edged. Lower band runs from top of head to corner of mouth and is white edged. Pectoral, dorsal, and anal fins are pale. Juveniles are yellow and black and have proportionally longer anterior section of dorsal fin and pelvic fin than adults. This species occurs in coastal waters in the western Atlantic from South Carolina and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of

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SCIAENIDAE 425

Mexico it is most common in the eastern and southern sections on hard bottoms. It is associated with bays and sounds, coral reefs, and hard-bottom areas. Maximum known size is 250 mm TL.

references: Böhlke and Chaplin 1968; Hastings et al. 1976; Hoese and Moore 1977, 1998; Ogren and Brusher 1977; Chao 1978, 2002; Hastings 1979; Uyeno and Sato 1983a; Robins and Ray 1986; Boschung 1992; Randall 1996; Schaldach et al. 1997; Smith-Vaniz et al. 1999.

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426 PERCIFORMES

Equetus punctatus Spotted drum

(Bloch and Schneider, 1801)

Equetus punctatus is oblong and moderately compressed, with back elevated and strongly tapering to narrow caudal peduncle, and broad oblique bars and narrow longitudinal stripes on body. Dorsal profile of head is nearly straight, and snout is blunt and broadly rounded. Mouth is small, inferior, and oblique. Posterior naris is oval and slightly larger than anterior naris. Jaw teeth are villiform and arranged in broad bands. Snout has three marginal and five rostral pores. Chin has one small medial pore and two pairs of large mental pores. Preopercular margin is membranous. First gill arch has 5 short gill rakers on upper limb and 10 to 13 on lower limb. Measurements are expressed as percent of SL: head length 28%–35%, snout length 9%–10%, eye diameter 6%–9%, upper jaw length 10%– 12%, length of longest dorsal fin spine 30%– 51%, body depth 34%– 41%. Pectoral fin has 18 rays. Dorsal fin has 11 or 12 spines in an-

terior section and 1 spine and 45 to 47 rays in posterior section. Anal fin has 2 spines, with second spine rather stout, and 6 to 8 rays. Caudal fin is lanceolate. Body and much of head are covered with ctenoid scales. Most of posterior sections of dorsal and anal fins are covered with small ctenoid scales. Gas bladder is simple and carrot shaped, without well-developed anterior horns or diverticula. Color is tan, with three broad black bands and two stripes above and one below posterior band. Posterior band runs from tip of dorsal fin to base of caudal fin. Middle band runs from nape to base of pelvic fin. Anterior band runs from top of head to corner of mouth. Pectoral fin, anterior section of dorsal fin, and pelvic fin are dark. Posterior sections of dorsal, anal, and caudal fins are dark brown with white spots. Juveniles are white and black and have proportionally longer anterior section of dorsal fin and pelvic fin than adults.

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SCIAENIDAE 427

This species occurs in coastal waters in the western Atlantic from Florida to Brazil, including the eastern Gulf of Mexico, the Bahamas, and the Antilles. It is rare to absent along the west coast of Florida but has been reported from Alabama. It has been listed from Bermuda, but the records have not been confirmed (Smith-Vaniz et al. 1999). This spe-

cies is associated with coral reefs. Maximum known size is 250 mm TL. references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Chao 2002; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997.

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428 PERCIFORMES

Larimus breviceps

(Cuvier, 1830)

Larimus breviceps is relatively deep bodied, with a large, oblique mouth and a plain-colored body. Head has prominent cavernous canals. Posterior naris is oval shaped. Snout is moderately blunt, and dorsal profile is slightly elevated. Mouth is terminal, but lower jaw projects beyond upper jaw. Maxilla extends beyond posterior margin of orbit. Snout has five marginal pores, and chin has four minute mental pores. Jaw teeth are villiform and arranged in one or two rows. Preopercular margin is nearly smooth. Operculum has 2 rather strong spines on upper portion. Gill rakers on first arch are long and slender and number 9 to 11 on upper limb and 19 to 22 on lower limb. Measurements are expressed as percent of SL: head length 31%–35%, snout length 6%–7%, eye diameter 7%– 8%, upper jaw length 15%– 16%, pectoral fin length 29%–31%, body depth 36%–39%. Pectoral fin has 16 rays. Dorsal fin is deeply notched and has 9 or 10 spines in anterior section and 1 spine and 26 to 29 rays in posterior section. Anal fin has

2 spines, with second spine long and stout, and 6 or 7 rays. Body and most of head are covered with ctenoid scales. Basal half of rayed section of dorsal fin and anal fin is scaled. Gas bladder is simple and carrot shaped, with short anterior appendages. Color is silvery and darker dorsally than ventrally, with a brown spot at pectoral fin base and pelvic and anal fins yellow. This species occurs in the western Atlantic from the southern Gulf of Mexico, the Antilles, and the coast of Costa Rica to southeastern Brazil. In the Gulf of Mexico there is an unconfirmed record from the western coast of Campeche, at Celestún. This species is associated with mud and sandy mud bottoms from near shore to 60 m, and juveniles occur in estuaries. Food consists of small shrimps. Maximum known size is 310 mm TL. references: Chao 1978, 2002; Uyeno and Sato 1983a; Cervigón 1993b; Smith 1997; VegaCendejas et al. 1997.

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SCIAENIDAE 429

Larimus fasciatus Banded drum

Holbrook, 1855

Larimus fasciatus is relatively deep bodied, with a large, oblique mouth and seven to nine dark vertical bars on body. Head has prominent cavernous canals. Posterior naris is oval shaped. Snout is moderately blunt, and dorsal profile is slightly elevated. Mouth is terminal, but lower jaw projects beyond upper jaw. Maxilla extends beyond posterior margin of orbit. Snout has five marginal pores, and chin has four minute mental pores. Jaw teeth are villiform and arranged in one or two rows. Preopercular margin is finely serrated. Operculum has 2 rather strong spines on upper portion. Gill rakers on first arch are long and slender and number 11 to 13 on upper limb and 22 to 25 on lower limb. Measurements are expressed as percent of SL: head length 30%–32%, snout length 6%–7%, eye diameter 7%– 8%, upper jaw length 15%–16%, pectoral fin length 23%–27%, body depth 32%–34%. Pectoral fin has 16 rays. Dorsal fin is deeply notched and has 10 spines in anterior section and 1 spine and 24 to 27 rays in posterior section. Anal fin has 2 spines, with second spine long and stout, and 6 or 7 rays. Body and most of head are

covered with ctenoid scales. Basal half of rayed section of dorsal fin and anal fin is scaled. Gas bladder is simple and carrot shaped and lacks anterior appendages. Color is grayish olive dorsally and silvery ventrally, with vertical bars extending to or below lateral line. Distal parts of pelvic and caudal fins are yellow. This species occurs in the western Atlantic from Massachusetts to Veracruz in the Gulf of Mexico, but it is rare to absent around southern Florida. It is associated with mud and sandy mud bottoms from near shore to 100 m and rarely occurs in estuaries. Food consists of small shrimps. In the northern Gulf of Mexico maturity is reached between 80 and 139 mm TL near the end of the first year. Spawning occurs between September and November and, to a lesser degree, between April and June. Maximum known size is 250 mm TL. references: Hoese and Moore 1977, 1998; Chao 1978, 2002; Johnson 1978; Standard and Chittenden 1983; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999.

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430 PERCIFORMES

Leiostomus xanthurus Spot

Lacepède, 1802

Leiostomus xanthurus is relatively deep bodied, with a small, nearly horizontal mouth; a dark blotch behind upper section of operculum; and oblique dark streaks on flank. Dorsal profile of head is elevated above eye. Posterior naris is slitlike and larger than anterior naris. Maxilla extends to about center of pupil. Snout has five marginal and five rostral pores. Chin has about five mental pores. Jaw teeth are villiform and arranged in bands. Preopercular margin is membranous. Gill rakers on first arch are moderately short and slender and number 8 to 12 on upper limb and 20 to 24 on lower limb. Measurements are expressed as percent of SL: head length 29%–32%, snout length 7%–9%, eye length 7%– 8%, upper jaw length 10%–11%, pectoral fin length 27%–31%, body depth 32%–36%. Pectoral fin has 19 to 24 rays. Dorsal fin is deeply notched and has 9 to 11 spines in anterior section and 1 spine and 29 to 35 rays in posterior section. Anal fin has 2 spines and 12 or 13 rays. Caudal fin is concave. Body and most of head are covered with ctenoid scales. Basal parts of rayed sections of

dorsal and anal fins are scaled. Gas bladder is simple and carrot shaped and lacks appendages. Color is silvery gray and darker dorsally than ventrally, with 11 to 15 dark streaks extending to below lateral line and a black blotch, equal to size of iris, behind upper end of operculum. Dorsal and caudal fins are dusky, and other fins are pale or yellowish. This species occurs in the western Atlantic from Massachusetts to Veracruz but is possibly absent from southeastern Florida. It is common in the northern and western Gulf of Mexico to Veracruz. This species is associated with sandy or muddy bottoms from near shore to 60 m and uses estuaries as nursery grounds. Juveniles may remain in estuaries year-round. Food consists of detritus, polychaetes, and small crustaceans. Maximum known size is 360 mm TL. references: Hastings et al. 1976; Hoese and Moore 1977, 1998; Chao 1978, 2002; Johnson 1978; Robins and Ray 1986; Boschung 1992; Castro-Aguirre et al. 1999.

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SCIAENIDAE 431

Menticirrhus americanus Southern kingfish

(Linnaeus, 1758)

Menticirrhus americanus is elongate, with a small, nearly horizontal mouth; a nearly straight ventral profile; and seven or eight dusky bars on body. Dorsal profile of head is slightly convex. Rostral fold on lower margin of snout is deeply notched. Posterior naris is oval shaped and larger than anterior naris. Mouth is inferior. Maxilla extends to about center of pupil. Snout has five marginal and three rostral pores. Chin has a single short, rigid barbel and one medial pore and two pairs of mental pores. Jaw teeth are villiform and arranged in broad bands, with those in outer row of upper jaw slightly enlarged. Preopercular margin is serrated. Gill rakers on first arch are knoblike and number about 10 but largely disappear with growth. Measurements are expressed as percent of SL: head length 23%– 30%, snout length 8%–9%, eye diameter 4%– 6%, upper jaw length 8%–9%, length of longest dorsal fin spine 16%–24%, pectoral fin length 22 to 26%, body depth 21%–26%. Pectoral fin has 21 to 24 rays. Dorsal fin is deeply notched and has 10 or 11 spines in anterior section and 1 spine and 22 to 26 rays in posterior section. Anal fin has 1 spine and 6 to 8 rays. Posterior margin of caudal fin is concave along upper half and convex along lower half. Body and most of head are covered with ctenoid

scales. Scales on chest are about same size as those on remainder of body. Gas bladder is vestigial. Color is grayish brown on dorsal half and silvery gray on lower half of body, with seven or eight obscure, dusky diagonal bands on flank. Second and third bars form a V below nape and spinous section of dorsal fin. Pectoral, pelvic, and anal fins are dusky. Lining of gill cavity is dusky. This species occurs in coastal waters in the western Atlantic from New York to Argentina, including the northern and southern Gulf of Mexico and the Greater Antilles. It is associated with sandy, muddy bottoms and hard mud bottoms from the surf zone to estuaries. Food consists of polychaetes and benthic crustaceans. In the northern Gulf of Mexico it matures at 150 to 220 mm TL at the end of, or shortly after, the first year. Spawning occurs between February and November but most intensely from February to April and from August to November. Maximum known size is 600 mm TL. references: Hoese and Moore 1977, 1998; Chao 1978, 2002; Johnson 1978; Uyeno and Sato 1983a; Robins and Ray 1986; Harding and Chittenden 1987; Boschung 1992; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999.

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432 PERCIFORMES

Menticirrhus littoralis Gulf kingfish

(Holbrook, 1847)

Menticirrhus littoralis is elongate, with a small, nearly horizontal mouth; a nearly straight ventral profile; and relatively small scales on the chest. Dorsal profile of head is slightly convex. Rostral fold on lower margin of snout is deeply notched. Posterior naris is oval shaped and larger than anterior naris. Mouth is inferior. Maxilla extends to about center of pupil. Snout has five marginal and three rostral pores. Chin has a single short, rigid barbel and one medial pore and two pairs of mental pores. Jaw teeth are villiform and arranged in broad bands, with those in outer row of upper jaw slightly enlarged. Preopercular margin is serrated. Gill rakers on first arch are knoblike and number 3 to 12. Measurements are expressed as percent of SL: head length 26%–31%, snout length 9%–11%, eye diameter 4%– 6%, upper jaw length 9%–10%, length of longest dorsal fin spine 18%–21%, pectoral fin length 18%– 21%, body depth 22%–24%. Pectoral fin has 18 to 21 rays. Dorsal fin is deeply notched and has 10 or 11 spines in anterior section and

1 spine and 19 to 26 rays in posterior section. Anal fin has 1 spine and 6 to 8 rays. Posterior margin of caudal fin is concave along upper half and convex along lower half. Body and most of head are covered with ctenoid scales, with those on chest relatively small. Gas bladder is vestigial. Color is silvery, with tip of dorsal fin often dusky and tip of caudal fin blackish. Lining of gill cavity is dusky. This species occurs in coastal waters in the western Atlantic from Virginia to Brazil, including the northern and southern Gulf of Mexico. It is associated with sandy and muddy bottoms and is most abundant in the surf zone. Occasionally it occurs in estuaries but is rare in water less saline than 21‰. Food consists of polychaetes and benthic crustaceans. Maximum known size is 600 mm TL. references: Hoese and Moore 1977, 1998; Chao 1978, 2002; Johnson 1978; Robins and Ray 1986; Boschung 1992; Vega-Cendejas et al. 1997; CastroAguirre et al. 1999.

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SCIAENIDAE 433

Menticirrhus saxatilis Northern kingfish

(Bloch and Schneider, 1801)

Menticirrhus saxatilis is elongate, with a small, nearly horizontal mouth; a nearly straight ventral profile; and five or six conspicuous dusky bars on body. Dorsal profile of head is slightly convex. Rostral fold on lower margin of snout is deeply notched. Posterior naris is oval shaped and larger than anterior naris. Mouth is inferior. Maxilla extends to about center of pupil. Snout has five marginal and three rostral pores. Chin has a single short, rigid barbel and one medial pore and two pairs of mental pores. Jaw teeth are villiform and arranged in broad bands, with those in outer row of upper jaw slightly enlarged. Preopercular margin is serrated. Gill rakers on first arch are knoblike and number 3 to 12 but disappear with growth. Measurements are expressed as percent of SL: head length 25%–29%, snout length 8%–9%, eye length 5%– 6%, upper jaw length 8%– 10%, length of longest dorsal fin spine 25%– 39%, pectoral fin length 20%–25%, body depth 22%–25%. Pectoral fin has 20 or 21 rays. Dorsal fin is deeply notched and has 10 or 11 spines in anterior section and 1 spine and 22 to 27 rays in posterior section. Anal fin has 1 spine and 7 to 9 rays. Posterior margin of caudal fin is concave along upper half and con-

vex along lower half. Body and most of head are covered with ctenoid scales. Scales on chest are about the same size as those on remainder of body. Gas bladder is vestigial in adults but well developed in juveniles. Color is grayish brown on dorsal half and silvery gray on lower half of body, with five or six distinct diagonal bars on flank. Second and third bars form a Vshaped marking below spinous section of dorsal fin. Lining of gill cavity is dusky. This species occurs in shallow coastal waters in the western Atlantic from Massachusetts to the southern Gulf of Mexico. It is most common along the coast of North America between Cape Cod and Cape Hatteras. It is associated with sandy and muddy bottoms from the surf zone to estuaries. Food consists of polychaetes and benthic crustaceans. Juveniles enter low-salinity estuaries. Maximum known size is 460 mm TL. references: Ginsburg 1952a (as M. folcaliger); Hoese and Moore 1977, 1998; Chao 1978, 2002; Johnson 1978; Hastings 1979; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Castro-Aguirre et al. 1999.

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434 PERCIFORMES

Micropogonias furnieri (Desmarest, 1823)

Micropogonias furnieri is moderately elongate, with a nearly horizontal mouth, three or four pairs of minute chin barbels, and oblique or wavy stripes on side. Dorsal profile of head is slightly elevated. Posterior naris is oval shaped and larger than anterior naris. Mouth is subterminal. Maxilla extends to about middle of eye. Snout has five marginal and five rostral pores. Chin has five mental pores. Jaw teeth are villiform and arranged in broad bands, with outer row in upper jaw slightly enlarged. Preopercular margin is serrated, serrations increase in size ventrally, and 2 or 3 strong spines are located at angle. Gill rakers on first arch are short and slender and number 21 to 25, with 7 to 9 on upper limb and 12 to 15 on lower limb. Measurements are expressed as percent of SL: head length 32%–34%, snout length 10%– 11%, eye diameter 6%– 8%, interorbital width 7%– 8%, upper jaw length 10%–11%, length of longest dorsal fin spine 19%–22%, pectoral fin length 24%–27%, body depth 26%–29%. Pectoral fin has 17 to 19 rays. Dorsal fin is deeply notched and has 10 spines in anterior section and 1 spine and 26 to 30 rays in posterior section. Anal fin has 2 spines and 7 to 9 rays. Posterior margin of caudal fin is double emarginate in adults. Body and most of head

are covered with ctenoid scales. Scale rows between origin of dorsal fin and lateral line number six or seven. Vertebrae number 25: 10 precaudal and 15 caudal. Gas bladder is carrot shaped, with a pair of tubelike diverticula originating laterally on posterior half of main chamber and extending to septum transversum. Color is silvery, with rows of dark spots on side above and slightly below the lateral line forming stripes and a dark spot on operculum. Spinous section of dorsal fin is black edged, and other fins are pale to yellowish. This species occurs in coastal waters to 80 m in the western Atlantic from Costa Rica to Argentina, including the Antilles. It has been reported from Tuxpan, Veracruz, and Celestún, Yucatán, in the Gulf of Mexico, but some investigators attribute these records to M. undulatus. This species is associated with sandy and muddy bottoms along the coast and in estuaries. Juveniles and young adults occur in estuaries year-round. Maximum known size is about 900 mm TL.

references: Hildebrand 1955; Chao 1978, 2002; Uyeno and Sato 1983a; Smith 1997; VegaCendejas et al. 1997.

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SCIAENIDAE 435

Micropogonias undulatus Atlantic croaker

(Linnaeus, 1766)

Micropogonias undulatus is moderately elongate, with a nearly horizontal mouth, three to five pairs of minute chin barbels, and irregular oblique stripes on side. Dorsal profile of head is slightly elevated. Posterior naris is oval shaped and larger than anterior naris. Mouth is subterminal. Maxilla extends to about middle of eye. Snout has five marginal and five to seven rostral pores. Chin has five mental pores. Jaw teeth are villiform and arranged in broad bands. Preopercular margin is serrated, serrations increase in size ventrally, and three to five strong spines are present at angle. Gill rakers on first arch are short and slender and number 22 to 30 (usually 23 to 26). Measurements are expressed as percent of SL: head length 30%–34%, snout length 10%–12%, eye length 6%–7%, interorbital width 7%– 8%, upper jaw length 10%–12%, length of longest dorsal fin spine 17%–18%, pectoral fin length 20%–25%, body depth 28%–30%. Pectoral fin has 17 to 19 rays. Dorsal fin is deeply notched and has 10 spines in anterior section and 1 spine and 26 to 30 rays in posterior section. Anal fin has 2 spines and 7 to 9 rays. Posterior margin of caudal fin is double emarginate in adults. Body and most

of head are covered with ctenoid scales. Scale rows between origin of dorsal fin and lateral line number eight or nine. Vertebrae number 25: 10 precaudal and 15 caudal. Gas bladder is carrot shaped, and a pair of tubelike diverticula originate laterally on posterior half of main chamber and extend to septum transversum. Color is silvery, with dark spots on dorsal fin and rows of dark spots on sides above and slightly below the lateral line that form irregular broken lines but not oblique or wavy lines. This species occurs in the western Atlantic from Massachusetts to the southern Gulf of Mexico. It occurs in the entire Gulf of Mexico from near shore to about 60 m. It is associated with sandy and muddy bottoms along the coast and in estuaries. Estuaries serve as nursery grounds. Food consists of polychaetes, benthic crustaceans, and small ray-finned fishes. Maximum known size is 500 mm TL.

references: Chao and Musick 1977; Chao 1978, 2002; Hoese and Moore 1977, 1998; White and Chittenden 1977; Johnson 1978; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; CastroAguirre et al. 1999.

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436 PERCIFORMES

Odontoscion dentex Reef croaker

(Cuvier, 1830)

Odontoscion dentex is oblong, with a truncate to double-emarginate caudal fin, and a large black blotch on the pectoral fin base. Dorsal profile is evenly convex. Posterior naris is oval shaped. Mouth is large and terminal to slightly subterminal. Maxilla extends to about posterior margin of eye. Snout has three rostral and five marginal pores, and chin has four mental pores. Jaw teeth are large, conical, and arranged in a single series. Two large canine teeth occur at tip of lower jaw. Preopercular margin is finely denticulate. Gill rakers on first arch are long and stiff and number 19 to 25. Measurements are expressed as percent of SL: head length 31%–34%, snout length 7%–9%, interorbital width 7%– 8%, eye diameter 9%– 10%, upper jaw length 14%–16%, length of longest dorsal fin spine 14%–17%, pectoral fin length about 17%, body depth 27%–29%. Pectoral fin has 15 to 17 rays. Dorsal fin is deeply notched and has 11 or 12 spines in anterior section and 1 spine and 23 to 26 rays in posterior section. Anal fin has 2 spines, and 8 to 10 rays. Body and most of head are covered

with ctenoid scales. Gas bladder has yokeshaped anterior chamber and carrot-shaped posterior chamber but lacks appendages. Color is silvery to brownish gray, with dark spots on scales and a large dark spot on pectoral fin base. Inner surface of opercular cavity is dark. This species occurs in coastal waters in the western Atlantic from southern Florida and Cuba to Brazil, including the Gulf of Mexico, the Florida Keys, and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from southwestern Florida, the Dry Tortugas, Veracruz, and the west coast of the Yucatán Peninsula. It is absent along the Caribbean coasts of Belize and Honduras. It is associated with sandy mud bottoms and coral reefs. Food consists of shrimps and small ray-finned fishes. Maximum known size is 300 mm TL. references: Longley and Hildebrand 1941; Chao 1978, 2002; Uyeno and Sato 1983a; Robins and Ray 1986; Randall 1996; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997.

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SCIAENIDAE 437

Paralonchurus brasiliensis

(Steindachner, 1875)

Paralonchurus brasiliensis is elongate, with a tuft and a parallel series of barbels on lower jaw. Dorsal profile of head is slightly elevated. Mouth is small, subterminal, and nearly horizontal. Posterior naris is oval shaped and larger than anterior naris. Maxilla extends slightly beyond center of pupil. Snout has five marginal and three or four rostral pores. Lower jaw has five mental pores, three pairs of small barbels at tip, and 10 to 12 barbels along each side. Jaw teeth are villiform and arranged in bands. Gill rakers on first arch are short and stout and number three to five on upper limb and six to nine on lower limb. Measurements are expressed as percent of SL: head length 23%– 25%, snout length 6%– 8%, eye diameter 4%– 5%, interorbital width 7%–9%, upper jaw length 7%–9%, pectoral fin length 22%–25%, body depth 25%–26%. Pectoral fin is short, does not extend beyond tip of pelvic fin, and has 19 rays. Dorsal fin is deeply notched and has 10 spines in anterior section and 1 spine and 28 to 31 rays in posterior section. Anal fin

has 2 spines and 7 to 9 rays. Posterior margin of caudal fin is pointed. Body and most of head are covered with ctenoid scales. Vertebrae number 29: 11 precaudal and 18 caudal. Gas bladder has two anterior lateral diverticula, each with a long, slender branch extending posteriorly to near posterior extension of bladder at vent. Males have a drumming muscle associated with gas bladder. Color is green to bluish gray dorsally and whitish ventrally, with a dark spot on lateral line behind operculum and seven to nine bars or saddles on upper flank. This species occurs in coastal waters in the western Atlantic from the Caribbean coast of Panama to the mouth of the Amazon. There is one unconfirmed record from the western coast of Campeche, at Celestún, in the Gulf of Mexico. Maximum known size is 300 mm TL.

references: Chao 1978, 2002; Cervigón 1993b; Smith 1997; Vega-Cendejas et al. 1997.

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438 PERCIFORMES

Pareques acuminatus High-hat

(Bloch and Schneider, 1801)

Pareques acuminatus is oblong, with back elevated and tapering posteriorly to caudal peduncle, and sides with four to seven relatively broad longitudinal stripes. Dorsal profile of head is slightly concave. Mouth is small, inferior, and nearly horizontal. Posterior naris is oval and slightly larger than anterior naris. Jaw teeth are villiform and arranged in broad bands, except those in outer row of upper jaw are enlarged and recurved. Snout has three large marginal pores and three small rostral pores. Chin has five pores: one medial pore and two pairs of mental pores. Preopercular margin is membranous. First gill arch has 5 to 7 gill rakers on upper limb and 9 to 14 on lower limb. Measurements are expressed as percent of SL: head length 31%–36%, snout length 9%–11%, eye diameter 8%–10%, upper jaw length 12%–13%, length of longest dorsal fin spine 18%–19%, body depth 33%– 39%. Pectoral fin has 16 or 17 rays. Dorsal fin has 8 or 9 spines in anterior section and 1 spine and 37 to 41 rays in posterior section. Anal fin has 2 spines, with second spine rather stout, and 7 or 8 rays. Caudal fin is convex. Body and

much of head are covered with ctenoid scales. Basal half of posterior section of dorsal fin and anal fin is covered with small ctenoid scales. Gas bladder is simple and carrot shaped, without distinctly developed anterior horns or diverticula. Color is whitish, with dark brown stripes more or less evenly spaced along body. Stripes vary slightly in width, but some are equal to width of pupil. Anterior section of head is spotted. Fins are dusky to dark. Prejuveniles have three stripes, a large black spot at tip of snout and lower jaw, and a black spot on anal fin that is continuous with ventral stripe. Juveniles have four broad and three narrow stripes. Stripe immediately above eye coalesces with stripe of opposite side, and snout has one large and two small black spots. This species occurs in coastal waters in the western Atlantic from Virginia to Brazil, including the northern and southern Gulf of Mexico and the Bahamas. Records from the eastern Gulf of Mexico may apply to E. umbrosus (Boschung 1992). This species has been reported from Bermuda, but these records have not been confirmed (Smith-Vaniz et al. 1999).

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SCIAENIDAE 439

It is associated with clear water near coral reefs and hard-bottom areas. Maximum known size is 250 mm TL. references: Haburay et al. 1969; Chao 1978, 2002; Uyeno and Sato 1983a; Robins and Ray

1986 (as Equetus acuminatus); Miller and Woods 1988 (as E. acuminatus); Boschung 1992 (as E. acuminatus); Randall 1996; Schaldach et al. 1997 (as E. acuminatus); Smith 1997; CastroAguirre et al. 1999.

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440 PERCIFORMES

Pareques iwamotoi Blackbar drum

Miller and Woods, 1988

Pareques iwamotoi is oblong and moderately compressed, with back elevated and tapering posteriorly to caudal peduncle, and body with broad band below first dorsal fin and narrow dark stripe posterior to band. Dorsal profile of head is nearly straight, and snout is moderately blunt and rounded. Mouth is small, inferior, and nearly horizontal. Posterior naris is oval and slightly larger than anterior naris. Jaw teeth are villiform and arranged in broad bands, except those in outer row of upper jaw are enlarged and recurved. Snout has three large marginal pores and three small rostral pores. Chin has one medial pore and two pairs of lateral mental pores. Preopercular margin is membranous. First gill arch has 4 to 7 gill rakers on upper limb, 1 in corner, and 9 to 12 on lower limb. Measurements are expressed as percent of SL: head length 31%–33%, snout length 8%–10%, eye diameter 8%–10%, upper jaw length 10%–12%, length of longest dorsal fin spine 16%–21%, body depth 36%– 41%. Pectoral fin has 15 to 18 rays. Dorsal fin has 8 to 10 spines in anterior section and 1 spine and 33 to 40 rays in posterior section. Anal fin has 2 spines, with second spine rather stout, and 7 or 8 rays. Caudal fin is convex.

Body and much of head are covered with ctenoid scales. Basal half of rayed section of dorsal fin and anal fin is covered with small ctenoid scales. Vertebrae number 25: 10 precaudal and 15 caudal. Gas bladder is simple, carrot shaped, and without distinct anterior horns or diverticula. Color is light to dark brown, with a broad, oblique, and very dark band extending from base of spiny section of dorsal fin, behind pectoral fin, to abdomen, and a narrow dark stripe extending from band to end of caudal fin. Band and stripe are most distinct on specimens ranging from 95 to 110 mm SL, less distinct on specimens from 170 to 213 mm SL, and obscure on specimens from 247 to 298 mm SL. Fins are dusky to black. This species occurs in the western Atlantic from North Carolina to Brazil, including the entire Gulf of Mexico, from 37 to 184 m. It is associated with reefs and hard bottoms. Food consists of gastropods. Maximum known size is 330 mm SL. references: Hoese and Moore 1977 (as Equetus sp.), 1998; Chao 1978 (as E. sp.), 2002; Robins and Ray 1986; Miller and Woods 1988; Boschung 1992; Schaldach et al. 1997.

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SCIAENIDAE 441

Pareques umbrosus Cubbyu

(Jordan and Eigenmann, 1889)

Pareques umbrosus is oblong and moderately compressed, with back elevated and tapering posteriorly to caudal peduncle, and side has seven to ten narrow longitudinal stripes. Dorsal profile of head is slightly concave. Mouth is small and inferior. Posterior naris is oval and slightly larger than anterior naris. Jaw teeth are villiform and arranged in broad bands, except those in outer row of upper jaw are enlarged and recurved. Snout has five marginal and five rostral pores. Chin has one medial pore and two pairs of mental pores. Preopercular margin is finely serrated. First gill arch has 4 to 6 gill rakers on upper limb and 10 to 12 on lower limb. Measurements are expressed as percent of SL: head length 30%–33%, snout length 9%–10%, eye diameter 7%– 8%, upper jaw length 10%–12%, length of longest dorsal fin spine 16%–20%, body depth 35%–38%. Pectoral fin has 14 to 16 rays. Dorsal fin has 9 or 10 spines in anterior section and 1 spine and 38 to 40 rays in posterior section. Anal fin has 2 spines, with second spine rather stout, and 7 rays. Caudal fin is convex. Body and much of head are covered with ctenoid scales. Basal half of posterior section of dorsal fin and anal

fin is covered with small ctenoid scales. Gas bladder is simple and carrot shaped, without well-developed anterior horns or diverticula. Color is dusky brown, with seven distinct stripes alternating with six very narrow interspaces. All stripes are narrower than pupil width. Stripe immediately above eye extends anteriorly to or near tip of snout. Fins are very dark. Prejuveniles have three stripes on body and a small spot on snout, and anal fin is dusky and not connected to ventral stripe. This species occurs from near shore to 91 m in the western Atlantic from North Carolina and Bermuda to Brazil, including the entire Gulf of Mexico. Many or all of the records of E. acuminatus from the eastern Gulf of Mexico may refer to this species (Darovec 1983). It is associated with muddy and sandy bottoms. Maximum known size is 250 mm TL. references: Hoese and Moore 1977, 1998; Chao 1978, 2002; Darovec 1983; Robins and Ray 1986; Miller and Woods 1988; Boschung 1992; Cervigón 1993b; Smith 1997; Smith-Vaniz et al. 1999.

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442 PERCIFORMES

Pogonias cromis Black drum

(Linnaeus, 1766)

Pogonias cromis is oblong, with a subterminal and nearly straight mouth and 10 to 13 pairs of chin barbels. Dorsal profile of head is slightly elevated. Posterior naris is oval shaped and larger than anterior naris. Maxilla extends to about center of pupil. Snout has five marginal and five rostral pores. Lower jaw has five mental pores and 10 to 13 pairs of barbels along inner margins, with barbels increasing in size posteriorly. Jaw teeth are villiform and arranged in broad bands. Preopercular margin is entire. Gill rakers on first arch are short and stout and number 16 to 21. Measurements are expressed as percent of SL: head length 31%– 34%, snout length 10%–12%, eye diameter 6%–9%, interorbital width 7%–9%, upper jaw length 10%–12%, length of longest dorsal fin spine 19%–24%, pectoral fin length 27%– 31%, body depth 33%–36%. Pectoral fin has 17 to 19 rays. Dorsal fin is deeply notched and has 10 spines in anterior section and 1 spine and 19 to 22 rays in posterior section. Anal fin has 2 spines, with second spine relatively stout and long, and 5 to 7 rays. Posterior margin of caudal fin is truncate to slightly emarginate in adults. Body and most of head are covered with ctenoid scales. Vertebrae number 24: 10

precaudal and 14 caudal. Gas bladder is single chambered, with many complex diverticula along lateral margin. Diverticula communicate with main chamber and with each other and become increasingly developed with growth. Color is silvery gray to dark gray. Juveniles and young adults have four or five broad black bars that are widest on back, and last bar does not extend below lateral line. Bars become obscure in large individuals. Fins are blackish. This species occurs in coastal waters in the western Atlantic from Nova Scotia to northern Mexico, including the entire Gulf of Mexico and the Antilles. It is rare in the Antilles and western Caribbean. This species is associated with sand and sandy mud bottoms, especially near large river mouths and estuaries. Juveniles use estuaries as nursery grounds. Food consists of benthic crustaceans, mollusks, and small ray-finned fishes. Maximum known size is 170 cm TL. references: Hoese and Moore 1977, 1998; Chao 1978, 2002; Johnson 1978; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; VegaCendejas et al. 1997; Castro-Aguirre et al. 1999.

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SCIAENIDAE 443

Sciaenops ocellata (Linnaeus, 1766) Reddrum

Sciaenops ocellata is elongate, with a nearly straight ventral profile and one or more black ocellated blotches on posterior section of body. Dorsal profile of head is evenly convex. Mouth is subterminal and nearly horizontal. Posterior naris is oval shaped and larger than anterior naris. Maxilla extends to about posterior margin of orbit. Snout has five marginal and five rostral pores. Chin has five mental pores. Jaw teeth are villiform and arranged in broad bands, with those in outer row of upper jaw slightly enlarged. Preopercular margin is finely serrated in juveniles but entire in adults. Gill rakers on first arch are short and slender and number 12 to 14. Measurements are expressed as percent of SL: head length 30%–32%, snout length 6%–10%, eye diameter 5%–7%, interorbital width 6%–7%, upper jaw length 12%– 13%, length of longest dorsal fin spine 12%– 14%, pectoral fin length 16%–18%, body depth 24%–27%. Pectoral fin has 17 to 19 rays. Dorsal fin is deeply notched and has 10 spines in anterior section and 1 spine and 23 to 25 rays in posterior section. Anal fin has 2 spines, with second relatively stout and long, and 8 or 9 rays. Posterior margin of caudal fin

is truncate in adults. Body and most of head are covered with ctenoid scales. Vertebrae number 25: 10 precaudal and 15 caudal. Gas bladder is single chambered, with a pair of small tubeshaped appendages that develop additional appendages with growth. Color is bronze and darker dorsally than ventrally, with dark centers of scales forming obscure stripes and dark ocellated spot or spots on caudal peduncle. This species occurs in the western Atlantic from Massachusetts to Veracruz, Mexico. In the Gulf of Mexico it is common in shallow water, although it is apparently not common south of the Laguna Madre. This species is associated with sand and sandy mud bottoms from the surf zone to estuaries. Juveniles use estuaries as nursery grounds. Food consists of benthic crustaceans, mollusks, and small ray-finned fishes. Maximum known size is 160 cm TL. references: Hastings et al. 1976; Hoese and Moore 1977, 1998; Chao 1978, 2002; Johnson 1978; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Castro-Aguirre et al. 1999.

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444 PERCIFORMES

Stellifer colonensis Colon stardrum

Meek and Hildebrand, 1925

Stellifer colonensis is oblong in profile, has a broad head, is slightly concave at nape, and possesses cavernous canals. Posterior naris is oval shaped and slightly larger than anterior naris. Mouth is relatively large, oblique, and subterminal. Maxilla extends beyond posterior margin of eye. Snout has three rostral and five marginal pores, and chin has six minute mental pores. Jaw teeth are villiform and arranged in bands, with outer row in upper jaw slightly enlarged. Preopercular margin has five or six stout spines, with lower spines strongest. Gill rakers on first arch are long and slender and number 27 to 34. Dorsal fin is deeply notched and has 11 spines in anterior section and 1 spine and 22 or 23 rays in posterior section. Anal fin has 2 spines and 8 or 9 rays. Caudal fin is lanceolate. Ctenoid scales occur on body, and cycloid scales occur on head. Basal parts of rayed sections of dorsal and anal fins are

scaled. Gas bladder consists of yoke-shaped anterior chamber and carrot-shaped posterior chamber, with small diverticula on posterolateral aspect of anterior chamber. Color is grayish dorsally and pale ventrally, with a dark blotch above operculum. Fins are pale to dusky, and spinous section of dorsal fin is dark. This species occurs between the shoreline and 20 m in the western Atlantic from the southern Gulf of Mexico and the Greater Antilles to Venezuela. In the Gulf of Mexico it occurs from Veracruz to northern Yucatán. This species is associated with hard sandy mud bottoms, edges of coral reefs, and estuaries. Food consists of small crustaceans. Maximum known size is 200 mm TL.

references: Meek and Hildebrand 1925; Chao 1978, 2002.

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SCIAENIDAE 445

Stellifer lanceolatus Star drum

(Holbrook, 1855)

Stellifer lanceolatus is oblong, has a broad head, is slightly concave at nape, and possesses cavernous canals. Posterior naris is oval shaped and slightly larger than anterior naris. Mouth is relatively large, oblique, and subterminal to terminal. Maxilla extends to about posterior margin of eye. Snout has three rostral and five marginal pores, and chin has six minute mental pores. Jaw teeth are villiform and arranged in bands, with outer row in upper jaw slightly enlarged. Preopercular margin has 4 to 6 stout spines. Gill rakers on first arch are long and slender and number 32 to 36. Measurements are expressed as percent of SL: head length 30%–32%, snout length 8%–9%, eye diameter 5%– 6%, interorbital width 10%–11%, upper jaw length 13%–15%, pectoral fin length 29%–31%, length of longest dorsal fin spine 15%–18%, body depth 28%–34%. Pectoral fin has 18 to 20 rays. Dorsal fin is deeply notched and has 11 or 12 spines in anterior section and 1 spine and 20 to 25 rays in posterior section. Anal fin has 2 spines and 8 or 9 rays. Caudal fin is lanceolate. Ctenoid scales occur on body, and cycloid scales occur on

head. Basal parts of rayed sections of dorsal and anal fins are scaled. Gas bladder consists of yoke-shaped anterior chamber and carrotshaped posterior chamber, with small diverticula on posterolateral aspect of anterior chamber. Color is silvery olive to brown dorsally and pale ventrally. Fins are pale to dusky, and spinous section of dorsal fin is dark. This species occurs between the shoreline and 20 m in the western Atlantic from Virginia to the Gulf of Mexico. It is common in the western Gulf, present but uncommon off Alabama and the Florida Panhandle in the eastern Gulf or off Veracruz in the southwestern Gulf, and rare or absent along the Florida peninsula (R. E. Matheson, pers. comm., August 2002). This species is associated with hard sandy mud bottoms and is common in estuaries. Food consists of crustaceans. Maximum known size is 200 mm TL. references: Hoese and Moore 1977, 1998; Chao 1978, 2002; Johnson 1978; Robins and Ray 1986; Boschung 1992; Smith 1997; Castro-Aguirre et al. 1999.

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446 PERCIFORMES

Umbrina coroides Sand drum

Cuvier, 1830

Umbrina coroides is slightly elongate, with a nearly straight ventral profile and a short, blunt chin barbel. Dorsal profile of head is moderately elevated. Posterior naris is oval shaped and larger than anterior naris. Mouth is moderately small, subterminal, and nearly horizontal. Maxilla extends beyond center of pupil. Snout has five marginal and three rostral pores. Chin has five mental pores: one in tip of barbel and two on each side behind barbel. Jaw teeth are villiform and arranged in broad bands. Preopercular margin is finely serrated. Gill rakers on first arch are short and number 13 to 15. Measurements are expressed as percent of SL: head length 26%–30%, snout length 8%– 10%, eye diameter 6%– 8%, interorbital width 6%– 8%, upper jaw length 9%–11%, length of longest dorsal fin spine 14%–20%, pectoral fin length 17%–19%, body depth 26%–31%. Pectoral fin has 16 to 19 rays. Dorsal fin is deeply notched and has 9 or 10 spines in anterior section and 1 spine and 26 to 31 rays in posterior section. Anal fin has 2 spines and

6 rays, with second spine relatively long. Posterior margin of caudal fin is truncate or emarginate. Body and most of head are covered with ctenoid scales. Gas bladder has a single chamber and is carrot shaped. Color is silvery gray dorsally and pale yellowish ventrally, with 9 or 10 narrow dark bars and a number of narrow wavy stripes. This species occurs along the coast in the western Atlantic from Virginia and the Bahamas to Brazil, including the western Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs from Texas to Laguna de Términos. This species is associated with sandy beaches, muddy bottoms, estuaries, and occasionally coral reefs. Food consists of benthic organisms. Maximum known size is 350 mm TL. references: Hoese and Moore 1977, 1998; Chao 1978, 2002; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999.

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447

MULLIDAE

Goatfishes

Goatfishes are oblong and moderately compressed, with a convex dorsal profile and a flattened ventral side; two long, unbranched hyoid barbels on the chin; and two dorsal fins. Snout is of moderate length. Nostril is paired. Eye is located near dorsal margin of head. Mouth is terminal and rather small, and upper jaw is slightly protrusible. Jaw teeth are in one to several series and are either villiform or conical. Teeth are present or absent in vomer and palatine. Hyoid barbel fits into median groove on throat. Maxilla is partially covered by suborbital bone. Supramaxilla is absent. Preopercular margin is entire and generally membranous. Operculum generally has flat spine on posterior margin. Branchiostegal rays number 4. Pectoral fin is rather high on flank and short, and has a nearly vertical base and 15 to 18 rays. Dorsal fins are widely separated; first consists of six to eight rather slender spines, and second consists of one spine and 8 or 9 rays. Pelvic fin is relatively large and thoracic and consists of one spine and 5 rays. Anal fin has one or two spines and 5 to 8 rays. Caudal fin is forked. Scales are cycloid, with attached spines on posterior margins. Axillary process is present on base of pelvic fin. Vertebrae number 24: 10 precaudal and 14 caudal. Maximum known size is 400 mm TL. Goatfishes occur worldwide in tropical to warm temperate seas. They are benthic on continental and insular shelves and occasionally occur in estuaries. Hyoid barbels possess chemosensory cells and are used to probe sediment and perceive small benthic organisms on which these fishes feed. Eggs, larvae, and juveniles are pelagic. There are about 55 species in six genera, and four species in separate genera occur in the western Central Atlantic and the Gulf of Mexico. references: Fritzsche and Fuiman 1982; Johnson 1984; Nelson 1994; Smith-Vaniz et al. 1999; Randall 2002a.

Key to the Species of the Gulf of Mexico (Adapted from Randall 2002a) 1a. Operculum with spine on margin; snout pointed; maxilla not extending to anterior margin of eye; no teeth on roof of mouth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Operculum without spine; snout blunt; maxilla reaching anterior margin of eye; teeth present on roof of mouth. . . 3 2a. Three dark blotches on lateral line; blue diagonal line on head; juveniles with a single series of jaw teeth, adults with two irregular series of teeth and a few caninelike teeth in upper jaw; lateral line scales 27 to 31 . . . . . . . . . . . . . . . . . . . . . Pseudupeneus maculatus p. 458

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448 PERCIFORMES

2b. Broad yellow stripe on side; three series of jaw teeth, all teeth small and blunt; lateral line scales 34 to 39 . . . . . . . . . . . . . . . . . . . . Mulloidichthys martinicus p. 449 3a. Teeth absent in upper jaw; first dorsal fin spines eight, first very small; dorsal profile of head strongly convex; interorbital space broad and flat; two broad yellow stripes on side . . . . . . . . . . . . . . . . . . . . . . . . . Mullus auratus p. 450 3b. Teeth present in upper jaw; first dorsal fin spines seven; dorsal profile of head angulated anterior to eye; interorbital space narrow and concave; several yellow stripes on side . . . . . . . . . . . . . . . . . . . . . . Upeneus parvus p. 452

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MULLIDAE 449

Mulloidichthys martinicus Yellow goatfish

(Cuvier, 1829)

Mulloidichthys martinicus is oblong, with a small spine on the opercular margin and a bright yellow stripe on side. Snout is relatively pointed, and dorsal profile of head is moderately convex. Posterior naris is slitlike and located close to anterior margin of orbit. Posterior margin of maxilla fails to reach anterior margin of orbit. Upper and lower jaw teeth are villiform and arranged in three rows anteriorly, two rows laterally, and one row posteriorly. Teeth are lacking in vomer and palatine. Gill rakers on first arch number 26 to 33. Measurements are expressed as percent of SL: head length 27%–28%, snout length 11%–13%, eye diameter 6%–7%, upper jaw length 8%– 10%, pectoral fin length 21%–22%, body depth 26%–28%. Pectoral fin rays number 15 to 17. First dorsal fin has eight spines, with first very short, and second dorsal fin has one spine and 8 rays. Anal fin has two spines and 6 rays. Lateral line scales number 34 to 39.

Color is olive gray to tan, darker dorsally and whitish ventrally, with bright yellow stripe extending from eye to caudal fin base. Median fins are yellow. This species occurs in coastal waters in the western Atlantic from southern Florida and Bermuda to Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is a schooling species associated with sandy bottoms near reefs. Food consists of small ray-finned fishes and benthic invertebrates, including polychaetes, bivalves, crustaceans, and brittle stars. Feeding takes place early in the morning. Maximum known size is 400 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996, 2002a; Hoese and Moore 1977, 1998; Matsuura 1983g; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1997; SmithVaniz et al. 1999.

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450 PERCIFORMES

Mullus auratus Red goatfish

Jordan and Gilbert, 1882

Mullus auratus is oblong in profile, with a broad, flat interorbital space and two distinct yellow to reddish bands on flank. Snout is relatively blunt, and dorsal profile of head is strongly convex. Posterior naris is slitlike and located close to anterior margin of orbit. Posterior margin of maxilla extends to anterior margin of orbit. Jaw teeth are villiform, arranged in a narrow band in lower jaw, and absent in upper jaw. Teeth are present in vomer and palatine. Opercular margin lacks spine. Gill rakers on first arch number 18 to 21. Measurements are expressed as percent of SL: head length 27%–29%, snout length 12%–13%, eye diameter 7%– 8%, upper jaw length 9%– 10%, pectoral fin length 19%–23%, body depth 25%–28%. Pectoral fin rays number 16 or 17. First dorsal fin has eight spines, with first minute. Second dorsal fin has one spine and 8 rays. Anal fin has two spines and 6 rays. Lateral line scales number 34 to 37. Color is bright scarlet or crimson, with two distinct yel-

low to reddish stripes on side. Upper stripe extends from opercular margin to end of second dorsal fin, and lower stripe extends from opercular margin to caudal fin base. Spinous dorsal fin is pale, with orange stripe at base and yellow to brownish stripe near tip, and second dorsal fin has series of yellowish to red spots forming stripe. This species occurs between 9 and 91 m in the western Atlantic from Cape Cod to Guyana, including the entire Gulf of Mexico and the Antilles. It is recorded from Bermuda, but these records have not been confirmed (SmithVaniz et al. 1999). This species is associated with muddy bottoms. Food consists of infaunal invertebrates. Maximum known size is 270 mm TL. references: Hoese and Moore 1977, 1998; Johnson 1978; Matsuura 1983g; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Cervigón 1993a,b; Randall 2002a.

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MULLIDAE 451

Pseudupeneus maculatus Spotted goatfish

(Bloch, 1793)

Pseudupeneus maculatus is oblong, with a small spine on the opercular margin and three dark blotches on lateral line. Snout is relatively pointed, and dorsal profile of head is slightly convex. Posterior naris is slitlike and located close to anterior margin of orbit. Posterior margin of maxilla fails to reach anterior margin of orbit. Jaw teeth are villiform and arranged in two irregular series, with some in outer row of upper jaw enlarged and curved outward. In juveniles, jaw teeth are in a single row. Teeth are lacking in vomer and palatine. Gill rakers on first arch number 26 to 32. Measurements are expressed as percent of SL: head length 30%–34%, snout length 16%–18%, eye diameter 6%–9%, upper jaw length 11%–12%, pectoral fin length 20%–24%, body depth 25%–28%. Pectoral fin rays number 13 to 16. First dorsal fin has eight spines, with first very short, and second dorsal fin has one spine and 8 rays. Anal fin has two spines and 6 rays. Lateral line scales number 27 to 31. Color is pale with edges of scales reddish, darker dor-

sally, and silvery on sides and ventrally, with diagonal blue lines on head. Anterior blackish blotch is located behind opercular margin, second is below first dorsal fin, and third is below second dorsal fin. Occasionally yellow stripes are present on lower flank. This species occurs in coastal waters to about 50 m in the western Atlantic from New Jersey and Bermuda to Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is associated with sandy and rocky bottoms near reefs. Juveniles are associated with grass beds. Food consists of sanddwelling polychaetes and crustaceans. Maximum known size is 300 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996, 2002a; Hoese and Moore 1977, 1998; Johnson 1978; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1997; SmithVaniz et al. 1999.

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452 PERCIFORMES

Upeneus parvus Dwarf goatfish

Poey, 1852

Upeneus parvus is oblong, with interorbital space narrow and concave and four or five oblique stripes on caudal fin. Snout is relatively blunt, and dorsal profile of head is moderately convex. Posterior naris is oval shaped and located close to anterior margin of orbit. Posterior margin of maxilla extends to anterior margin of orbit. Upper jaw teeth are villiform and arranged in two or three series anteriorly and a single series posteriorly. Lower jaw teeth are villiform and arranged in two or three series anteriorly and three or four rows posteriorly. Teeth are present in vomer and palatine. Gill rakers on first arch number 26 to 32. Measurements are expressed as percent of SL: head length 29%–31%, snout length 10%–11%, eye diameter 7%– 8%, upper jaw length 10%– 11%, pectoral fin length 21%–23%, body depth 23%–26%. Pectoral fin rays number 15 or 16. First dorsal fin has seven spines, with

first longest, and second dorsal fin has one spine and 8 rays. Anal fin has two spines and 6 rays. Lateral line scales number 36 to 40. Color is vermilion dorsally and silvery white ventrally, with yellow markings on flanks. Dorsal fins have bronze stripes, and caudal fin has four or five oblique orangish brown to dusky or black stripes. This species occurs between 40 and 100 m in the western Atlantic from North Carolina to Brazil, including the entire Gulf of Mexico and the Antilles but not the Bahamas. It is associated with muddy and sandy bottoms. Food consists of benthic invertebrates. Maximum known size is 200 mm TL. references: Hoese and Moore 1977, 1998; Matsuura 1983g; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1997; Randall 2002a.

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453

PEMPHERIDAE

Sweepers

Sweepers are oblong or ovate in profile, deep bodied, and very compressed, with a large terminal mouth, large eyes, and a single dorsal fin. Snout is acute. Nostril is paired. Mouth is terminal, oblique, and relatively large, with lower jaw slightly projecting beyond upper jaw. Upper jaw is slightly protrusible, and maxilla extends to about center of eye. Lachrymal bone does not cover maxilla when mouth is closed, and supramaxilla is absent. Jaw teeth are small, conical, and arranged in a single series or in a narrow band. Teeth occur in vomer and palatine. Four gill arches are present, and slit is present behind fourth arch. Branchiostegal rays number 7, and gill membranes are separate and free of isthmus. Pectoral fin is shorter than head, is pointed, and has 16 to 18 rays. Dorsal fin is single, originates anterior to and is shorter than anal fin, and has four to six increasingly larger spines and 7 to 13 rays. Spines are closely applied to front of dorsal fin. Pelvic fin inserts below pectoral fin base and has one spine and 5 rays. Anal fin has three small spines and 17 to 45 rays. Short axillary process at base of pelvic fin is present or absent. Caudal fin is truncate to shallowly forked. Body and most of head are covered with cycloid or ctenoid scales. Scales are absent from snout. Lateral line runs high along flank and extends onto caudal fin or to caudal fin margin is some species. Several species possess luminous organs. Vertebrae number 25: 10 precaudal and 15 caudal. Gas bladder is present. Maximum known size is about 150 mm TL. Sweepers occur over inner continental shelves in tropical to subtropical seas of the western Atlantic, Pacific, and Indian Oceans. They are common on coral reefs and in caves or other cavities but are seldom seen because they are nocturnally active. Individuals of species often form large schools. There are about 25 species in two genera, 2 species occur in the western Central Atlantic, and 1 of these occurs in the Gulf of Mexico. references: Fritzsche and Fuiman 1982; Johnson 1984; Nelson 1994; Smith 1997; Smith-Vaniz et al. 1999; R. Mooi (pers. comm., December 20, 2001).

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454 PERCIFORMES

Pempheris schomburgkii Müller and Troschel, 1848 Glassy sweeper

Pempheris schomburgkii is oblong in profile and deep bodied, with relatively large eyes and a long anal fin. Dorsal profile is slightly convex. Nostril is divided by flap. Preopercular margin is irregular, and opercular margin lacks spines. Gill rakers on first arch number 5 to 7 on upper limb and 16 to 20 on lower limb. Measurements are expressed as percent of SL: head length 31%–33%, snout length 6%– 8%, orbit diameter 12%–13%, upper jaw length 16%–17%, pectoral fin length 21%–27%, body depth 44%– 48%. Pectoral fin has slightly oblique base and 17 or 18 rays. Dorsal fin has five (rarely four or six) spines and 8 to 10 rays. Anal fin has 29 to 36 rays. Caudal fin is moderately forked. Lateral line is strongly arched anteriorly, runs near dorsal midline, and has 50 to 61 pored scales. Head and body are cop-

per colored, with a dark streak along base of anal fin. Juveniles are nearly transparent and silvery over abdomen. This species occurs from near shore to about 20 m in the western Atlantic from southeastern Florida and Bermuda to Brazil, including southern Florida and the Dry Tortugas in the Gulf of Mexico, the Bahamas, and the Antilles. It is associated with crevices and caves in reefs during the day and the water column above reefs at night. Food consists of zooplankton. Maximum known size is 72 mm SL. references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Robins and Ray 1986; Cervigón 1993a,b; Randall 1996; Smith 1997; Smith-Vaniz et al. 1999; R. Mooi 2002b.

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455

BATHYCLUPEIDAE

Bathyclupeids are elongate, deep anteriorly, and compressed, with large eyes, a single short dorsal fin, and a long anal fin. Dorsal profile is nearly straight. Head is very cavernous. Snout is short and oblique. Nostril is paired, and nares are close set and small. Eyes are very large and laterally located. Mouth is superior and very oblique, with lower jaw projecting beyond upper jaw. Upper jaw is bordered by both premaxilla and maxilla. Small villiform teeth occur in jaws, vomer, and palatine. Preoperculum and operculum are fragile and have entire margins. Four gill arches are present, and gill openings are large. Gill rakers on first arch are lathlike and range from 15 to 19. Gill membranes are separate and free of isthmus. Branchiostegal rays number 7. Pseudobranch is well developed. Pectoral fin is large and has 26 to 30 rays, with lower rays filamentous. Dorsal fin is short, consists of one spine and 8 to 10 rays, and is located posterior to midlength. Pelvic fin base is anterior to pectoral fin base, and fin is very small and has one spine and five rays. Anal fin is very long and has a single spine and 24 to 39 rays. Caudal fin is emarginate to forked. Body is covered with large, deciduous cycloid scales. Lateral line is complete and nearly straight. Vertebrae number 31: 9 or 10 precaudal and 20 or 21 caudal. Bathyclupeids occur in oceanic waters in the western Atlantic, western Pacific, and Indian Oceans at mesopelagic and bathypelagic depths. There are about four species in a single genus, and two species occur in the western Central Atlantic and the Gulf of Mexico. references: Fritzsche and Fuiman 1982; Johnson 1984; Heemstra 1986l; Nelson 1994; Paxton 2002.

Key to the Species of the Gulf of Mexico 1a. Anal fin with one spine and 36 to 39 rays . . . . . . . . . . . . . . . . . . . . . . Bathyclupea schroederi p. 457 1b. Anal fin with one spine and fewer than 34 rays . . . . . . . . . . . . . . . . . . . . . . . Bathyclupea argentea p. 456

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456 PERCIFORMES

Bathyclupea argentea

Goode and Bean, 1896

Bathyclupea argentea is elongate, compressed, and tapering posteriorly, with a relatively short anal fin. Snout is acute and shorter than eye diameter. Upper jaw extends slightly beyond anterior margin of orbit. Jaw teeth are conical, slightly recurved, and arranged in a single row. Vomerine and palatine teeth are of similar shape and are arranged in single rows. Gill rakers on first arch are long and slender and number 16 to 19, including rudiments. Measurements are expressed as percent of SL: head length 30%–34%, snout length 8%–10%, eye diameter 10%–13%, snout to dorsal fin origin 50%–59%, snout to anal fin origin 45%–

46%, body depth 20%–22%. Pectoral fin extends to beyond origin of anal fin and has 27 to 30 rays. Dorsal fin has 9 rays. Anal fin has a single spine and 28 to 31 rays. Caudal fin is forked. Lateral line has about 33 scales. Body is darkly pigmented. This species occurs between 366 and 640 m in the western Atlantic, including the northern and southern Gulf of Mexico and the Caribbean Sea. Maximum known size is about 211 mm SL. references: Goode and Bean 1896; Shimizu 1983b; Paxton 2002.

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BATHYCLUPEIDAE 457

Bathyclupea schroederi

Dick, 1962

Bathyclupea schroederi is elongate, compressed, and tapering posteriorly, with a relatively long anal fin. Snout is acute and shorter than eye diameter. Upper jaw extends slightly beyond anterior margin of orbit. Jaw teeth are conical, slightly recurved, and arranged in a single row. Vomerine and palatine teeth are of similar shape and are arranged in single rows. Gill rakers on first arch are long and slender and number 3 on upper limb and 16 on lower limb. Measurements are expressed as percent of SL: head length 29%–32%, snout length 8%–9%, eye diameter 11%–12%, snout to dorsal fin origin 51%–56%, snout to anal fin

origin 43%– 48%, body depth 23%–28%. Pectoral fin extends to beyond origin of anal fin and has 29 or 30 rays. Dorsal fin has 9 or 10 rays. Anal fin has a single spine and 36 to 39 rays. Lateral line has about 38 scales. Body is darkly pigmented. This species occurs between 366 and 549 m in the western Atlantic in the southern Gulf of Mexico and the Caribbean Sea. Maximum known size is about 144 mm SL.

references: Dick 1962; Shimizu 1983b; Paxton 2002.

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458

CHAETODONTIDAE

Butterflyfishes

Butterflyfishes are deep bodied, oval to orbicular or subrhomboid in profile, and strongly compressed, with a slightly to greatly prolonged snout and comblike jaw teeth. Dorsal profile is concave anterior to eye and straight to slightly convex posterior to eye. Nostril is paired, and nares are close set. Mouth is terminal and small, with upper jaw protrusible and posterior section of maxilla covered by lachrymal bone when mouth is closed. Jaw teeth are long and slender and arranged in brushlike bands. Vomer and palatine lack teeth. Preopercular margin is often serrated but lacks spine at angle. Gill membranes are narrowly attached to isthmus. Branchiostegal rays number 6. Gill rakers are short and not numerous. Pectoral fin is moderate to well developed, has strongly oblique base, and inserts on lower half of flank. Dorsal fin is single and entire or slightly notched, and has 6 to 16 spines and 15 to 30 rays, with membranes incised between spines. Pelvic fin has ancillary process, inserts below pectoral fin base, and has 1 spine and 5 rays. Anal fin has 3 to 5 spines and 14 to 23 rays. Caudal fin is rounded to emarginate and has 17 principal rays. Scales are ctenoid and small to large, and cover body and rayed sections of dorsal and anal fins. Lateral line either extends to caudal fin base or ends under rayed section of dorsal fin. Vertebrae number 24: 10 or 11 precaudal and 13 or 14 caudal. Gas bladder is present and has two anteriorly directed processes. Larval stages of most species (tholichthys) are covered with bony plates. Butterflyfishes occur worldwide in shallow tropical to subtropical seas. They are generally associated with coral reefs to a depth of 30 m, but some are found to 200 m and others occur in brackish or freshwater. Food generally consists of small invertebrates, such as coral polyps and copepods, and algae that are browsed from hard substrates. Some species are pelagic and feed on plankton. Juveniles of some species are cleaners, removing ectoparasites from larger fishes. Eggs and larvae are pelagic. There are about 114 species in 10 genera, 8 species in two genera in the western Central Atlantic, and 6 species in two genera in the Gulf of Mexico. references: Burgess 1974, 1978, 2002a; Johnson 1984; Heemstra 1986h; Nelson 1994; Smith-Vaniz et al. 1999.

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Key to the Species of the Gulf of Mexico (Adapted in part from Hoese and Moore 1998; Smith-Vaniz et al. 1999) 1a. Dark band extending from nape to eye and from eye to side of snout; snout 42% to 48% of head length . . . . . . . . . . . . . . . . . . . . . . Prognathodes aculeatus p. 468 1b. Dark band extending from nape through eye to ventral margin of cheek; snout length 26% to 37% of head length .............................................2 2a. Dorsal fin spines 13 or 14 (rarely 15) . . . . . . . . . . . . . . . . 3 2b. Dorsal fin spines 12 (rarely 13) . . . . . . . . . . . . . . . . . . . . 5 3a. Dorsal fin rays 18; anal fin rays 15 or 16; black band extending from base of spinous portion of dorsal fin to posterior portion of base of anal fin . . . . . . . . . . . . . . . . . . . . . . . . . . Prognathodes aya p. 470 3b. Dorsal fin rays 19 to 23; anal fin rays 16 to 19; no black band extending from base of spinous portion of dorsal fin to posterior portion of base of anal fin . . . . . . . . . . . . . . . 4 4a. Dorsal fin rays 19 or 20; anal fin rays 16 or 17; adults with large, black, white-edged spot on flank below posterior section of dorsal fin; juveniles with additional spot on posterior section of dorsal fin. . . Chaetodon capistratus p. 464 4b. Dorsal fin rays 21 to 23; anal fin rays 18 or 19; no ocellated spots on flank or on posterior section of dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon sedentarius 5a. Dorsal fin rays 21 or 22; side of body with four distinct dark bands. . . . . . . . . . . . . . . . . Chaetodon striatus p. 466 5b. Dorsal fin rays 19 to 21; side of body with one or two distinct dark bands . . . . . . . . . . . Chaetodon ocellatus p. 462

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460 PERCIFORMES

Chaetodon capistratus Foureye butterflyfish

Linnaeus, 1758

Chaetodon capistratus is deep bodied and compressed, with a short, pointed snout and a large ocellus on body below rayed section of dorsal fin. Dorsal profile is slightly concave from snout to eye and slightly convex from eye to dorsal fin origin. Jaws are slightly prolonged. Anterior naris is tubelike, and posterior naris is slightly elongated. Lachrymal bone, below eye, is free of scales and has a smooth margin. Preopercular margin is very finely serrated, with corner rounded. Operculum has broad-based spine on posterior margin. Gill rakers on lower limb of first arch are short and slender and number about 14 or 15. Measurements are expressed as percent of SL: head length 29%– 34%, snout length 9%–10%, eye diameter 9%–10%, upper jaw length 4%–7%, pectoral fin length 24%–25%, body depth 63%–77%.

Pectoral fin has 14 or 15 rays. Dorsal fin has 13 spines and 19 or 20 rays, with third or fourth spines longer than remainder. Anal fin has 3 spines and 16 or 17 rays, with second spine longest. Caudal fin is truncate. Lateral line is incomplete, forms high angular arc, and has 31 to 35 pored scales. Scales form deep sheath at base of dorsal fin. Color is whitish, with a dark bar running from nape through eye and lower section of interoperculum, and a large ocellus on upper part of body below rayed section of dorsal fin. Scale rows are marked with dark dashes forming pattern of diagonal lines that meet along midflank at about a 45 angle. Rayed sections of dorsal, anal, and caudal fins have dark submarginal stripes. Juveniles have an additional spot on posterior section of dorsal fin.

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This species occurs in shallow water in the western Atlantic from Massachusetts and Bermuda to northern South America, including the entire Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with shallow reefs. Food consists of gorgonians, tunicates, and polychaetes. Maximum known size is 80 mm SL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Haburay et al. 1969; Smith et al. 1975; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Burgess 1978, 2002a; Hastings 1979; Allen 1980; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Randall 1996; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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462 PERCIFORMES

Chaetodon ocellatus Bloch, 1787 Spotfin butterflyfish

Chaetodon ocellatus is deep bodied and compressed, with a short, pointed snout, one or two bars on body, and a large spot on dorsal fin. Dorsal profile is slightly concave from snout to eye and slightly convex from eye to dorsal fin origin. Jaws are slightly prolonged. Anterior naris has enlarged flap on posterior margin, and posterior naris is slightly elongated. Lachrymal bone, below eye, is free of scales and has a smooth margin. Preopercular margin is very finely serrated, with corner rounded. Operculum has a broad-based spine on posterior margin. Gill rakers on lower limb of first arch are short and slender and number about 14 or 15. Measurements are expressed as percent of SL: head length 34%–35%, snout length 11%–13%, eye diameter 9%– 10%, upper jaw length 8%–9%, pectoral fin length 24%–28%, body depth 63%– 80%.

Pectoral fin has 14 or 15 rays. Dorsal fin has 12 (rarely 13) spines and 19 to 21 rays, with fourth or fifth spines longer than remainder. Anal fin has 3 spines and 16 to 18 rays, with second spine longest. Caudal fin is truncate. Lateral line is incomplete, forms high angular arc, and has 33 to 34 pored scales. Scales form deep sheath at base of dorsal fin. Color is whitish yellow, with a black bar extending from nape through eye to lower section of interoperculum, a narrow yellow band extending from gill opening to pectoral fin base, and a large dull spot at base of rayed section of dorsal fin. Dull spot at base of dorsal fin becomes black during night and may disappear or become lighter than background during daylight. Lips and fins are yellowish. Males have a small black spot on posterior edge of dorsal fin. Juveniles have an additional black band extend-

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ing from margin of rayed section of dorsal fin to base of anal fin. This species occurs from near shore to 30 m in the western Atlantic from New Jersey and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is rare to absent in the Antilles. This species is associated with shallow reefs. Maximum known size is 150 mm SL.

references: Longley and Hildebrand 1941; Caldwell 1959; Böhlke and Chaplin 1968; Haburay et al. 1969; Hastings et al. 1976; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Burgess 1978, 2002a; Shipp and Hopkins 1978; Hastings 1979; Allen 1980; Matsuura 1983i; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Randall 1996; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999.

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464 PERCIFORMES

Chaetodon sedentarius Reef butterflyfish

Poey, 1860

Chaetodon sedentarius is deep bodied and compressed, with a short, pointed snout and two broad bars on body. Dorsal profile is slightly concave from snout to eye and slightly convex from eye to dorsal fin origin. Jaws are slightly prolonged. Anterior naris has enlarged flap on posterior margin, and posterior naris is slightly elongated. Lachrymal bone, below eye, is free of scales and has a smooth margin. Preopercular margin is very finely serrated, with a rounded corner. Operculum has a broad-based spine on posterior margin. Gill rakers on lower limb of first arch are short and slender and number 14 or 15. Measurements are expressed as percent of SL: head length 27%–33%, snout length 9%–11%, eye diameter 8%–10%, upper jaw length 6%–7%, pectoral fin length 21%–25%, body depth 59%–71%. Pectoral fin has 15 or 16 rays. Dorsal fin has 13 (rarely

14) spines and 21 to 23 rays, with fourth or fifth spines longer than remainder. Anal fin has 3 spines and 18 or 19 rays, with second spine longest. Caudal fin is truncate. Lateral line is incomplete, forms high angular arc, and has 36 to 39 pored scales. Body is cream colored, with one dark brown to black band extending from origin of dorsal fin through eye to lower edge of interoperculum, and another dark brown to black band extending from margin of rayed section of dorsal fin across caudal peduncle to rear edge of anal fin. Caudal fin is light yellow with a white base. Dorsal section of body may be yellowish. This species occurs between 15 and 40 m in the western Atlantic from North Carolina and Bermuda to northern Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with

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CHAETODONTIDAE 465

shallow reefs. Food consists of hydroids, polychaetes, and amphipods. Maximum known size is 150 mm SL. references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Bright and Cashman

1974; Hastings et al. 1976; Hoese and Moore 1977, 1998; Burgess 1978, 2002a; Shipp and Hopkins 1978; Hastings 1979; Allen 1980; Matsuura 1983i; Robins and Ray 1986; Cervigón 1993b; Randall 1996; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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466 PERCIFORMES

Chaetodon striatus Linnaeus, 1758 Banded butterflyfish

Chaetodon striatus is deep bodied and compressed, with a short, pointed snout and four broad bands on body. Dorsal profile is concave from snout to eye and moderately convex from eye to dorsal fin origin. Jaws are slightly prolonged. Anterior naris has enlarged flap on posterior margin, and posterior naris is slightly elongated. Lachrymal bone, below eye, is free of scales and has a smooth margin. Preopercular margin is very finely serrated, with corner slightly produced and rounded. Operculum has broad-based spine on posterior margin. Gill rakers on lower limb of first arch are short and slender and number 13 or 14. Measurements are expressed as percent of SL: head length 34%–37%, snout length 11%–13%, eye diameter 10%–12%, upper jaw length 8%– 9%, pectoral fin length 26%–28%, body depth 65%–70%. Pectoral fin has 15 or 16 rays. Dor-

sal fin has 12 spines and 21 or 22 rays, with fourth or fifth spines longer than remainder. Anal fin has 3 spines and 16 to 18 rays, with second spine longest. Caudal fin is truncate. Lateral line is incomplete, forms high angular arc, and has 35 or 36 pored scales. Body is whitish, with first dark band on body extending from nape through eye to margin of interoperculum, second extending from origin of dorsal fin to belly, third extending from margin of spiny section of dorsal fin to margin of anal fin, and fourth extending from rayed section of dorsal fin through caudal peduncle to base of anal fin. Submarginal dark band runs disjunctly along distal margin of dorsal fin, midsection of caudal fin, and posterior margin of anal fin. Juveniles have an ocellated spot on rayed section of dorsal fin. This species occurs in the tropical Atlan-

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tic between 3 and 20 m. In the western Atlantic it occurs from Massachusetts (juveniles) or North Carolina (adults) and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. It is associated with shallow reefs. Food consists of coral polyps, polychaetes, crustaceans, and mollusk eggs. Maximum known size is 150 mm SL.

references: Longley and Hildebrand 1941; Caldwell and Briggs 1957; Caldwell 1959; Böhlke and Chaplin 1968; Haburay et al. 1969; Randall 1968a, 1996; Bright and Cashman 1974; Smith et al. 1975; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Burgess 1978, 2002a; Hastings 1979; Allen 1980; Robins and Ray 1986; Boschung 1992; Cervigón 1993b; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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468 PERCIFORMES

Prognathodes aculeatus Longsnout butterflyfish

(Poey, 1860)

Prognathodes aculeatus is deep bodied and compressed, with a long, beaklike snout and a dark bar between eye and snout. Dorsal profile is strongly concave from snout to eye and almost straight from eye to dorsal fin origin. Jaws resemble forceps. Anterior naris is tubelike, and posterior naris is elongated. Preopercular margin is finely serrated, with corner produced and rounded. Operculum has a broad-based spine on posterior margin. Gill rakers on lower limb of first arch are short and slender and number about 15. Measurements are expressed as percent of SL: head length 38%–39%, snout length 14%–15%, eye diameter 9%–11%, upper jaw length about 10%, pectoral fin length 27%–28%, body depth 56%– 63%. Pectoral fin has 13 to 15 rays. Dorsal fin has 13 spines and 18 or 19 rays, with third, fourth, and fifth spines longer than remainder, making dorsal fin margin deeply incised. Anal fin has 3 spines

and 14 to 16 rays, with third spine longest. Caudal fin is truncate. Lateral line is incomplete, forms high angular arc, and has 22 to 28 pored scales. Scales form low sheath at base of dorsal fin. Color is golden or bronze yellow dorsally, shading to black on dorsal fin base, and pale to white ventrally. Dark brown bands run from snout to anterior margin of eye and from dorsal margin of eye to nape. Orange band runs from eye to side of snout, and orange bar is located on caudal peduncle. This species occurs between 1 and 60 m in the western Atlantic from southern Florida and Bermuda to northern South America, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It occurs in the southern and northern Gulf of Mexico. This species is most common over deep reefs. Food consists of tentacles of tube-dwelling polychaetes, pedicellariae of sea urchins, and

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small crustaceans. Maximum known size is 80 mm SL. references: Böhlke and Chaplin 1968; Bright and Cashman 1974 (as Chaetodon aculeatus); Hoese

and Moore 1977, 1998 (as C. aculeatus); Burgess 1978 (as C. aculeatus); Allen 1980 (as C. aculeatus); Robins and Ray 1986 (as C. aculeatus); Randall 1996; Schaldach et al. 1997 (as C. aculeatus); Smith 1997 (as C. aculeatus); Smith-Vaniz et al. 1999 (as C. aculeatus).

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470 PERCIFORMES

Prognathodes aya Bank butterflyfish

(Jordan, 1886)

Prognathodes aya is deep bodied and compressed, with a long beaklike snout and two broad, diagonal blackish bands on body. Dorsal profile is strongly concave from snout to eye and slightly convex from eye to dorsal fin origin. Jaws resemble forceps. Anterior naris is tubelike, and posterior naris is elongated. Preopercular margin is finely serrated, with corner produced and rounded. Operculum has a broad-based spine on posterior margin. Gill rakers on lower limb of first arch are short and slender and number about 14 or 15. Measurements are expressed as percent of SL: head length 35%–38%, snout length 14%–16%, eye diameter 9%–10%, upper jaw length 9%– 10%, pectoral fin length 21%–25%, body depth 54%–58%. Pectoral fin has 14 or 15 rays. Dorsal fin has 13 spines and 18 rays, with third, fourth, and fifth spines longer than remainder, making dorsal fin margin deeply incised. Anal fin has 3 spines and 15 or 16 rays, with second and third spines about equal in length. Caudal fin is truncate. Lateral line is

incomplete, forms high angular arc, and has about 26 pored scales. Scales form low sheath at base of dorsal fin. Color is white to yellowish white, with a narrow black bar extending from below lower jaw through eye to dorsal fin origin and a broad black bar extending from middle of spiny section of dorsal fin to posterior section of anal fin base. Bars form inverted-V-shaped pattern on body. Pelvic fin, rayed section of dorsal fin, anal fin, and caudal fins are often yellow. This species occurs between 20 and 167 m in the western Atlantic from North Carolina to Quintana Roo in the northwestern Caribbean Sea, including the northern and southern Gulf of Mexico. It is associated with rocky bottoms. Maximum known size is 150 mm SL. references: Longley and Hildebrand 1941; Hoese and Moore 1977, 1998; Burgess 1978, 2002a; Allen 1980; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; SmithVaniz et al. 1999.

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471

POMACANTHIDAE

Angelfishes

Angelfishes are oval to subrhomboid in profile, deep bodied, and strongly compressed, with a strong spine at angle of preoperculum and comblike jaw teeth. Dorsal profile is generally straight or convex anterior to eye and straight to slightly convex posterior to eyes. Mouth is terminal and small, with upper jaw slightly protrusible. Nostril is paired. Jaw teeth are long and slender, generally tricuspid, and arranged in brushlike bands. Vomer and palatine lack teeth. Preopercular margin is often serrated above spine at angle. Gill membranes are free of isthmus. Branchiostegal rays number 6. Gill rakers are short and not numerous. Pectoral fin is moderate to well developed and inserts on lower half of flank at a strongly oblique angle. Dorsal fin is entire and has 9 to 15 spines and 15 to 37 rays. Pelvic fin lacks ancillary process, inserts below pectoral fin base, and has 1 spine and 5 rays. Anal fin has 3 spines and 14 to 25 rays. Dorsal and anal fins bear elongate posterior extensions in some species. Caudal fin is rounded to strongly emarginate and has 17 principal rays. Scales are ctenoid, and ctenii are continuous with posterior margin of scale and cover body and rayed section of dorsal and anal fin. Vertebrae number 24: 10 or 11 precaudal and 13 or 14 caudal. Gas bladder is present and lacks anteriorly directed processes. Larval stage lacks bony plates. Angelfishes occur worldwide in shallow tropical to subtropical seas. They are generally associated with coral reefs and are seldom found below 50 m. Food generally consists of small invertebrates, such as coral polyps and copepods, and algae that are browsed from hard substrates. All species have bright, striking color patterns, and juveniles are often very differently patterned than adults. Eggs and larvae are pelagic. There are 74 species in nine genera, 7 species in three genera in the western Central Atlantic, and 6 species in three genera in the Gulf of Mexico. references: Burgess 1974, 1978, 2002b; Johnson 1984; Nelson 1994.

Key to the Species of the Gulf of Mexico (Adapted from Hoese and Moore 1998; Burgess 2002b) 1a. Dorsal fin spines 9 or 10 . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Dorsal fin spines 12 to 15. . . . . . . . . . . . . . . . . . . . . . . . . 3 2a. Dorsal fin spines 10; dorsal fin rays 29 to 31; caudal fin convex; adults with posterior margin of caudal fin dark; juveniles with a yellow stripe on midline of head extending to upper lip and a round dark spot on caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . . Pomacanthus paru p. 481 2b. Dorsal fin spines 9; dorsal fin rays 31 to 33; caudal fin convex in juveniles and truncate in adults; adults with poste-

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472 PERCIFORMES

3a.

3b.

4a.

4b.

5a.

5b.

rior margin of caudal fin light; juveniles with a yellow stripe on midline of head extending to chin and a semicircular to rectangular dark spot on caudal fin . . . . . . . . . . . . . . . . . . . . . . . Pomacanthus arcuatus p. 479 Dorsal fin rays 15 or 16; preorbital bone beneath eye with 2 or 3 posteriorly directed spines; caudal fin bluish black . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centropyge argi p. 473 Dorsal fin rays 17 to 21; preorbital bone beneath eye angular or with 1 blunt spine; caudal fin yellow or with yellow margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Adults with black body posterior to pectoral fin base and upper margin of caudal fin elongated; juveniles (30 mm SL) with body mostly yellow and large black spot on upper side . . . . . . . . . . . . . . . . . . . . . Holacanthus tricolor p. 478 Adults without black body posterior to pectoral fin base and without upper margin of caudal fin elongated; juveniles (45 mm SL) with body mostly dark blue and narrow bluish white bars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Adults mostly tan to purplish, body scales with pale edges, proximal half of pectoral fin blue, no black markings on body or fins, and posterior margin of caudal fin yellow; juveniles dark blue, with bluish white bars on head and body and second bar straight . . . . . . . . . . . . . . . . . . . . Holacanthus bermudensis p. 474 Adults bluish to greenish, body scales with orange spots or triangles, large black blotch on pectoral fin base and on last rays of dorsal and anal fins, and caudal fin uniformly yellow; juveniles dark blue, with light blue bars on head and body and second bar curved . . . . . . . . . . . . . . . . . . . . . . . . . Holacanthus ciliaris p. 476

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POMACANTHIDAE 473

Centropyge argi Cherubfish

Woods and Kanazawa, 1951

Centropyge argi is oval in profile, moderately deep bodied, and strongly compressed, with 2 or 3 large spines on preorbital bone and a narrow blue ring around eye. Dorsal profile of head is slightly convex. Snout is blunt. Anterior and posterior nares are close set. Maxilla fails to reach or just reaches anterior margin of orbit. Margin of preoperculum is strongly serrated, and a small spine is located on ventral margin, below large spine on corner. Operculum has a very broad, triangular spine on margin. Gill rakers on first arch number 22 to 24. Measurements are expressed as percent of SL: head length 28%–33%, snout length 7%– 8%, eye diameter 7%–10%, upper jaw length 5%– 7%, pectoral fin length 22%–27%, body depth 50%–56%. Pectoral fin has 15 or 16 rays. Dorsal fin has 14 or 15 spines, with margin behind first 4 spines deeply incised, and 15 or 16 rays. Anal fin has 17 rays, and membranes are deeply incised between first two spines. Dorsal and anal fins are not extended as fila-

mentous lobes. Caudal fin is rounded. Lateral line extends to rayed section of dorsal fin and consists of 32 to 34 pored scales. Color is dark blue, with yellowish or yellowish orange snout, cheek, chest, and pectoral fins. Eye is surrounded by a blue ring, and a blue smudge is located behind corner of mouth. This species occurs from near shore to 30 m in the western Atlantic from West Palm Beach, FL, and Bermuda to northern South America, including the northern and southern Gulf of Mexico, the Florida Keys, the Bahamas, and the Antilles. Food is thought to be algae. Maximum known size is 50 mm TL.

references: Woods and Kanazawa 1951; Böhlke and Chaplin 1968; Randall 1968a, 1996; Feddern 1972; Hoese and Moore 1977, 1998; Rezak et al. 1985; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Burgess 2002b.

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474 PERCIFORMES

Holacanthus bermudensis Blue angelfish

Goode, 1876

Holacanthus bermudensis is subrhomboid in profile, deep bodied, and strongly compressed, with posterior filamentous extensions of dorsal and anal fins, and caudal fin margin yellow. Dorsal profile of head is nearly straight to center of eye and moderately convex from eye to origin of dorsal fin. Snout is blunt. Anterior naris has flap on posterior margin, and posterior naris is elongated. Margin of preorbital bone is serrated. Maxilla fails to reach anterior margin of orbit. Margin of preoperculum has a series of small spines on margin, with spines increasing slightly in size toward corner. Operculum has a very broad, triangular spine on margin. Gill rakers on first arch number 18 to 21. Measurements are expressed as percent of SL: head length 30%–32%, snout length 11%–13%, eye diameter 7%– 8%, upper jaw length 8%–9%, pectoral fin length 23%–25%, body depth 53%– 67%. Pectoral fin has 19 rays. Dorsal fin has 14 spines, with margin behind first 2 spines deeply incised, and 19 to 21 rays. Anal fin has 19 to 21 rays, with membrane between first 2 spines deeply incised.

Caudal fin is rounded. Lateral line extends to rear portion of dorsal fin and consists of 45 to 49 pored scales. Color is bluish tan, scales on side of body have yellow edges, and chest and nape are deep purplish blue. Proximal half of pectoral fin is blue, and dorsal and anal fins are blue tipped, except for attenuated distal extensions that are yellow. Caudal fin has yellow margin. Juveniles are dark blue, with snout, chest, pectoral fin, and pelvic fin yellow. Rear edges of dorsal and anal fin change to yellow with growth. Four long bluish white bars occur on head and body, and second is straight. This species occurs in shallow water to 60 m in the western Atlantic from New Jersey (rare) or North Carolina and Bermuda to Yucatán, including the Gulf of Mexico, the Florida Keys, and the Bahamas. It is associated with coral reefs and oil platforms. Maximum known size is 450 mm TL. references: Longley and Hildebrand 1941 (as Holocanthus isabelita); Böhlke and Chaplin

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POMACANTHIDAE 475

1968; Feddern 1968, 1972; Randall 1968a (as Holocanthus isabelita), 1996; Haburay et al. 1969; Bright and Cashman 1974; Hastings et al. 1976; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Allen 1980; Burgess 1978 (as Holocanthus isa-

belita), 2002b; Shipp and Hopkins 1978; Hastings 1979; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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476 PERCIFORMES

Holacanthus ciliaris Queen angelfish

(Linnaeus, 1758)

Holacanthus ciliaris is subrhomboid in profile, deep bodied, and strongly compressed, with posterior corners of dorsal and anal fins filamentous, and a large black blotch on nape. Dorsal profile of head is nearly straight from snout to center of eye and moderately convex from eye to origin of dorsal fin. Snout is blunt. Anterior naris has flap on posterior margin, and posterior naris is elongated. Margin of preorbital bone is serrated. Maxilla fails to reach anterior margin of orbit. Margin of preoperculum has a series of small spines on margin, with spines increasing slightly in size toward corner. Operculum has a very broad, triangular spine on margin. Gill rakers on first arch number 18 to 21. Measurements are expressed as percent of SL: head length 29%–33%, snout length 11%–13%, eye diameter 9%–10%, upper jaw length 7%– 8%, pectoral fin length 24%–31%, body depth 53%– 67%. Pectoral fin has 18 to 20 rays. Dorsal fin has 14 spines, with margin behind first 2 spines deeply in-

cised, and 19 to 21 rays. Anal fin has 20 or 21 rays, and membrane between first 2 spines is deeply incised. Caudal fin is rounded. Lateral line extends to rear portion of dorsal fin and consists of 45 to 49 pored scales. Color is deep blue to greenish, with an orange spot or triangle on each scale, a yellow to orange caudal fin, and a yellow pelvic fin. A blue-ringed black spot, speckled with blue pigment, is located on nape. Large black blotches are located on base of pectoral fin and on bases of posterior rays of dorsal and anal fins. Tips of dorsal and anal fin rays are orange. Juveniles are dark blue, with snout, chest, pectoral fin, and pelvic fin yellow. Four long bluish white bars occur on head and body, first three are curved, and last is straight. This species occurs in shallow water to 60 m in the western Atlantic from North Carolina and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with

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POMACANTHIDAE 477

coral reefs and other hard structures. Maximum known size is 450 mm TL. references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Feddern 1968, 1972;

Randall 1968a, 1996; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Burgess 1978, 2002b; Allen 1980; Matsuura 1983j; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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478 PERCIFORMES

Holacanthus tricolor Rock beauty

(Bloch, 1795)

Holacanthus tricolor is subrhomboid in profile, deep bodied, and strongly compressed, with short filaments extending from dorsal and anal fins, and anterior section of body yellow and posterior section black. Dorsal profile of head is straight to slightly convex. Snout is moderately blunt. Anterior naris has flap on posterior margin, and posterior naris is elongated. Margin of preorbital bone is serrated. Maxilla fails to reach anterior margin of orbit. Margin of preoperculum has series of small spines on margin. Operculum has a very broad, triangular spine on margin. Gill rakers on first arch number 17 to 20. Measurements are expressed as percent of SL: head length 27%– 34%, snout length about 10%, eye diameter 6%–10%, upper jaw length 7%– 8%, pectoral fin length 21%–25%, body depth 53%– 67%. Pectoral fin has 17 or 18 rays. Dorsal fin has 14 spines, with margin behind first 4 spines deeply incised, and 17 to 19 rays. Anal fin has 18 to 20 rays, with membrane between first 2 spines deeply incised. Caudal fin is rounded, with small filament extending from upper and lower rays. Lateral line extends to rear portion of dorsal fin and consists of 43 to 46 pored

scales. Head, nape, and anterior part of body are bright yellow; lips and remainder of body are black. Posterior margins of dorsal and anal fins and entire caudal fin are bright yellow. Upper and lower sections of iris are bright blue. Pectoral fin and pelvic fin are yellow. Juveniles are yellow, with a large black spot surrounded by a blue ring below rayed section of dorsal fin. With growth, black coloration extends beyond spot until adult coloration is achieved. This species occurs in shallow water in the western Atlantic from North Carolina and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. It is associated with coral reefs and other hard structures. Juveniles are associated with colonies of stinging coral (Millepora). Maximum known size is 250 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Burgess 1978, 2002b; Allen 1980; Williams and Shipp 1980; Matsuura 1983j; Robins and Ray 1986; Boschung 1992; Smith-Vaniz et al. 1999.

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POMACANTHIDAE 479

Pomacanthus arcuatus (Linnaeus, 1758) Gray angelfish

Pomacanthus arcuatus is oval in profile, deep bodied, and strongly compressed, with long filaments extending from dorsal and anal fins, scales irregular in size and shape, and larger scales dark at base and light at margin. Dorsal profile of head is slightly convex. Snout is moderately blunt. Anterior naris has flap on posterior margin, and posterior naris is oval. Maxilla fails to reach anterior margin of orbit, and lower jaw projects beyond upper jaw. Margin of preoperculum has a series of very small spines on margin. Operculum has a very broad, rounded process on margin. Gill rakers on first arch number 17 to 19. Measurements are expressed as percent of SL: head length 27%–31%, snout length 11%–13%, eye diameter 7%–10%, upper jaw length about 8%, pectoral fin length 23%–25%, body depth 71%–77%. Pectoral fin has 19 or 20 rays.

Dorsal fin has nine spines and 31 to 33 rays. Anal fin has 23 to 25 rays. Posterior margins of dorsal and anal fins are evenly convex. Caudal fin is convex in juveniles and truncate, with acute upper and lower corners, in adults. Lateral line is moderately arched and extends to base of caudal fin. Color is gray to brownish, with lips and chin cream colored; medial side of pectoral fin yellowish; and posterior margins of dorsal, anal, and caudal fins yellow to transparent. Juveniles are blackish to dark brown, with four yellow bands on head and body, a median yellow stripe on nape extending to chin, and a rectangular to hemispherical black spot on caudal fin. Yellow bands may persist in small adults. This species occurs in shallow water in the western Atlantic from New York to Brazil, including the northern and southern Gulf of

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480 PERCIFORMES

Mexico, the Bahamas, and the Antilles. It is associated with coral reefs and other hard structures with high relief. Juveniles clean other fishes of ectoparasites. Maximum known size is 600 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a; Hoese and Moore 1977, 1998; Burgess 1978, 2002b; Allen 1980; Matsuura 1983j; Robins and Ray 1986; Boschung 1992; Smith 1997; Schaldach et al. 1997.

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POMACANTHIDAE 481

Pomacanthus paru (Bloch, 1787) French angelfish

Pomacanthus paru is oval in profile, deep bodied, and strongly compressed, with long filaments extending from dorsal and anal fins, scales irregular in size and shape, and yellow margins on most scales. Dorsal profile of head is slightly convex. Snout is moderately blunt. Anterior naris has flap on posterior margin, and posterior naris is oval. Maxilla fails to reach anterior margin of orbit, and lower jaw extends beyond upper jaw. Margin of preoperculum has a series of very small spines on margin. Operculum has very broad, rounded process on margin. Gill rakers on first arch number 17 to 19. Measurements are expressed as percent of SL: head length 25%–33%, snout length 11%–12%, eye diameter 6%–9%, upper jaw length 7%–9%, pectoral fin length 24%–28%, body depth 73%–78%. Pectoral fin has 19 or 20 rays. Dorsal fin has 10 spines and 29 to 31 rays. Anal fin has 22 to 24 rays. Posterior margins of dorsal and anal fins are evenly convex. Caudal fin is rounded. Lateral line is moderately arched and extends to base of caudal fin. Color is black except for golden yellow margins of scales on side of body. Base

of pectoral fin has yellow blotch or bar, filament of dorsal fin is yellow, and caudal fin has dark margin. Juveniles are blackish to dark brown, with four yellow bands on head and body, a median yellow stripe on nape extending to upper lip, a circular black spot on caudal fin, and a yellow margin on caudal fin. Yellow bands may persist on small adults. This species occurs in the western and Central Atlantic at Ascension Island. In the western Atlantic it occurs in shallow water from North Carolina and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. It is associated with coral reefs and other hard substrates with vertical relief. Juveniles clean other fishes of ectoparasites. Maximum known size is 400 mm TL.

references: Longley and Hildebrand 1941 (as Pomacanthus aureus); Böhlke and Chaplin 1968; Randall 1968a, 1996; Haburay et al. 1969; Bright and Cashman 1974; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Burgess 1978, 2002b; Allen 1980; Matsuura 1983j; Robins and Ray 1986; Schaldach et al. 1997; Smith-Vaniz et al. 1999.

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482

KYPHOSIDAE

Sea chubs

Sea chubs are oblong to oval in profile, moderately deep to deep, and moderately compressed, with a short head and an outer series of incisor-like teeth possessing deep roots. Dorsal profile of head is convex, and snout is blunt. Mouth is terminal, small, and horizontal. Upper jaw is moderately protrusible. Maxilla is covered by lachrymal bone when mouth is closed. Jaw teeth are thin, small, sharp, and arranged in bands. Teeth in outer row are narrow and blunt, and teeth in inner row are small and weak. Teeth also occur in vomer and palatine and occasionally on tongue. Preoperculum has a membranous margin, and operculum has a broad-based spine on posterior margin. Gill membranes are united and joined to isthmus. Branchiostegal rays number 7. Pectoral fin is relatively small, either smaller than or equal in size to pelvic fin, and inserts on lower half of flank. Dorsal fin is continuous and has 6 to 11 spines and 11 to 15 rays. Spinous section of dorsal fin is depressible into scaly groove. Pelvic fin inserts slightly behind pectoral fin insertion and has 1 spine and 5 rays. Anal fin has 3 spines and 10 to 14 rays. Caudal fin is emarginate to lunate and has 17 principal rays. Scales are ctenoid and cover body, head (except for snout), and rayed section of dorsal fin and anal fin. Scales on body are larger than those on head and on dorsal and anal fins. Maxilla is also scaled. Vertebrae number 25: 10 precaudal and 15 caudal. Gas bladder is divided into anterior and posterior sections and extends to tail. Maximum known size is 760 mm TL. Sea chubs occur in shallow tropical and subtropical seas worldwide. They form schools and are associated with coral reefs and rocky areas. Food consists of macroalgae. Juveniles are pelagic and are associated with floating seaweed. There are 11 species in four genera, and 2 species in the same genus occur in the western Central Atlantic and the Gulf of Mexico. references: Fritzsche and Fuiman 1982; Johnson 1984; Johnson and Fritzsche 1989; Nelson 1994; Sommer 1995; Smith 1997; Carpenter 2002c.

Key to the Species of the Gulf of Mexico (Adapted from Smith-Vaniz et al. 1999) 1a. Dorsal fin rays 11 to 13; anal fin rays 11 (rarely 12); total dorsal and anal fin rays 23 or 24 (rarely 22); gill rakers on first arch 22 to 27 in specimens greater than 50 mm SL . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus sectator p. 484 1b. Dorsal fin rays 13 to 15; anal fin rays 12 or 13; total dorsal and anal fin rays 25 to 28; gill rakers on first arch 27 to 30 in specimens greater than 50 mm SL . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus incisor p. 483

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KYPHOSIDAE 483

Kyphosus incisor Yellow chub

(Cuvier, 1831)

Kyphosus incisor is oval in profile and moderately compressed, with a short head, longitudinal brassy stripes on body, and a yellowedged whitish streak below eye. Dorsal profile of head is slightly convex. Nostril is paired, and nares are oblong. Villiform teeth are present on tongue. Gill rakers on first gill arch number 6 to 8 on upper limb and 19 to 22 on lower limb. Measurements are expressed as percent of SL: head length 28%–29%, snout length 8%–10%, eye diameter 7%– 8%, upper jaw length 5%–9%, pectoral fin length 17%–18%, body depth 45%– 46%. Pectoral fin has 18 to 20 rays. Dorsal fin has 11 spines and 13 to 15 rays. Anal fin has 12 or 13 rays. Caudal fin is forked. Lateral line scales number 54 to 62. Rayed sections of dorsal fin and other fins are densely covered with small scales. Color is dusky bluish, with bright, brassy yellow lines along scale rows. Opercular membrane is

slightly pigmented. Juveniles have large whitish spots on body. This species occurs in the tropical and warm temperate Atlantic. In the western Atlantic it occurs in shallow water from Cape Cod and Bermuda to Brazil, including the entire Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with rocky bottoms and coral reefs. Juveniles often occur in floating Sargassum rafts. Food consists mostly of benthic algae and Sargassum. Maximum known size is 670 mm TL.

references: Longley and Hildebrand 1941; Moore 1962; Böhlke and Chaplin 1968; Randall 1968a, 1996; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Carpenter 2002c.

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484 PERCIFORMES

Kyphosus sectator Bermuda chub

(Linnaeus, 1758)

Kyphosus sectator is oval shaped and moderately compressed, with a short head, longitudinal straw yellow stripes on body, and a yellow line below eye. Dorsal profile of head is slightly convex. Nostril is paired, and nares are oblong. Villiform teeth are present on tongue. Gill rakers on first gill arch number 6 to 8 on upper limb and 16 to 18 on lower limb. Measurements are expressed as percent of SL: head length 30%–33%, snout length 8%–11%, eye diameter 9%–11%, upper jaw length 8%– 10%, pectoral fin length 17%–20%, body depth 43%–53%. Pectoral fin has 17 to 19 rays. Dorsal fin has 11 spines and 11 to 13 (usually 12) rays. Anal fin has 10 to 12 rays. Caudal fin is forked. Lateral line scales number 51 to 58. Rayed sections of dorsal fin and other fins are densely covered with small scales. Color is dusky bluish, with straw yellow lines along scale rows; yellow line under eye extends from corner of mouth to margin of preoperculum. Upper section of opercular membrane

is blackish. Juveniles have large pale spots on head, body, and unpaired fins. This species occurs in the tropical and warm temperate seas of the Atlantic. In the western Atlantic it occurs in shallow water from Nova Scotia and Bermuda to Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is found throughout the Gulf of Mexico in shallow water over turtle-grass beds, sandy bottoms, and coral reefs. Juveniles often occur in floating Sargassum rafts. Food consists mostly of benthic algae and Sargassum. Maximum known size is 760 mm TL.

references: Longley and Hildebrand 1941; Moore 1962; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hastings et al. 1976; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Hastings 1979; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Carpenter 2002c.

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485

CIRRHITIDAE

Hawkfishes

Hawkfishes are oval to oblong and moderately compressed, with pelvic fin insertion relatively far behind pectoral fin insertion and lower rays of pectoral fin unbranched, usually thickened, and occasionally separate from each other. Nostril is paired, and anterior naris bears cirri on posterior margin. Mouth is terminal and moderate in size. Jaw teeth consist of an outer row of canines and an inner band of villiform teeth. Anterior upper jaw teeth are larger than posterior teeth, and the largest teeth in lower jaw are located along midlength of jaw. Vomer has teeth, and palatine either has or lacks teeth. Operculum has 2 weak, flat spines on margin. Gill membranes are broadly joined and have a free fold across isthmus. Branchiostegal rays number 6. Pectoral fin has 14 rays, and lower 5 to 7 rays are unbranched and usually thickened and deeply incised. Dorsal fin is single, is notched between spinous and rayed sections, and consists of 10 spines and 11 to 17 rays, with spinous section longer than rayed section. One or more cirri project from interspinous membranes near tips of dorsal spines. Pelvic fin has 1 spine and 5 rays. Anal fin is rather short and consists of 3 spines and 5 to 7 rays. Caudal fin has 15 principal rays. Scales are cycloid. Vertebrae number 26 to 28. Gas bladder is absent. Maximum size is 30 cm TL. Hawkfishes occur worldwide in tropical seas, but few species occur outside the Indo-West Pacific Ocean. They are benthic in shallow water. Thickened lower rays of pectoral fin are used for support and to wedge themselves into substrate in areas subjected to wave action. Food consists of benthic invertebrates and rayfinned fishes. There are 38 species in 10 genera, and 1 species occurs in the western Central Atlantic and in the Gulf of Mexico. references: Randall 1963b, 1986a, 2001b, 2002b; Nelson 1994; Tominaga et al. 1996; Smith-Vaniz et al. 1999.

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486 PERCIFORMES

Amblycirrhitus pinos Redspotted hawkfish

(Mowbray, 1927)

Amblycirrhitus pinos has an acute snout, a tuft of cirri behind each dorsal fin spine, and a scaled interorbital region. Dorsal profile from snout to eye is nearly straight. Mouth is slightly oblique, upper jaw slightly overhangs lower jaw, and maxilla extends to center of pupil. Preopercular margin is finely serrated and evenly rounded. Palatine teeth are present. Gill rakers on first arch number 4 or 5 on upper limb and 9 to 11 on lower limb. Measurements are expressed as percent of SL: head length 33%– 36%, snout length 9%–10%, eye diameter 7%– 8%, upper jaw length 11%–12%, pectoral fin length 34%– 40%, pelvic fin length 17%–22%, body depth 35%–38%. Pectoral fin consists of 9 upper rays united by membrane and 5 lower rays that are thickened and connected by membranes only along medial half of ray. Dorsal fin consists of 10 spines and 11 rays, with membrane deeply incised between spines. Anal fin has 6 rays. Lateral line scales number 41 to 44, scales between base of spinous dorsal fin and lateral line number 4, and scales between anal fin base and lateral line number 9. Cheek has four or five rows of scales.

Color is olive with five broad white bars, and last bar is on caudal fin base. The first three bars and the fifth one are bisected by yellowish brown bars. Large black spot is located on base of rayed section of dorsal fin, and the last two white bars are separated by a black ring on caudal peduncle. Head, anterior part of body, and dorsal fin are covered with small orangish red spots. This species occurs to a depth of 45 m in the western Atlantic from southern Florida and Bermuda to northern South America, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from southwestern Florida, the Flower Garden Banks, and Yucatán. This species is associated with coral reefs and rocky substrates. Maximum known size is 95 mm TL. references: Böhlke and Chaplin 1968; Bright and Cashman 1974; Randall 1986a, 1996, 2002b; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith 1997.

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487

POMACENTRIDAE

Damselfishes

Damselfishes are ovate to oblong in profile and compressed, with a single dorsal fin, generally unpaired nostrils, and a highly protrusible upper jaw. Mouth is small, terminal, and oblique. Jaw teeth are conical, incisor-like, or cardiform, and in one or two series. Teeth are absent on vomer and palatine. Lower pharyngeal bones are fused into a plate bearing teeth. Gill opening is unrestricted, but membranes are broadly united across isthmus. Pseudobranch is well developed. Pectoral fin is broad, with base on lower half of flank and nearly vertically oriented. Dorsal fin has 7 to 17 spines and 9 to 21 rays, with spinous section longer than rayed section. Pelvic fin inserts below pectoral fin base and has 1 spine and 5 rays. Anal fin is similar to rayed section of dorsal fin and has 2 spines and 9 to 16 rays. Caudal fin is shallow to deeply forked. Body and part of head are covered with fairly large ctenoid scales. Lateral line consists of tube-bearing scales that extend to end of dorsal fin and a separate series of pored scales that extend about to midlength of caudal fin. Gas bladder is present and usually has a posterior extension into caudal myomeres. Vertebrae number 26: 11 precaudal and 15 caudal. Most species are brightly colored, especially as juveniles. Maximum known size is 35 cm TL. Damselfishes occur worldwide in tropical to warm temperate seas but are most diverse in the Indo-West Pacific Ocean. They are commonly found on coral reefs and are often the most conspicuous fishes on reefs. They are also found on sand, rubble, and silty bottoms and in sea-grass beds. Most species are territorial and guard home ranges for feeding and reproductive purposes. Eggs are laid on the bottom and are guarded by males. Larvae are pelagic. Food consists of algae and small invertebrates, including zooplankton. There are about 315 species in 27 genera, 20 species in 4 genera in the western Central Atlantic, and 14 species in 4 genera in the Gulf of Mexico. references: Leis and Rennis 1983; Allen 1986, 1991, 2001b; Nelson 1994; Randall 1996; Tominaga et al. 1996; Smith-Vaniz et al. 1999; Carter 2002.

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488 PERCIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Emery and Burgess 1974; Smith-Vaniz and Emery 1980) 1a. Upper jaw teeth conical or incisor-like, not flexible and cardiform; no notch in lachrymal bone bordering dorsal margin of upper jaw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Upper jaw teeth flexible and cardiform; distinct notch in lachrymal bone bordering dorsal margin of upper jaw . . . . . . . . . . . . . . . . . . . Microspathodon chrysurus p. 497 2a. Jaw teeth incisor-like or flattened and in a single series; no procurrent spines on upper and lower margins of caudal fin base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2b. Jaw teeth conical and in band two to three teeth wide in anterior section of jaws; two or three procurrent spines on upper and lower margins of caudal fin base . . . . . . . . . . 10 3a. Preoperculum distinctly serrated; margins of teeth entire or with those in anterior section of jaws slightly emarginate .............................................4 3b. Preoperculum entire; margins of teeth distinctly bilobed .............................................9 4a. Cheek scales in three rows; coloration obliquely countercolored . . . . . . . . . . . . . . . . . . . . . Stegastes partitus p. 501 4b. Cheek scales in four rows; body coloration not obliquely countercolored . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5a. Profile of head from snout to level of midorbit straight; juveniles and some adults yellow, with dark spot on axil of pectoral fin, on dorsal fin, and on caudal peduncle; live specimens with bright yellow line above eye . . . . . . . . . . . . . . . . . . . . . . . . . Stegastes planifrons p. 502 5b. Profile of head from snout to level of midorbit convexly rounded; body dark brown or pale yellow, and if pale yellow, with blue lines above eye . . . . . . . . . . . . . . . . . . . . . 6 6a. Three major opercular scales between preoperculum and spine; adults dark blue or brown dorsally, grading to yellow ventrally; blue lines on snout and forehead; diffuse dark spot low on dorsal fin (not present in large adults); juveniles blue dorsally and yellow ventrally, often with dorsal fin spot ocellated . . . . . . . . . . Stegastes variabilis p. 503 6b. Two major opercular scales between preoperculum and spine; color pattern not as above (6a). . . . . . . . . . . . . . . . 7 7a. Fourth or lowest row of cheek scales reduced and not extending to margin of cheek; posterior suborbital bones with strong serrae or spines; color usually dark blue or brown dorsally and pale yellow ventrally; no distinct vertical bars on body; two prominent blue lines on forehead; large spot on pectoral axil . . . Stegastes leucostictus p. 500 7b. Fourth or lowest row of cheek scales normal and extending to margin; color not as above (7a) . . . . . . . . . . . . . . . . . . 8

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POMACENTRIDAE 489

8a. Anal fin extending, at most, to base of caudal peduncle or slightly beyond . . . . . . . . . . . . . . . Stegastes adustus p. 498 8b. Anal fin extending beyond base of caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . . Stegastes diencaeus p. 499 9a. Anal fin rays 12 or 13 (usually 12) . . . . . . . . . . . . . . . . . . . . . . . . . Abudefduf saxatilis p. 490 9b. Anal fin rays 9 or 10 (usually 10) . . . . . . . . . . . . . . . . . . . . . . . . . . Abudefduf taurus p. 491 10a. Dorsal fin spines 12; caudal fin deeply forked to lunate (caudal concavity 27% to 41% of SL); suborbital scales much higher than wide. . . . . . . . . . . . . . . . . . . . . . . . . . 11 10b. Dorsal fin spines 13; caudal fin slightly to moderately forked (caudal concavity 8% to 22% of SL); suborbital scales nearly circular . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11a. Bony tubercle on maxilla projecting above upper lip; pectoral fin rays 18 to 29; gill rakers on lower limb of first gill arch 22 to 25; body greenish gray or brown, with large dark spot on axil of pectoral fin; dorsal fin with pale margin . . . . . . . . . . . . . . . . . . Chromis multilineata p. 495 11b. No bony tubercle on maxilla projecting above upper lip; pectoral fin rays 16 to 18; gill rakers on lower limb of first gill arch 21 or 22; body blue, with no dark spot on axil of pectoral fin; dorsal fin with dark margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chromis cyanea p. 492 12a. Caudal fin pale (yellow, gray, or dusky); juveniles, and often adults, with bright blue V-shaped line on snout and head terminating below spinous section of dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . Chromis enchrysura p. 493 12b. Caudal fin dark gray, blue, or olive; juveniles and adults without bright blue V-shaped line on snout and head (if bright blue V-shaped line is present, it terminates at posterior margin of orbit) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13a. Adults dark olive gray, with dark pectoral spot; juveniles lime green dorsally and abruptly dull olive ventrally; last dorsal spine 12% to 16% of SL; anal fin rays 11 or 12 (usually 11). . . . . . . . . . . . . . . . . . Chromis insolata p. 494 13b. Adults and juveniles dark to bright blue, countershaded or not countershaded; pectoral spot small or absent; last dorsal fin spine 15% to 19% of SL; anal fin rays 11 to 13 (usually 12) . . . . . . . . . . . . . . . . . . . Chromis scotti p. 496

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490 PERCIFORMES

Abudefduf saxatilis Sergeant major

(Linnaeus, 1758)

Abudefduf saxatilis has a smooth suborbital shelf and five broad black bars on side of body. Lachrymal bone, above mouth, is not notched. Jaw teeth are incisor-like and arranged in a single row, and in large individuals they are notched. Preopercular margin is entire. Gill rakers on first arch are moderately long and slender and number 7 or 8 on upper limb and 17 to 19 on lower limb. Measurements are expressed as percent of SL: head length 29%– 33%, snout length 7%–9%, eye diameter 7%– 10%, upper jaw length 8%–9%, pectoral fin length 32%–36%, body depth 57%– 61%. Pectoral fin has 18 or 19 rays. Dorsal fin has 13 spines and 12 or 13 (usually 13) rays. Anal fin has 12 or 13 rays. Caudal fin is strongly forked. Lateral line scales number 19 to 23 (usually 21). Color is bright yellow dorsally and silvery gray on sides and ventrally, with five dark brown to black bands on sides, each tapering to a point ventrally. Fourth band is located immediately posterior to spinous section of dorsal fin. Small dark spot is located on upper base of pectoral fin. It also has a dark color phase in which body is dark bluish gray and bands are indistinct.

This species occurs in tropical to warm temperate waters of the Atlantic Ocean. In the western Atlantic it occurs in shallow water from Rhode Island and Bermuda to Uruguay, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with coral and rocky reefs to 14 m and is common along jetties and shallow reefs in the Gulf of Mexico. Juveniles are associated with pelagic Sargassum and range far offshore. Food consists of benthic algae, colonial anemones, copepods, larval invertebrates, pelagic tunicates, and small ray-finned fishes. Eggs are attached by adhesive filaments to coral or rock surfaces and are guarded by the males. Maximum known size is 150 mm SL.

references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Emery 1973; Castro-Aguirre and MárquezEspinoza 1981; Robins and Ray 1986; Allen 1991; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; SmithVaniz et al. 1999; Carter 2002.

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POMACENTRIDAE 491

Abudefduf taurus (Müller and Troschel, 1848) Night sergeant

Abudefduf taurus has a smooth suborbital shelf and five very broad dark brown bars on side of body. Lachrymal bone, above mouth, is not notched. Jaw teeth are incisor-like and arranged in a single row, and in large individuals they are conspicuously notched. Preopercular margin is entire. Gill rakers on first arch are moderately long and slender and number 6 to 8 on upper limb and 11 or 12 on lower limb. Measurements are expressed as percent of SL: head length about 35%, snout length about 8%, eye diameter about 10%, upper jaw length about 8%, pectoral fin length about 26%, body depth 50%–55%. Pectoral fin has 18 or 19 rays. Dorsal fin has 12 spines and 11 or 12 rays. Anal fin has 9 or 10 (usually 10) rays. Caudal fin is forked. Lateral line scales number 19 or 20. Color is pale to yellowish brown dorsally and on sides, with five very wide dark brown bands running to upper belly. Light markings

between bands are narrower than band width. Sixth band is occasionally present on caudal peduncle. Large black spot generally occurs on axil of pectoral fin. This species occurs in very shallow water in the western Atlantic from southern Florida to northern South America, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with turbulent wave-swept areas and occasionally low-salinity waters. Food consists of benthic algae and submerged vascular plants. Juveniles feed on copepods. Maximum known size is 250 mm TL.

references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Emery 1973; Hoese and Moore 1977, 1998; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Allen 1991; Boschung 1992; Schaldach et al. 1997; Castro-Aguirre et al. 1999; Carter 2002.

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492 PERCIFORMES

Chromis cyanea Blue chromis

(Poey, 1860)

Chromis cyanea has 2 or 3 procurrent spines projecting from upper and lower margins of caudal peduncle and is uniformly blue in color. Bony tubercle is absent from anterior end of maxilla. Lachrymal bone, above mouth, is not notched. Jaw teeth are conical and arranged in two or three rows. Preopercular margin is entire. Gill rakers on first arch are long and slender and number 7 or 8 on upper limb and 21 or 22 on lower limb. Measurements are expressed as percent of SL: head length 29%– 32%, snout length 7%– 8%, eye diameter 9%– 10%, upper jaw length 10%–12%, pectoral fin length 29%–32%, body depth 39%– 44%. Pectoral fin has 16 to 18 rays. Dorsal fin has 12 spines and 12 rays. Anal fin has 12 rays. Caudal fin is deeply forked and has long filamentous tips. Lateral line scales number 16 to 18. Color in life is dark blue dorsally, with broad black margin on spinous dorsal fin and

along upper and lower margins of caudal fin. Color in preservative is brown dorsally and tan ventrally. This species occurs in shallow water in the western Atlantic from Florida and Bermuda to Venezuela, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs off southern Florida and on the Flower Garden Banks. It is generally associated with coral reefs. Food consists of zooplankton, such as copepods, pelagic eggs, siphonophores, and juvenile fishes. Prey are captured by sight during daylight. Maximum known size is 250 mm TL. references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Emery 1973; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Robins and Ray 1986; Allen 1991; Smith 1997; Smith-Vaniz et al. 1999; Carter 2002.

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POMACENTRIDAE 493

Chromis enchrysura Yellowtail reeffish

Jordan and Gilbert, 1882

Chromis enchrysura has several spines projecting from upper and lower margins of caudal peduncle and a thin raised line running from snout over eye to base of dorsal fin. Lachrymal bone, above mouth, is not notched. Jaw teeth are conical and arranged in two rows. Preopercular margin is entire. Gill rakers on first arch are long and slender and number 17 to 21 on lower limb. Measurements are expressed as percent of SL: head length 29%–33%, snout length 7%–9%, eye diameter 9%–11%, upper jaw length 8%–9%, pectoral fin length 30%– 32%, body depth 49%–54%. Pectoral fin has 18 rays. Dorsal fin has 13 spines and 11 to 13 rays. Anal fin has 11 to 13 (usually 12) rays. Caudal fin is forked. Lateral line scales number 15 to 19 (usually 16 or 17). Scale rows on cheek number three, and accessory scales in upper row number five to eight. Color in life is dark blue dorsally and silvery white ventrally,

with a thin, raised brilliant blue line running from upper jaw, between nares, and over eye to dorsal fin base. Caudal fin is yellow. Color in preservative is brown for adults and juveniles. This species occurs in the western Atlantic from southern Florida and Bermuda; the Bahamas; the eastern, northern, and southern Gulf of Mexico; and the Antilles. It is associated with steep slopes on coral reefs and patch reefs between 20 and 100 m. Food consists of benthic algae and zooplankton, such as copepods, decapod shrimp larvae, and molluskan egg masses. Maximum known size is 105 mm SL. references: Emery 1968, 1973; Hoese and Moore 1977, 1998; Emery and Smith-Vaniz 1982; Robins and Ray 1986; Allen 1991; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Carter 2002.

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494 PERCIFORMES

Chromis insolata Sunshinefish

(Cuvier, 1830)

Chromis insolata has several spines projecting from upper and lower margins of caudal peduncle and is countershaded olive green. Lachrymal bone, above mouth, is not notched. Jaw teeth are conical and arranged in two rows. Preopercular margin is entire. Gill rakers on first arch are long and slender and number 19 to 22 on lower limb. Measurements are expressed as percent of SL: head length 29%– 31%, snout length 6%– 8%, eye diameter 10%–12%, upper jaw length 7%–9%, pectoral fin length 30%–35%, body depth 45%– 52%. Pectoral fin has 16 to 18 rays. Dorsal fin has 13 spines and 12 (rarely 11) rays. Anal fin has 11 (rarely 12) rays. Caudal fin is forked. Lateral line scales number 15 to 17 (usually 16). Preorbital scales number 5 to 10. Cheek has three major rows of scales and a series of six or seven accessory scales to upper row. Pectoral fin is scaled over most of length. Scales extend onto medial half of dorsal and anal fin. Color in life is dull olive green dorsally and

dull cream or gray ventrally, usually with a large dark spot on upper base of pectoral fin. Juveniles are lime yellow dorsally and gray ventrally. Color in preservative is yellow brown dorsally and light gray ventrally. This species occurs in shallow water in the western and Central Atlantic from southern Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, the Lesser Antilles, and St. Helena. In the Gulf of Mexico it is known from southwestern Florida, Stetsons Bank off Texas, and Campeche Bay. It is associated with coral reefs. Food consists of copepods, ostracods, shrimp larvae, pelagic tunicates, and zooplankton. Maximum known size is 125 mm SL. references: Böhlke and Chaplin 1968; Emery 1968, 1973; Smith et al. 1975; Hoese and Moore 1977, 1998; Robins and Ray 1986; Allen 1991; Boschung 1992; Smith 1997; Smith-Vaniz et al. 1999; Carter 2002.

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POMACENTRIDAE 495

Chromis multilineata Yellowedge cromis

(Guichenot, 1853)

Chromis multilineata has a bony tubercle projecting from anterior section of maxilla and several spines projecting from upper and lower margins of caudal peduncle. Lachrymal bone, above mouth, is not notched. Jaw teeth are conical and arranged in two or three rows. Preopercular margin is finely serrated. Gill rakers on first arch are long and slender and number 8 or 9 on upper limb and 22 to 25 on lower limb. Measurements are expressed as percent of SL: head length 27%–32%, snout length 6%– 8%, eye diameter 8%–9%, upper jaw length 9%–10%, pectoral fin length 25%– 30%, body depth 34%– 41%. Pectoral fin has 18 to 20 rays. Dorsal fin has 12 spines and 12 rays. Anal fin has 11 to 13 rays. Caudal fin is deeply forked. Lateral line scales number 19 or 20. Color in life is greenish gray to brown dorsally and silvery gray ventrally, with a black spot on axil of pectoral fin, a broad yellow

margin on dorsal fin, and a yellow margin on upper and lower edges of caudal fin. Color in preservative is yellow brown dorsally and light tan ventrally. This species occurs from near shore to 40 m in the western Atlantic from Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs off southwestern Florida, on the Flower Garden Banks, and in Campeche Bay. It is associated with coral reefs. Food consists of zooplankton, such as copepods. Maximum known size is 115 mm SL. references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Emery 1973; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Robins and Ray 1986; Allen 1991; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Carter 2002.

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496 PERCIFORMES

Chromis scotti Emery, 1968 Purple chromis

Chromis scotti has several spines projecting from upper and lower margins of caudal peduncle and dark blue coloration. Lachrymal bone, above mouth, is not notched. Jaw teeth are conical and arranged in two rows. Preopercular margin is finely serrated. Gill rakers on first arch are long and slender and number 18 to 20 on lower limb. Measurements are expressed as percent of SL: head length 29%– 34%, snout length about 8%, eye diameter 10%–14%, upper jaw length 8%–9%, pectoral fin length 31%–36%, body depth 51%– 56%. Pectoral fin has 17 to 19 (usually 18) rays. Dorsal fin has 13 spines and 11 or 12 (usually 12) rays. Anal fin has 11 or 12 (usually 12) rays. Caudal fin is forked. Lateral line scales number 15 to 18 (usually 17). Preorbital scales number five or six. Cheek has three major rows of scales and a series of three to seven (usually four) scales accessory to upper row. Pectoral fin has scales over most of length. Scales extend onto medial half of dorsal and anal fin. Color in life is dark blue with a slight

magenta tinge, with bright blue spots on anterior part of body. Juveniles are countershaded to a greater degree. Color in preservative is dark brown in adults and light brown in juveniles. This species occurs between 5 and 50 m in the western Atlantic from North Carolina and Bermuda to northeastern Brazil, including the Gulf of Mexico, the Florida Keys, the Bahamas, and Jamaica. In the Gulf of Mexico it occurs off the Florida and Alabama coasts, on the Flower Garden Banks, and in Campeche Bay. It is associated with steep outer reef slopes and patch reefs. Food consists of zooplankton, copepods, and pelagic tunicates. Maximum known size is 70 mm SL.

references: Emery 1968, 1973; Randall 1968a, 1996; Bright and Cashman 1974; Hensley and Smith 1977; Hoese and Moore 1977, 1998; Emery and Smith-Vaniz 1982; Robins and Ray 1986; Allen 1991; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Carter 2002.

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POMACENTRIDAE 497

Microspathodon chrysurus Yellowtail damselfish

(Cuvier, 1830)

Microspathodon chrysurus has a notched and very broad lachrymal bone. Jaw teeth in upper jaw are flexible, brushlike, and arranged in a single row. Teeth in lower jaw are incisor-like, stout, and arranged in a single row to form cutting edge. Operculum has a small, acute spine on upper margin. Gill rakers on first arch are short and moderately slender and number 10 on upper limb and 15 to 20 on lower limb. Measurements are expressed as percent of SL: head length 33%–34%, snout length 12%– 14%, eye diameter 8%–9%, upper jaw length 9%–10%, pectoral fin length 29%–31%, body depth 50%–59%. Pectoral fin has 20 to 22 rays. Dorsal fin has 12 spines and 14 or 15 (usually 15) rays. Anal fin has 12 or 13 rays. Caudal fin is forked. Lateral line scales number 20 to 22. Scales extend over most of dorsal and anal fins except for posterior tip. Cheek scales are in four rows in addition to accessory scales. Opercular scales are arranged in three series, with anterior one incomplete. Color in life is dark yellowish brown, with edges of scales

darker. Dorsal part of head and body and dorsal fin are scattered with iridescent blue spots. Caudal fin is bright yellow. Juveniles are blue to bluish brown, with relatively large metallic blue spots. Color in preservative is dark brown, with tail light colored. This species occurs in very shallow water in the western Atlantic from southern Florida and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with coral reefs. Young feed on zooplankton, and adults feed on benthic algae, sponges, harpacticoid copepods, ostracods, isopods, amphipods, and shrimps. Maximum known size is 160 mm SL.

references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Emery 1973; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Carter 2002.

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498 PERCIFORMES

Stegastes adustus Dusky damselfish

(Troschel, 1865)

Stegastes adustus has a coarsely serrated preopercular margin and a relatively short anal fin. Lachrymal bone, above mouth, is not notched. Jaw teeth are incisor-like and close set to form cutting edge. Gill rakers on first arch are short and moderately slender and number 16 to 18, with 9 or 10 on lower limb. Measurements are expressed as percent of SL: head length 30%– 32%, snout length 8%–10%, eye diameter 8%–9%, upper jaw length 7%– 8%, pectoral fin length 28%–32%, body depth 51%–54%. Pectoral fin has 20 to 22 rays. Dorsal fin has 12 spines and 14 to 17 rays. Anal fin has 13 to 15 rays. Caudal fin is forked. Lateral line scales number 18 to 21. Cheek scales are arranged in four major rows, in addition to accessory scales to upper row. Opercular scales are arranged in three columns, with those in second and third columns about the same size. Scales extend over most of dorsal and anal fins except for posterior tips. Color in life is dark gray to blackish, with vertical black lines on body following scale margins. A small black spot is often present on axil of pectoral fin. Fins are dusky except for pectoral fin, which is pale. Faint blue spots occur on head and anterior section of body. Juveniles are lighter, and nape,

spinous dorsal fin, and adjacent upper back are orange red. A large blue-edged black spot occurs on base of dorsal fin at junction of spinous and rayed sections, and a small blue-edged black spot occurs on upper section of caudal peduncle. Color in preservative is dark brown in adults and light brown in juveniles. This species occurs to a depth of 5 m in the western Atlantic from southern Florida to Venezuela, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. There is only a single record of this species from Bermuda. It is associated with rocky shores that are exposed to wave action. Food consists of benthic algae and detritus and, to a lesser extent, polychaetes and harpacticoid copepods. Maximum known size is 100 mm SL. references: Böhlke and Chaplin 1968 (as Eupomacentrus dorsopunicans and E. sp.); Randall 1968a, 1996 (as Stegastes dorsopunicans); Emery 1973 (as E. fuscus); Greenfield and Woods 1974 (as E. dorsopunicans); Hoese and Moore 1977, 1998 (as Pomacentrus fuscus); Castro-Aguirre and Márquez-Espinoza 1981 (as P. fuscus in part); Robins and Ray 1986 (as P. fuscus); Allen 1991 (as S. dorsopunicans); Smith 1997 (as S. dorsopunicans); Smith-Vaniz et al. 1999; Carter 2002.

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POMACENTRIDAE 499

Stegastes diencaeus Longfin damselfish

(Jordan and Rutter, 1897)

Stegastes diencaeus has a relatively long anal fin and spots on scales that form horizontal stripes. Lachrymal bone, above mouth, is not notched. Jaw teeth are incisor-like and close set to form cutting edge. Preopercular margin is finely serrated. Gill rakers on first arch are short and moderately slender and number 6 to 10 on upper limb and 11 to 13 on lower limb. Measurements are expressed as percent of SL: head length 30%–33%, snout length 8%–9%, eye diameter 9%–10%, upper jaw length 7%– 9%, pectoral fin length 28%–31%, body depth 49%–56%. Pectoral fin has 18 to 21 rays. Dorsal fin has 12 or 13 spines (usually 12) and 14 to 16 (usually 15) rays. Anal fin has 12 to 14 rays. Caudal fin is moderately forked. Lateral line scales number 18 to 20. Cheek scales are arranged in four major rows, in addition to accessory scales to upper row. Opercular scales are arranged in three columns, with those of second and third columns about the same size. Scales extend over most of dorsal fin and anal fin except for posterior tips. Color in life is

dark brown to black, with leading edge of anal fin dark iridescent purple. Juveniles are yellow, with a large black spot on upper back and base of posterior section of spinous dorsal fin, and blue lines on series of spots on head and upper part of body. Color of adults in preservative is dark brown to black, with dark spots on scales forming horizontal stripes. This species occurs in the western Atlantic from southern Florida and the Bahamas to Venezuela, including the Gulf of Mexico and the Greater and Lesser Antilles. It is known from East Flower Garden Bank in the Gulf of Mexico. This species is associated with coral reefs in 2 to 45 m, generally in quiet water. Maximum known size is 125 mm TL. references: Böhlke and Chaplin 1968 (as Eupomacentrus sp.); Greenfield and Woods 1974 (as E. diencaeus); Robertson and Allen 1981; Robins and Ray 1986; Randall 1996; Smith 1997; Carter 2002.

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500 PERCIFORMES

Stegastes leucostictus Beau gregory

(Müller and Troschel, 1848)

Stegastes leucostictus has a serrated preopercular margin, a relatively long anal fin, and dark coloration on top of head and nape. Lachrymal bone, above mouth, is not notched. Jaw teeth are incisor-like and close set to form cutting edge. Suborbital shelf is slightly serrated. Preopercular margin is coarsely serrated. Operculum has 2 spines on upper margin, with lower one at angle rather strong. Gill rakers on first arch are short and moderately slender and number 13 to 15, with 7 to 9 on lower limb. Measurements are expressed as percent of SL: head length 26%–30%, snout length 7%– 8%, eye diameter 6%– 8%, upper jaw length 6%– 7%, pectoral fin length 21%–24%, body depth 42%– 48%. Pectoral fin has 17 to 19 (usually 18) rays. Dorsal fin has 12 spines and 13 to 16 rays. Anal fin has 12 to 14 rays. Caudal fin is forked. Lateral line scales number 18 to 20. Cheek scales occur in four major rows, in addition to accessory scales to upper row. Opercular scales occur in three major columns, with those of third column largest. Scales extend over most of dorsal fin and anal fin except for posterior tips. Color in life is blue on head and most of dorsal fin and upper back, and bright yellow on remainder of body. A black spot is located on blue section of dorsal fin. Rows of blue spots occur on top of head, dorsal fin, and

upper back. White spots occur on lower body and anal fin. Large specimens are uniformly dusky. Spot on dorsal fin is best developed in juveniles and disappears with growth. Color in preservative is dark brown except for rayed sections of dorsal and anal fins and caudal fin, which are light. Light spots occur on body and on dorsal and anal fins. Juveniles are lighter. This species occurs to a depth of 5 m in the tropical to warm temperate Atlantic Ocean. In the western Atlantic it occurs from Maine and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. Food consists of benthic algae, harpacticoid copepods, nemerteans, and polychaetes. It is associated with sea-grass beds, coral, and rocky reefs and sandy areas. Maximum known size is 77.5 mm SL.

references: Böhlke and Chaplin 1968 (as Eupomacentrus leucostictus); Randall 1968a, 1996; Emery 1973 (as E. leucostictus); Emery and Burgess 1974 (as E. leucostictus); Castro-Aguirre and Márquez-Espinoza 1981 (as Pomacentrus leucostictus); Robins and Ray 1986 (as P. leucostictus); Allen 1991; Boschung 1992; Schaldach et al. 1997 (as P. leucostictus); Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Carter 2002.

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POMACENTRIDAE 501

Stegastes partitus Bicolor damselfish

(Poey, 1868)

Stegastes partitus has a serrated preopercular margin, a relatively long anal fin, and a bicolored body. Lachrymal bone, above mouth, is not notched. Jaw teeth are incisor-like and close set to form cutting edge. Suborbital shelf is entire. Preopercular margin is coarsely serrated. Operculum has 2 spines on upper margin, with lower one at angle rather strong. Gill rakers on first arch are short and moderately slender and number 17 to 20. Measurements are expressed as percent of SL: head length 27%–33%, snout length 6%– 8%, eye diameter 8%–9%, upper jaw length 6%– 8%, pectoral fin length 26%–30%, body depth 44%– 47%. Pectoral fin has 18 to 20 rays. Dorsal fin has 12 spines and 14 to 17 rays. Anal fin has 13 to 15 rays. Caudal fin is deeply forked. Lateral line scales number 18 to 21. Scales extend over most of dorsal fin and anal fin except for posterior tips. Cheek scales are in three rows, in addition to accessory scales to upper row. Opercular scales are arranged in three columns, with those in second and third columns about the same size. Color in life is dark brown on head and anterior two-fifths of body, and abruptly orange shading to white on remain-

der of body. Demarcation between anterior and posterior colors is diagonal. Deepwater specimens are black anteriorly and gray posteriorly, and the demarcation between colors is also diagonal but less sharp than in shallow-water pattern. Color in preservative is dark brown anteriorly and tan posteriorly. This species occurs to a depth of 45 m in the western Atlantic from North Carolina and Bermuda to Venezuela, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with shallow coral reefs and patch reefs in deeper water. Food consists of benthic algae and several species of pelagic copepods. Maximum known size is 70 mm SL.

references: Böhlke and Chaplin 1968 (as Eupomacentrus partitus); Randall 1968a, 1996; Emery 1973 (as E. partitus); Bright and Cashman 1974 (as E. partitus); Hoese and Moore 1977, 1998 (as Pomacentrus partitus); Robins and Ray 1986 (as P. partitus); Allen 1991; Boschung 1992; Schaldach et al. 1997 (as P. partitus); Smith 1997; SmithVaniz et al. 1999; Carter 2002.

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502 PERCIFORMES

Stegastes planifrons Threespot damselfish

(Cuvier, 1830)

Stegastes planifrons has a serrated preopercular margin, a relatively long anal fin, and a broad saddle-shaped blotch on top of caudal peduncle. Lachrymal bone, above mouth, is not notched. Jaw teeth are incisor-like and close set to form cutting edge. Suborbital shelf is entire. Preopercular margin is coarsely serrated. Operculum has 2 spines on upper margin, with lower one at angle rather strong. Gill rakers on first arch are short and moderately slender and number 17 to 20. Measurements are expressed as percent of SL: head length 31%–34%, snout length 8%–10%, eye diameter 8%–9%, upper jaw length 7%– 8%, pectoral fin length 28%–31%, body depth 48%– 59%. Pectoral fin has 18 to 20 rays. Dorsal fin has 12 spines and 15 to 17 rays. Anal fin has 13 or 14 rays. Caudal fin is forked. Lateral line scales number 18 to 20. Scales extend over most of dorsal fin and anal fin except for posterior tips. Cheek scales are in four rows, in addition to accessory scales to upper row. Opercular scales are arranged in three columns, with those in second and third columns about the same size. Color in life is brownish gray, with a yellowish cast and vertical dark lines along scale margins. A large dark spot is located on pectoral fin base, and a large saddle-shaped

blotch on upper surface of caudal peduncle is lost with growth. Juveniles are bright yellow, with a large dark spot on base of dorsal fin between spinous and rayed sections, a small spot on upper part of pectoral fin base, and a dark saddle-shaped blotch on caudal peduncle. Color in preservative is dark brown, with dark vertical lines, spot on pectoral fin base, and saddle-shaped blotch evident. This species occurs to a depth of 30 m in the western Atlantic from North Carolina and Bermuda to Venezuela, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with coral reefs. Food consists of benthic algae, hydroids, sponges, mollusks, polychaetes, and harpacticoid copepods. Maximum known size is 90 mm SL.

references: Böhlke and Chaplin 1968 (as Eupomacentrus planifrons); Randall 1968a, 1996; Emery 1973 (as E. planifrons); Bright and Cashman 1974 (as Pomacentrus planifrons); Hoese and Moore 1977, 1998 (as P. planifrons); Robins and Ray 1986 (as P. planifrons); Allen 1991; Boschung 1992; Smith 1997; Smith-Vaniz et al. 1999; Carter 2002.

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POMACENTRIDAE 503

Stegastes variabilis Cocoa damselfish

(Castelnau, 1855)

Stegastes variabilis has a serrated preopercular margin, a relatively long anal fin, and a narrow saddle-shaped blotch on top of caudal peduncle. Lachrymal bone, above mouth, is not notched. Jaw teeth are incisor-like and close set to form cutting edge. Suborbital shelf is attached to cheek and is entire. Preopercular margin is coarsely serrated. Operculum has 2 spines on upper margin, with lower one at angle rather strong. Gill rakers on first arch are short and moderately slender and number 15 or 16, with 8 or 9 on lower limb. Measurements are expressed as percent of SL: head length 31%–33%, snout length 7%–9%, eye diameter 7%–9%, upper jaw length 8%–9%, pectoral fin length 25%–30%, body depth 45%–50%. Pectoral fin has 18 to 21 (usually 19 or 20) rays. Dorsal fin has 12 spines and 14 to 17 rays. Anal fin has 12 to 15 rays. Caudal fin is forked. Lateral line scales number 18 to 20. Scales extend over most of dorsal fin and anal fin except for posterior tips. Cheek scales are arranged in four rows, in addition to accessory scales to upper row. Opercular scales are arranged in three columns, with all scales about the same size. Color in life is dark brown to bluish on upper half of body and bright yellow on lower half, with dark vertical lines

along scale margins and a small saddle-shaped blotch on upper margin of caudal peduncle. Pectoral fins are yellow, with a small blackish spot on upper base. Juveniles are blue dorsally and yellow ventrally, with a large black spot on rayed section of dorsal fin and extending onto back. Color in preservative is dark brown, with dark vertical lines and a saddle-shaped blotch usually evident. This species occurs to a depth of 30 m in the western Atlantic from North Carolina and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with coral reefs. Food consists of benthic algae, sponges, hydroids, anemones, and ascidians. Maximum known size is 88 mm SL.

references: Böhlke and Chaplin 1968 (as Eupomacentrus variabilis); Randall 1968a, 1996; Emery 1973 (as E. variabilis); Bright and Cashman 1974 (as E. variabilis); Greenfield and Woods 1974 (as E. variabilis); Hoese and Moore 1977, 1998 (as Pomacentrus variabilis); Robins and Ray 1986 (as P. variabilis); Allen 1991; Boschung 1992; Schaldach et al. 1997; Smith Vaniz et al. 1999; Carter 2002.

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504

LABRIDAE

Wrasses

Wrasses are oblong to elongate and slightly to strongly compressed, with a protrusible upper jaw and a single lower pharyngeal plate bearing conical, nodular, or molariform teeth. Nostril is paired, and anterior and posterior nares are generally close set. Mouth is terminal and small to moderate in size, usually with prominent and often fleshy lips. Posterior end of maxilla is covered by lachrymal bone. Jaw teeth are usually separate and caniniform (rarely incisor-like), generally with one or two pairs of enlarged canine teeth in front of jaws, and rarely with canine tooth at corner of mouth. Teeth are absent in palatine but occasionally present in vomer. Posterior margin of preoperculum is serrated or entire. Opercular margin lacks spine on posterior margin. Slit behind fourth gill arch is restricted or absent. Branchiostegal rays number 5 or 6. Gill membranes are united anteriorly and are either joined or free of isthmus. Pseudobranch is well developed. Pectoral fin has oblique base and varies from broad and paddlelike to long and falcate. Dorsal fin is continuous except in several species and consists of 8 to 21 spines and 6 to 21 rays. Pelvic fin inserts under pectoral fin base and has 1 spine and 5 rays. Anal fin consists of 2 to 6 spines and 7 to 14 rays. Caudal fin is moderately convex, truncate, or double convex. Body and often cheeks and operculum are covered with large cycloid scales. Lateral line is continuous or interrupted. Vertebrae number 23 to 41. Most species are brightly colored, and color patterns often vary with age and sex. Maximum known size is 70 cm TL. Wrasses occur worldwide in tropical to warm temperate seas between the shore zone and 100 m. Many species are associated with coral reefs, but some occur in Thalassia beds, sandy patch reefs, sandy bottoms, and rocky flats. They are active during daylight hours, and most species feed on benthic invertebrates, including gastropods and bivalves, and ray-finned fishes, although some feed on zooplankton. Some species are territorial on reefs, and some small species clean larger fishes of ectoparasites. Most species are hermaphrodites and function as females before changing to males. Color patterns within species may vary with sex. There are about 500 species in at least 60 genera, 20 species in 8 genera occur in the western Central Atlantic, and 17 to 19 species in 8 genera occur in the Gulf of Mexico. references: Fritzsche and Fuiman 1982; Randall 1986a; Nelson 1994; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2001, 2002a.

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LABRIDAE 505

Key to the Species of the Gulf of Mexico (Adapted from Randall and Böhlke 1965; Böhlke and Chaplin 1968; Westneat 2002a) 1a. Dorsal fin spines more than 10. . . . . . . . . . . . . . . . . . . . . 2 1b. Dorsal fin spines less than 10 . . . . . . . . . . . . . . . . . . . . . . 6 2a. Dorsal fin spines 14, with first three of four dorsal fin spines terminating as filaments . . . Lachnolaimus maximus p. 522 2b. Dorsal fin spines 11 or 12, with none terminating as filaments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. Dorsal fin spines 11 . . . . . . . . . . Decodon puellaris p. 511 3b. Dorsal fin spines 12 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4a. Snout pointed; front of jaws with strong canine teeth; posterior canine tooth at corner of jaw; color combination of red or violet and golden or yellow . . . . . . . . . . . . . . . . . . 5 4b. Snout blunt; jaws with relatively small and weak canine teeth; posterior canine teeth absent; color violet . . . . . . . . . . . . . . . . . . . . . . . . . . . Clepticus parrae p. 510 5a. Gill rakers on first gill arch 17 to 19; pectoral fin not darkly pigmented along outer section of upper margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bodianus rufus p. 509 5b. Gill rakers on first gill arch 15 or 16; pectoral fin darkly pigmented along outer section of upper margin . . . . . . . . . . . . . . . . . . . . . . . . Bodianus pulchellus p. 508 6a. Lateral line continuous and steeply curved below posterior of dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6b. Lateral line interrupted posteriorly, with posterior section on midlateral segment of caudal peduncle . . . . . . . . . . . 16 7a. Dorsal fin spines 8; no canine tooth in corner of mouth . . . . . . . . . . . . . . . . . . . . . Thalassoma bifasciatum p. 523 7b. Dorsal fin spines 9; canine tooth in corner of mouth . . . . 8 8a. Two pairs of enlarged canine teeth in anterior section of lower jaw; anal fin rays 12; gill rakers on first arch 15 to 23 .............................................9 8b. One pair of enlarged canine teeth in anterior section of lower jaw; anal fin rays 11; gill rakers on first arch 13 to 15 . . . . . . . . . . . . . . . . . . . . . Halichoeres maculipinna p. 518 9a. Dorsal fin rays 11; broad dark stripe on upper half of body absent; small black spot on nape absent . . . . . . . . . . . . . 10 9b. Dorsal fin rays 12; broad dark stripe on upper half of body extending to caudal fin; small black spot on nape (may be part of dark mark extending from eye or may be faint) . . . . . . . . . . . . . . . . . . . Halichoeres cyanocephalus p. 516 10a. Gill rakers on first arch 15 to 20; body depth 22% to 30% of SL; narrow dark bands radiating from eye absent; small black spot at upper part of pectoral fin base absent; juvenile specimens without five pale bars on upper part of body and no blackish areas on back; no black spot above

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506 PERCIFORMES

midline at base of caudal fin rays; maximum length about 200 mm SL. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10b. Gill rakers on first arch 21 to 23; body depth 28% to 37% of SL; narrow dark bands radiating from eye; small black spot at upper part of pectoral fin base; juvenile specimens with five pale bars on upper part of body and one to four blackish areas on back; small specimens with black spot above midline at base of caudal fin rays; maximum length about 420 mm SL . . . . . . . . . . Halichoeres radiatus p. 521 11a. Anterior lateral line scales with more than one pore; caudal fin slightly rounded (double emarginate in large specimens of H. poeyi). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11b. All lateral line scales with a single pore; large adults with double emarginate caudal fin . . . . . . . . . . . . . . . . . . . . . 14 12a. Body without dark brown or black stripes (juvenile H. garnoti have a single midlateral blue stripe, which is dark edged in preserved specimens) . . . . . . . . . . . . . . . . . . . . 13 12b. Body with two dark brown or blackish stripes or rows of close-set spots, with upper one more prominent and extending from eye to caudal fin and lower one extending from pectoral fin base to caudal peduncle but faint or absent in preserved specimens . . . . . . . . . . . . . . . . . . . . . . . Halichoeres bivittatus p. 514 13a. Black spot behind eye; small black spot on posterior base of dorsal fin; dark line on base of pectoral fin . . . . . . . . . . . . . . . . . . . . . . . . . . Halichoeres poeyi p. 520 13b. Adults with two dark lines extending posteriorly from eye and many scales on nape with single black spots; large adults with dark band on back below rayed section of dorsal fin; juveniles yellow with midlateral blue stripe . . . . . . . . . . . . . . . . . . . . . . . . . Halichoeres garnoti p. 517 14a. No dark spot on head behind eye; snout 33% to 37% of head length; eye not close to dorsal profile . . . . . . . . . . . 15 14b. Dark spot on head behind eye; snout 27% to 33% of head length; eye close to dorsal profile . . . . . . . . . . . . . . . . . . . . . . . . Halichoeres caudalis p. 515 15a. Row of 5 to 7 suborbital pores rimming lower half of eye (exclusive of pores on snout and inclusive of only one of pair of pores perpendicular to eye); 11 to 13 teeth in upper jaw (exclusive of enlarged anterior canine teeth and tooth at rictus); large canine teeth at front of upper jaw not outcurved; one to three dark spots above pectoral fin, beneath fourth lateral line scale . . . Halichoeres bathyphilus p. 513 15b. Row of 8 to 11 suborbital pores rimming lower half of eye (exclusive of pores on snout and inclusive of only one of pair of pores perpendicular to eye); 8 to 10 teeth in upper jaw (exclusive of enlarged canine teeth at front of upper jaw and tooth at rictus); large canine teeth at front of upper jaw outcurved; no dark spot(s) between pectoral fin

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LABRIDAE 507

and lateral line; large adult males with large black spot on caudal peduncle . . . . . . . . . . . . . Halichoeres pictus p. 519 16a. Snout pointed; sides of head below and behind eye covered with scales; posterior canine teeth present . . . . . . . . . . . . . . . . . . . . . . Doratonotus megalepis p. 512 16b. Snout blunt; sides of head below and behind eye mostly naked; posterior canine teeth absent. . . . . . . . . . . . . . . . 17 17a. Five scales between origin of dorsal fin and lateral line; separate posterior section of lateral line with six pored scales; diagonal row of scales below eye extending anteriorly to center of eye; pelvic fin of adults not elongate . . . . . . . . . . . . . . . . . . . . . . . . . Xyrichtys novacula p. 525 17b. Three or four scales between origin of dorsal fin and lateral line; separate posterior section of lateral line with five pored scales; diagonal row of scales below eye not extending to center of eye; pelvic fin of adults elongate. . . . . . . . . . . . 18 18a. Caudal fin rounded; adult male specimens with black spot on side of body; axil of pectoral fin not darker than remainder of body; gill rakers on first arch 17 to 22 . . . . . . . . . . . . . . . . . . . . . . . . . Xyrichtys splendens p. 526 18b. Caudal fin truncate or slightly rounded; no black spot on side of body; axil of pectoral fin dusky to dark brown; gill rakers on first gill arch 21 to 25 . . . . . . . . . . . . . . . . . . . . . . Xyrichtys martinicensis p. 524

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508 PERCIFORMES

Bodianus pulchellus Spotfin hogfish

(Poey, 1860)

Bodianus pulchellus is moderately deep, with a pointed snout, canine teeth in corner of jaws, and a dark blotch of pigment on upper margin of pectoral fin. Dorsal profile of head is nearly straight. Maxilla extends to anterior margin of eye. Upper jaw has two large, slightly projecting canine teeth on either side of symphysis; a small, curved canine in corner of mouth; and a row of stout, blunt teeth behind canines and forming cutting edge. Lower jaw has two large canine teeth on either side of symphysis and a row of short, stout teeth behind canines and forming cutting edge. Preopercular margin is slightly serrated. Gill rakers on first gill arch are short and number 15 or 16. Measurements are expressed as percent of SL: head length 32%– 33%, snout length 10%–12%, eye diameter 6%–7%, upper jaw length 9%–12%, pectoral fin length 20%–24%, body depth 29%–37%. Pectoral fin is winglike and has 15 or 16 rays. Dorsal fin has 11 or 12 (usually 12) spines and 9 to 11 rays. Anal fin is similar in size and shape to rayed section of dorsal fin and has 3 spines and 11 to 13 rays. Posterior rays of dorsal and anal fins are elongated into filaments in adults. Caudal fin is truncate to slightly convex, with upper and lower corners extended as filaments in adults. Scales cover bases of dorsal and anal fins. Lateral line is gently curved

and continuous and has 29 to 31 pored scales. Color is red, with broad, diffuse white stripe and large yellow area along upper half of body from posterior section of dorsal fin to upper lobe of caudal fin. Black spot is located on anterior section of dorsal fin, eye is red, outer section of leading edge of pectoral fin is black, and pelvic and anal fins are red. Juveniles are mostly yellow, with spinous section of dorsal fin black, and often with two dark stripes radiating from posterior margin of eye. This species occurs between 10 and 120 m from South Carolina and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is rare along the northern Gulf of Mexico and southeastern United States. Food consists of small crabs and mollusks. Juveniles are cleaners of parasites on large fishes of many species. Maximum known size is 214 mm SL.

references: Feddern 1963; Böhlke and Chaplin 1968; Randall 1968a, 1996; Haburay et al. 1969; Bright and Cashman 1974; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Shipp and Hopkins 1978; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002a.

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LABRIDAE 509

Bodianus rufus Spanish hogfish

(Linnaeus, 1758)

Bodianus rufus is moderately deep, with a pointed snout, canine teeth in corner of jaws, and an immaculate pectoral fin. Dorsal profile of head is nearly straight. Maxilla extends to anterior margin of eye. Upper jaw has two large, slightly projecting canine teeth on either side of symphysis; a small, curved canine in corner of mouth; and a row of stout, blunt teeth behind canines forming cutting edge. Lower jaw has two large canine teeth on either side of symphysis and a row of short, stout teeth behind canines forming cutting edge. Preopercular margin is slightly serrated. Gill rakers on first gill arch are short and number 17 to 19. Measurements are expressed as percent of SL: head length 33%–35%, snout length 10%– 13%, eye diameter 6%–7%, upper jaw length 10%–13%, pectoral fin length 24%–25%, body depth 29%–37%. Pectoral fin is fanlike and has 15 or 16 rays. Dorsal fin has 11 or 12 (usually 12) spines and 9 to 11 rays. Anal fin is similar in size and shape to rayed section of dorsal fin and has 3 spines and 11 to 13 rays. Posterior rays of dorsal and anal fins are elongated into filaments in adults. Caudal fin is truncate to slightly convex, with upper and lower corners extended as filaments in adults. Scales cover bases of dorsal and anal fins. Lat-

eral line is gently curved and continuous and has 29 to 31 pored scales. Color is bluish purple, reddish, or plum on anterior two-thirds of body, and yellow on ventral and posterior one-third of body. Large adults may be bluish black. Jaws are gold, orange, or reddish; eye is red; and black spot is located on anterior section of dorsal fin. Juveniles are similar in color, but entire anterior half of body may be dark. This species occurs to 40 m in the western Atlantic from North Carolina and Bermuda to southern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is common on offshore banks in the Gulf of Mexico. This species is associated with coral reefs. Food consists of mollusks, sea urchins, brittle stars, and small crabs. Juveniles are cleaners of ectoparasites on large fishes of many species. Maximum known size is 500 mm TL.

references: Feddern 1963; Böhlke and Chaplin 1968; Randall 1968a, 1996; Haburay et al. 1969; Bright and Cashman 1974; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002a.

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510 PERCIFORMES

Clepticus parrae Creole wrasse

(Bloch and Schneider, 1801)

Clepticus parrae is moderately slender, with dorsal and ventral profiles equally curved, a blunt snout, and a deeply forked caudal fin. Dorsal profile of head is nearly straight. Maxilla fails to reach anterior margin of eye. Anterior jaw teeth are small and blunt, with two canine teeth on side of symphysis in upper jaw and one canine on side of symphysis in lower jaw. Preopercular margin is serrated. Gill rakers on first gill arch are slender and short and number 26 to 28. Measurements are expressed as percent of SL: head length 29%–31%, snout length 8%–9%, eye diameter 5%–7%, upper jaw length 7%– 8%, pectoral fin length 27%– 31%, body depth 30%–37%. Pectoral fin is extremely winglike and has 17 or 18 rays. Dorsal fin has 12 spines and 10 rays. Anal fin has 3 spines and 12 or 13 rays. In adults, fifth, sixth, and seventh dorsal and anal rays are prolonged. Caudal fin is lunate. Scales cover body and head except for lips, snout, and basal parts of dorsal and anal fins. Lateral line is continuous and has 32 pored scales. Color is violet or purple, with wash of yellow over ventral two-

thirds of body most distinct on anal fin and caudal peduncle. Teeth and bones are pale blue. Juveniles less than 70 mm SL are purplish dorsally and silvery ventrally, with about six short, black mottled bars on side above lateral line. This species occurs in the western Atlantic from southern Florida and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with patch reefs, reef edges, and oil platforms. Juveniles commonly dwell in sponges at night, and adults rest in crevices of coral reefs at night. Feeding takes place in the water column, and food consists of zooplankton, including jellyfishes, copepods, pteropods, and pelagic tunicates. Maximum known size is 300 mm TL.

references: Stark and Davis 1966; Böhlke and Chaplin 1968; Randall 1968a, 1996; Collette and Talbot 1972; Bright and Cashman 1974; Greenfield 1975a; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002a.

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LABRIDAE 511

Decodon puellaris Red hogfish

(Poey, 1860)

Decodon puellaris is elongate and relatively slender, with a pointed snout and strong canine teeth in front of jaws. Anterior profile of head is slightly convex. Upper and lower jaws have two canine teeth near symphysis and a row of short, stout teeth behind canines. Preopercular margin is finely serrated. Gill rakers on first arch are short and number about 14. Measurements are expressed as percent of SL: head length 34%–36%, snout length 9%– 11%, eye diameter 7%– 8%, upper jaw length 9%–13%, pectoral fin length 16%–21%, body depth 25%–29%. Pectoral fin is fan shaped and has 16 rays. Dorsal fin is continuous and has 11 spines and 9 or 10 rays, with membrane behind spines extended as short filaments. Anal fin has 2 spines and 9 or 10 rays. Caudal fin is truncate, with upper and lower fin rays extended as short filaments. Body and most of

head are covered with large scales. Lateral line is continuous and has 27 to 29 pored scales. Color is red dorsally and white ventrally, with yellow spots. Lips are yellow, and yellow stripes run from nares to eye, from eye to operculum, and from eye across cheek. This species occurs between 18 and 275 m in the western Atlantic from southern Florida and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it is common on the Flower Garden Banks. Maximum known size is 150 mm TL. references: Longley and Hildebrand 1941; Gomon 1974; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith-Vaniz and Böhlke 1991; Boschung 1992; Smith 1997; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Westneat 2002a.

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512 PERCIFORMES

Doratonotus megalepis Dwarf wrasse

Günther, 1862

Doratonotus megalepis is relatively deep bodied, with a pointed snout, anterior section of dorsal fin notched, and lateral line interrupted. Anterior profile of head is nearly uniformly elevated to dorsal fin origin. Jaw teeth are arranged in two rows anteriorly and one row posteriorly, with those in outer row decreasing in size posteriorly and those in inner row small and limited to anterior section of jaws. Upper jaw possesses small canine tooth at corner of mouth. Preopercular margin is smooth. Gill rakers on first arch are short and number 15 or 16. Measurement is expressed as percent of SL: body depth 32%– 40%. Pectoral fin has 11 or 12 rays. Dorsal fin has nine spines and 10 rays, with middle three spines relatively short. Anal fin has three spines and 9 rays. Caudal fin is rounded. Body and most of head are covered with large cycloid scales. Scales are lacking from top of head and side of head anterior to eye. Lateral line has 17 pored scales in anterior section and 4 in posterior section on caudal pe-

duncle. Color is pale green, with oblique white band on cheek, small light spots on head and body, and small reddish or yellowish markings on head and body. Occasionally, ventral section of body is yellowish. Juveniles are blotched with brown pigment. This species occurs in the tropical Atlantic Ocean. In the western Atlantic it occurs from the Florida Keys and Bermuda to northern South America, including the eastern Gulf of Mexico off Florida and Alabama, the Bahamas, and the Lesser Antilles. It is associated with shallow turtle-grass beds between the shoreline and 15 m. Food consists of invertebrates and small ray-finned fishes. Maximum known size is 75 mm TL. references: Longley and Hildebrand 1941; Woods and Kanazawa 1951; Böhlke and Chaplin 1968; Hastings 1979; Robins and Ray 1986; Boschung 1992; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002a.

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LABRIDAE 513

Halichoeres bathyphilus Greenband wrasse

(Beebe and Tee-Van, 1932)

Halichoeres bathyphilus is moderately elongate, with a pointed snout, a complete lateral line, and a dark spot between lateral line and pectoral fin. Upper and lower profiles of body are nearly symmetrical. Mouth has a prominent downward-projecting flap extending from lower lip. Upper jaw has a single downwarddirected canine near symphysis, a row of 11 to 13 smaller teeth, and a small canine in corner. Lower jaw has two pairs of forward-directed canines near symphysis and a row of smaller teeth. Preopercular membrane is filamentous. Gill membranes are broadly joined and united with isthmus. Gill rakers on first arch are short and number 17 to 20. Measurements are expressed as percent of SL: head length 30%– 32%, snout length 9%–10%, eye diameter 5%– 6%, upper jaw length 5%–7%, pectoral fin length 15%–16%, body depth 21%–30%. Pectoral fin is fanlike and has 14 rays. Dorsal fin is continuous and has nine spines and 11 rays, with membrane behind spines forming short tabs. Anal fin has three spines and 12 rays. Caudal fin is slightly rounded to truncate in juveniles and double emarginate in adults.

Body and base of caudal fin are covered with scales, but head is naked. Lateral line is strongly arched posteriorly and consists of 27 pored scales. Color in life is light greenish tan dorsally and pale ventrally, with a stripe from snout to end of caudal fin, a turquoise spot above pectoral fin, and a pale blue caudal fin. Stripe is yellowish brown on head and yellowish orange on body, and caudal fin has yellowish and green stripes. Color in preservative is light brown dorsally and paler ventrally, with a spot between lateral line and pectoral fin and occasionally a dark spot on midcaudal base. This species occurs between 27 and 155 m in the western Atlantic from North Carolina and Bermuda to Yucatán, including the Gulf of Mexico and the Bahamas. In the Gulf of Mexico it occurs off Florida and Alabama and in Campeche Bay. Maximum known size is 230 mm TL. references: Randall and Böhlke 1965; Böhlke and Chaplin 1968; Randall 1968a; Robins and Ray 1986; Boschung 1992; Smith-Vaniz et al. 1999; Westneat 2002a.

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514 PERCIFORMES

Halichoeres bivittatus Slippery dick

(Bloch, 1791)

Halichoeres bivittatus is moderately elongate, with a pointed snout, a complete lateral line, and two dark stripes on body. Upper and lower profiles of body are nearly symmetrical. Mouth has a prominent downward-projecting flap extending from lower lip. Upper jaw has a single outward-directed canine near symphysis, a row of smaller teeth, and a small canine in corner. Lower jaw has two pairs of forward-directed canines near symphysis and a row of smaller teeth. Preopercular membrane is filamentous. Gill membranes are broadly joined and united with isthmus. Gill rakers on first arch are short and number 16 to 19. Measurements are expressed as percent of SL: head length 26%– 32%, snout length 7%–11%, eye diameter 4%–5%, upper jaw length 6%–9%, pectoral fin length 15%–19%, body depth 22%–30%. Pectoral fin is fanlike and has 13 rays. Dorsal fin has nine spines and 11 rays, with membrane behind spines forming short tabs. Anal fin has three spines and 12 rays. Caudal fin is slightly rounded. Body and base of caudal fin are covered with scales, but head is largely naked. Lateral line is strongly arched posteriorly and consists of 27 pored scales. Anterior lateral line scales have more than one pore (often four or five). Color is pale green dorsally

and light greenish yellow on sides and ventrally. Two dark stripes are present on flank; upper one runs from snout to caudal fin base, and lower one is less distinct and runs along lower flank. Upper opercular margin has a bicolored spot, and a small black spot is located at posterior base of dorsal fin. Tips of caudal fin are black in adults. This species occurs in shallow water in the western Atlantic from the Carolinas and Bermuda to Brazil, including the eastern, northern, and southern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with shallow reefs, sandy areas around reefs, and sea-grass beds. Food consists of polychaetes, mollusks, sea urchins, brittle stars, crabs, and ray-finned fishes. Maximum known size is 237 mm SL.

references: Springer and Hoese 1958; Randall and Böhlke 1965; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hastings and Bortone 1976; Hastings 1979; Castro-Aguirre and MárquezEspinoza 1981; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; VegaCendejas et al. 1997; Smith-Vaniz et al. 1999; Westneat 2002a.

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LABRIDAE 515

Halichoeres caudalis Painted wrasse

(Poey, 1860)

Halichoeres caudalis is moderately elongate, with a pointed snout, a complete lateral line, and eye close to dorsal profile of head. Upper and lower profiles of body are nearly symmetrical. Mouth has a prominent downwardprojecting flap extending from lower lip. Upper jaw has a single outward-directed canine near symphysis, a row of 9 to 13 smaller teeth, and a small canine in corner. Lower jaw has two pairs of forward-directed canines near symphysis and a row of smaller teeth. Preopercular membrane is filamentous. Gill membranes are broadly joined and united with isthmus. Gill rakers on first arch are short and number 16 to 19. Measurements are expressed as percent of SL: head length 29%–31%, snout length 9%– 11%, eye diameter 4%–5%, upper jaw length 6%– 8%, pectoral fin length 18%–21%, body depth 23%–28%. Pectoral fin is fanlike and has 13 rays. Dorsal fin has nine spines and 11 rays, with membrane behind spines forming short tabs. Anal fin has three spines and 12 rays. Caudal fin is slightly rounded to truncate.

Body and base of caudal fin are covered with scales, but head is largely naked. Lateral line is strongly arched posteriorly and has 27 singlepored scales. Color in life is olive green dorsally and bright blue ventrally, with a yellow stripe on lower flank, and each scale possesses an olive spot surrounded by a blue ring. A small blue spot is located behind eye. Juveniles are tan with two streaks on side, and the area between streaks is pale orange. Color in preservative is light brown, with a small dark spot behind and slightly above level of center of eye. This species occurs between 26 and 73 m in the western Atlantic from North Carolina to Venezuela, including the northern and southern Gulf of Mexico and the Greater and Lesser Antilles. Maximum known size is 176 mm SL. references: Longley and Hildebrand 1941; Randall and Böhlke 1965; Hastings and Bortone 1976; Sonnier et al. 1976; Hastings 1979; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Westneat 2002a.

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516 PERCIFORMES

Halichoeres cyanocephalus Yellowcheek wrasse

(Bloch, 1791)

Halichoeres cyanocephalus is moderately elongate, with a pointed snout, a complete lateral line, and anterior lateral line scales possessing multiple pores. Upper and lower profiles of body are nearly symmetrical. Mouth has a prominent downward-projecting flap extending from lower lip. Upper jaw has a single outward-directed canine near symphysis, a row of smaller teeth, and a small canine in corner. Lower jaw has a row of small teeth, in addition to two pairs of canine teeth in front of jaw and a small canine in corner of mouth. Preopercular membrane is filamentous. Gill membranes are broadly joined and united with isthmus. Gill rakers on first arch are short and number 18 to 21. Measurements are expressed as percent of SL: head length 27%–30%, snout length 9%–10%, eye diameter about 4%, upper jaw length 7%–10%, pectoral fin length 17%–21%, body depth 26%–29%. Pectoral fin has 13 rays. Dorsal fin has nine spines and 12 rays, with membrane behind spines forming short tabs. Anal fin has three spines and 12 rays. Caudal fin is slightly rounded. Body and base of caudal fin are covered with scales, but head is naked. Lateral line is strongly arched posteriorly and has 27 pored scales. Color in

life is yellowish green dorsally, with a broad black stripe, and blue green ventrally. Scales bordering lower margin of black stripe bear single orange spots. Upper section of head is yellowish green, with a dark diagonal mark extending from nape to eye, and lower section of head is blue green. Color in preservative is pale, with a broad dark stripe on upper section of body. Dorsal section of head has a median dark stripe along midline between eyes and a dark mark extending from nape to eye. Dorsal fin of adults has a broad stripe along base. Juveniles are similarly colored, but stripe along upper body is narrower. This species occurs between 27 and 82 m in the western Atlantic from North Carolina to Brazil, including the Florida Keys and the Greater and Lesser Antilles. It has not been reported from the Gulf of Mexico, but the records from the Florida Keys suggest that it may occur there. Maximum known size is about 300 mm SL.

references: Randall and Böhlke 1965; Robins and Ray 1986; Randall 1996; Smith 1997; Westneat 2002a.

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LABRIDAE 517

Halichoeres garnoti Yellowhead wrasse

(Valenciennes, 1839)

Halichoeres garnoti is moderately elongate, with a pointed snout, a complete lateral line, and anterior lateral line scales possessing more than a single pore. Upper and lower profiles of body are nearly symmetrical. Mouth has a prominent downward-projecting flap extending from lower lip. Upper jaw has a single outward-directed canine near symphysis, a row of smaller teeth, and a small canine in corner. Lower jaw has two pairs of forward-directed canines near symphysis and a row of smaller teeth. Preopercular membrane is filamentous. Gill membranes are broadly joined and united with isthmus. Gill rakers on first arch are short and number 15 to 19. Measurements are expressed as percent of SL: head length 29%– 32%, snout length 8%–11%, eye diameter about 5%, upper jaw length 6%–9%, pectoral fin length 17%–20%, body depth 22%–30%. Pectoral fin is fanlike and has 13 rays. Dorsal fin has nine spines and 11 rays, with membrane behind spines forming short tab. Anal fin has three spines and 12 rays. Caudal fin is slightly rounded. Body and base of caudal fin are covered with scales, but head is naked. Lateral line is strongly arched posteriorly and has 27 pored scales. Adults are yellow on anterior upper part

of body, green on posterior upper part of body, and white ventrally. Dark bar separates dorsal color patterns and is continuous with broad black stripe running along base and posterior section of dorsal fin. Two slender and irregular dark lines run diagonally upward from posterior margin of eye. Small juveniles are bright yellow, with blue, dark-edged stripe. Large juveniles are yellowish brown dorsally and white ventrally. This species occurs in shallow water to 50 m in the western Atlantic from the east coast of Florida and Bermuda to Brazil, including the eastern, northern, and southern Gulf of Mexico, the Bahamas, and the Antilles. This species is associated with shallow coral and rocky reefs. Food consists of invertebrates and ray-finned fishes. Maximum known size is 150 mm TL.

references: Longley and Hildebrand 1941; Randall and Böhlke 1965; Böhlke and Chaplin 1968; Randall 1968a, 1996; Bright and Cashman 1974; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002a.

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518 PERCIFORMES

Halichoeres maculipinna Clown wrasse

(Müller and Troschel, 1848)

Halichoeres maculipinna is moderately elongate, with a pointed snout, a complete lateral line, and a single outcurved canine on each side of symphysis of lower jaw. Upper and lower profiles of body are nearly symmetrical. Mouth has a prominent downward-projecting flap extending from lower lip. Upper jaw has a single outward-directed canine near symphysis, a row of smaller teeth, and a small canine in corner. Lower jaw has a row of small teeth, in addition to a single canine in front of jaw, and a small canine in corner of mouth. Preopercular membrane is filamentous. Gill membranes are broadly joined and united with isthmus. Gill rakers on first arch are short and number 13 to 15. Measurements are expressed as percent of SL: head length 30%–33%, snout length 8%– 10%, eye diameter 5%– 6%, upper jaw length 6%–7%, pectoral fin length 20%–23%, body depth 22%–30%. Pectoral fin is fanlike and has 14 rays. Dorsal fin has nine spines and 11 rays, with membrane behind spines forming short tabs. Anal fin has three spines and 11 rays. Caudal fin is slightly rounded. Body and base of caudal fin are covered with scales, but

head is naked. Lateral line is strongly arched posteriorly and has 27 pored scales. Color in life is yellow dorsally and white ventrally, with a broad black stripe separating dorsal and ventral colors. Three transverse red bands occur on head, a dark spot is often present on dorsal fin between fifth and seventh interspinous membranes, and a small dark spot is located on posterior end of dorsal fin base. Large adult males are rosy green, with a prominent black spot on midflank and a large black spot on spinous section of dorsal fin. This species occurs to a depth of 25 m in the western Atlantic from North Carolina and Bermuda to Brazil, including the eastern and southern Gulf of Mexico, the Florida Keys, the Bahamas, and the Antilles. This species is associated with shallow coral and rocky reefs. Food consists of invertebrates and ray-finned fishes. Maximum known size is 120 mm TL. references: Longley and Hildebrand 1941; Randall and Böhlke 1965; Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002a.

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LABRIDAE 519

Halichoeres pictus Rainbow wrasse

(Poey, 1860)

Halichoeres pictus is moderately elongate, with a pointed snout, a complete lateral line, and a large dark spot on caudal peduncle of adult males. Upper and lower profiles of body are nearly symmetrical. Mouth has a prominent downward-projecting flap extending from lower lip. Upper jaw has a single moderately outward-directed canine near symphysis, a row of 8 to 10 smaller teeth, and a small canine in corner. Lower jaw has two forward-directed canines near symphysis and a row of smaller teeth. Preopercular membrane is filamentous. Gill membranes are broadly joined and united with isthmus. Gill rakers on first arch are short and number 17 or 18. Measurements are expressed as percent of SL: head length 27%–31%, snout length 8%–10%, eye diameter 5%– 6%, upper jaw length 6%– 8%, pectoral fin length 18%–20%, body depth 22%–30%. Pectoral fin is fanlike and has 13 rays. Dorsal fin has nine spines and 11 rays, with membrane behind spines forming short tab. Anal fin has three spines and 12 rays. Caudal fin is slightly rounded to truncate in juveniles, but double emarginate in adults. Body and base of caudal fin are covered with scales, but head is naked. Lateral line is strongly

arched posteriorly and has 27 single-pored scales. Color in life is white or yellow, with two yellowish brown stripes: one along upper back and the other along upper side. In juveniles, stripes are generally restricted to anterior part of body. Large males are bluish green dorsally and pale blue ventrally, with blue stripes on upper head and a large triangular orange stripe behind spot on caudal peduncle. This species occurs between 5 and 25 m in the western Atlantic from southern Florida and the Bahamas to northern South America, including the southern Gulf of Mexico, the Florida Keys, and the Greater and Lesser Antilles. It has putatively been captured in the southern Gulf of Mexico at Isla de Lobos; however, the specimen is a juvenile that can only tentatively be identified as this species, thus its presence in the Gulf of Mexico is questionable. This species is associated with coral reefs and feeds in the water column on zooplankton. Maximum known size is 125 mm TL. references: Randall and Böhlke 1965; Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002a.

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520 PERCIFORMES

Halichoeres poeyi Blackear wrasse

(Steindachner, 1867)

Halichoeres poeyi is moderately elongate, with a pointed snout, a complete lateral line, and a black spot behind eye. Upper and lower profiles of body are nearly symmetrical. Mouth has a prominent downward-projecting flap extending from lower lip. Upper jaw has a single outward-directed canine near symphysis, a row of smaller teeth, and a small canine in corner. Lower jaw has two canines near symphysis, a row of small teeth, and a small canine in corner of mouth. Preopercular membrane is filamentous. Gill membranes are broadly joined and united with isthmus. Gill rakers on first arch are short and number 17 to 20. Measurements are expressed as percent of SL: head length 29%–31%, snout length 9%–11%, eye diameter 4%–5%, upper jaw length 6%– 8%, pectoral fin length 16%–19%, body depth 22%–30%. Pectoral fin is fanlike and has 13 rays. Dorsal fin has nine spines and 11 rays, with membrane behind spines forming short tab. Anal fin has three spines and 12 rays. Caudal fin is slightly rounded. Body and base of caudal fin are covered with scales, but head is

naked. Lateral line is strongly arched posteriorly and has 27 pored scales, with anterior scales bearing multiple pores. Color in life is yellowish green, with red-edged black spot behind eye, small black spot at end of dorsal fin base, and dark blue line at pectoral fin base. Large males are dull green, with center of each scale marked by an orangish red spot. Irregularity in size of spots produces rose-colored bars on head and wavy diagonal bands on tail. This species occurs to a depth of 49 m in the western Atlantic from southeastern Florida and the Bahamas to Brazil, including the eastern and southern Gulf of Mexico and the Greater and Lesser Antilles. It is associated with sea-grass beds and, to a lesser extent, coral and rocky reefs. Maximum known size is 163 mm SL. references: Longley and Hildebrand 1941; Randall and Böhlke 1965; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hastings and Bortone 1976; Robins and Ray 1986; Boschung 1992; Smith 1997; Westneat 2002a.

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LABRIDAE 521

Halichoeres radiatus Puddingwife

(Linnaeus, 1758)

Halichoeres radiatus is moderately elongate, with a pointed snout, a complete lateral line, and anterior lateral line scales possessing multiple pores. Upper and lower profiles of body are nearly symmetrical. Mouth has a prominent downward-projecting flap extending from lower lip. Upper jaw has a single outwarddirected canine near symphysis, a row of smaller teeth, and a small canine in corner. Lower jaw has two canines near symphysis, a row of small teeth, and a small canine in corner of mouth. Preopercular membrane is filamentous. Gill membranes are broadly joined and united with isthmus. Gill rakers on first arch are short and number 21 to 23. Measurements are expressed as percent of SL: head length 30%–33%, snout length 7%–11%, eye diameter 5%– 6%, upper jaw length 7%– 8%, pectoral fin length 18%–21%, body depth 28%–37%. Pectoral fin has 13 rays. Dorsal fin has nine spines and 11 rays, with membrane behind spines forming short tabs. Anal fin has three spines and 12 rays. Caudal fin is slightly rounded. Body and base of caudal fin are covered with scales, but head is naked. Lateral line is strongly arched posteriorly and has 27 pored scales. Color in life is yellowish olive dorsally, shading to orangish yellow ventrally, with rows

of blue spots and pale blue bars on body, a small dark spot on upper base of pectoral fin, and blue lines on head. Juveniles are orange dorsally, with four dark saddles below dorsal fin, dark spot on caudal fin base, and two orange stripes: one extending from eye to caudal fin base and the other from pectoral fin base to caudal peduncle. Large males have a greenish tinge and a black-edged blue bar on midflank. This species occurs to a depth of 50 m in the western Atlantic from North Carolina and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It occurs in the northern and southern Gulf of Mexico but is rare along the coast of Florida. Food consists of polychaetes, sea urchins, brittle stars, mollusks, crabs, and ray-finned fishes. Maximum known size is 420 mm SL.

references: Hildebrand et al. 1964; Randall and Böhlke 1965; Böhlke and Chaplin 1968; Randall 1968a, 1996; Bright and Cashman 1974; Hastings and Bortone 1976; Hoese and Moore 1977, 1998; Hastings 1979; Castro-Aguirre and MárquezEspinoza 1981; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; SmithVaniz et al. 1999; Westneat 2002a.

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522 PERCIFORMES

Lachnolaimus maximus Hogfish

(Walbaum, 1792)

Lachnolaimus maximus is very deep bodied, with a long, pointed snout and first several dorsal fin spines extended as filaments. Dorsal profile of head is very steep and straight to slightly concave. Mouth has a longitudinal fold extending from upper lip and covering posterior section of mouth. Anterior jaw teeth are strong, caninelike, and directed anteriorly, and posterior teeth are short, stout, and arranged in a single row. Preopercular margin is membranous. Gill membranes are connected and joined to isthmus. Gill rakers on first arch number 15 to 17. Measurements are expressed as percent of SL: head length 33%–37%, snout length 11%–13%, eye diameter 7%– 8%, upper jaw length 12%–15%, pectoral fin length 20%–23%, body depth 43%–50%. Mature males have a longer snout than females and juveniles and a concave rather than straight dorsal profile of head. Pectoral fin has 15 or 16 rays. Dorsal fin is deeply notched and has 14 spines and 11 rays, with anterior 3 or (rarely) 4 spines very long and extended as filaments. Anal fin is similar to rayed section of dorsal fin and has 3 spines and 10 rays. Caudal fin is emarginate in juveniles and lunate in adults. Body and much of head are covered with cycloid scales. Lateral line is complete and con-

sists of 32 to 34 pored scales. Color is variable but is typically mottled brownish red, with a black spot at end of dorsal fin. Large males are abruptly maroon on upper half of head and nape, and median fins are blackish along bases. Juveniles are gray, reddish brown, or mottled gray and reddish brown, with a black blotch on posterior aspect of dorsal fin. This species occurs in the western Atlantic from North Carolina and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico. This species is associated with deep reefs but is most common on open bottoms where gorgonian soft corals are abundant. Food consists of sea urchins, bivalve mollusks, gastropods, barnacles, pelecypods, crabs, and hermit crabs. Maximum known size is 100 cm TL.

references: Longley and Hildebrand 1941; Gunter 1944; Randall and Böhlke 1965; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hastings et al. 1976; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Hastings 1979; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002a.

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LABRIDAE 523

Thalassoma bifasciatum Bluehead

(Bloch, 1791)

FPO

Thalassoma bifasciatum is moderately elongate, with a moderately pointed snout, a complete lateral line, and a truncate to lunate caudal fin. Upper and lower profiles of body are nearly symmetrical. Mouth has a prominent downward-projecting flap extending from lower lip and covering posterior section of mouth. Jaw teeth are arranged in a single row and increase in size anteriorly, with those near symphysis projecting anteriorly. Corner of mouth lacks canine tooth. Preopercular margin is membranous. Gill membranes are broadly joined and united with isthmus. Gill rakers on first arch are very short and number 17 to 21. Measurements are expressed as percent of SL: head length 29%–33%, snout length 7%– 8%, eye diameter 5%– 6%, upper jaw length 6%–7%, pectoral fin length 21%– 26%, body depth 23%–29%. Pectoral fin has 14 or 15 rays. Dorsal fin has eight spines and 12 or 13 rays, with membrane behind spines elongated into filaments. Anal fin has three spines and 10 or 11 rays. Caudal fin is slightly truncate to slightly rounded in juveniles and lunate in adults. Body and base of caudal fin are covered with scales, but head is naked. Lateral line is strongly arched posteriorly and consists of 26 pored scales. Color in life is yellow dorsally and white ventrally, with a broad,

dusky greenish stripe or series of square greenish blotches on midflank and a dark spot on anterior part of dorsal fin. Large males have a blue head and a green body, and the two colors are separated by two black bands that are in turn separated by a light blue band. This species occurs in the tropical and subtropical Atlantic. In the western Atlantic it occurs at shallow depths from eastern Florida and Bermuda to northern South America, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico and is associated with coral reefs. This species forms large aggregations over shallow reefs and reef flats. Food consists of zooplankton, benthic invertebrates, and ectoparasites of other fishes. Maximum known size is 180 mm TL.

references: Longley and Hildebrand 1941; Feddern 1965; Randall and Böhlke 1965; Böhlke and Chaplin 1968; Randall 1968a, 1996; Bright and Cashman 1974; Hastings and Bortone 1976; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002a.

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524 PERCIFORMES

Xyrichtys martinicensis Rosy razorfish

Valenciennes, 1840

Xyrichtys martinicensis is elongate and very compressed, with all dorsal fin spines slightly flexible and lateral line interrupted posteriorly. Anterior profile of head is moderately to very steep. Upper jaw is little protrusible. Jaw teeth adjacent to symphysis are caninelike and directed anteriorly, and remaining teeth are small, stout, and arranged in a single row. Preopercular margin is membranous. Gill membranes are united and joined to isthmus. Gill rakers on first arch are short and number 21 to 25, with 7 or 8 on upper limb. Measurements are expressed as percent of SL: head length 28%–31%, snout length 12%–18%, eye diameter 4%–5%, upper jaw length 7%–9%, pectoral fin length 17%–23%, pelvic fin length 17%–26%, body depth 26%–31%. Head depth, snout length, and body depth increase with body size. Pectoral fin has 12 rays, with 2 unbranched and the remainder branched. Dorsal fin has nine spines and 12 rays. Pelvic fin of adults is elongated. Anal fin has three spines and 12 or 13 rays. Caudal fin is truncate to slightly convex. Body and base of caudal fin are covered with scales, and scales occur on nape, upper part of operculum, and cheek. Lateral line has 19 to 22 pored scales in anterior section and 5 in posterior section on caudal peduncle. Diagonal row of scales on cheek extends anteriorly to below rear margin of eye or below center of eye. Scales between origin of dorsal fin and lateral line number 4.

Females and juveniles are rosy to pinkish, with a broad brownish to orangish stripe running from behind eye to base of caudal fin, a white patch on upper part of abdomen, and vertical white lines alternating with reddish lines on lower abdomen. Males are pale blue, with yellowish outlines around scales and eight pale blue bars on pale yellow head. Axil of pectoral fin possesses a multicolored (black, blue, and orange) spot, and pelvic fin is yellow. This species occurs from the shoreline to 21 m in the western Atlantic from southern Florida and the Bahamas to northern South America, including the eastern and southern Gulf of Mexico and the Antilles. It is associated with sandy bottoms, coral rubble, and sea-grass beds. Individuals hover over the bottom and dive headfirst into the sand when threatened. Food consists of mollusks and crustaceans. Abdominal body tissues are transparent in mature females, enabling the brightly colored ova to be visible externally. Maximum known size is 150 mm TL.

references: Böhlke and Chaplin 1968; Randall 1965b, 1968a, 1996; Castro-Aguirre and MárquezEspinoza 1981; Robins and Ray 1986; Cervigón 1993b; Boschung 1992 (all previous references except first as Hemipteronotus martinicensis); Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Westneat 2002a.

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LABRIDAE 525

Xyrichtys novacula Pearly razorfish

(Linnaeus, 1758)

Xyrichtys novacula is elongate and very compressed, with anterior two dorsal spines more flexible than remainder of spines and lateral line interrupted posteriorly. Anterior profile of head is nearly vertical. Upper jaw is little protrusible. Two jaw teeth adjacent to symphysis are caninelike and directed anteriorly; remaining teeth are small, stout, and arranged in a single row. Preopercular margin is membranous. Gill membranes are united and joined to isthmus. Gill rakers on first arch are short and number 18 to 21, with 6 or 7 on upper limb. Measurements are expressed as percent of SL: head length 27%–31%, snout length 12%– 17%, eye diameter 4%–5%, upper jaw length 7%– 8%, pectoral fin length 18%–20%, pelvic fin length 13%–15%, body depth 30%–36%. Pectoral fin has 12 rays, with 2 unbranched and remainder branched. Dorsal fin originates over eye and consists of nine spines and 12 rays, with first two spines separated by membranous gap from remainder of spines. Anal fin has three spines and 12 or 13 rays. Caudal fin is rounded. Body and base of caudal fin are covered with scales, and scales also occur on nape, upper part of operculum, and cheek. Lateral line has 19 to 22 scales in anterior section and 6 in posterior section on caudal pe-

duncle. Diagonal row of scales on cheek extends anteriorly to below center of eye. Scales between origin of dorsal fin and lateral line number 5. Color is dull green dorsally, grading to orange on side, with a blue mark on each scale. A red band is located just posterior to pectoral fin. Head has a number of alternating blue and orange vertical lines below eye. This species occurs in the tropical and subtropical Atlantic. In the western Atlantic it occurs between the shoreline and 80 m from North Carolina and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with sand and coral rubble. Individuals hovering over bottom avoid predation by diving headfirst into sandy bottoms. Food consists of mollusks, shrimps, and crabs. Maximum known size is 220 mm SL.

references: Randall 1965b, 1968a, 1996; Böhlke and Chaplin 1968; Hastings and Bortone 1976; Chittenden and Moore 1977; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997 (all previous as Hemipteronotus novacula); Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002a.

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526 PERCIFORMES

Xyrichtys splendens Green razorfish

Castelnau, 1855

Xyrichtys splendens is elongate and very compressed, with anterior two dorsal spines more flexible than remainder of spines and lateral line interrupted posteriorly. Anterior profile of head is rather steep and convex. Upper jaw is little protrusible. Two jaw teeth adjacent to symphysis are caninelike and are directed anteriorly; remaining teeth are small, stout, and arranged in a single row. Preopercular margin is membranous. Gill membranes are united and joined to isthmus. Gill rakers on first arch are short and number 17 to 22, with 6 or 7 on upper limb. Measurements are expressed as percent of SL: head length 30%–33%, snout length 8%–10%, eye diameter 6%–7%, upper jaw length 7%–9%, pectoral fin length 19%– 22%, pelvic fin length 19%–33%, body depth 30%–36%. Pectoral fin has 12 rays, with 2 unbranched and the remainder branched. Dorsal fin has nine spines and 12 rays, with first two spines separated from remainder of spines by slight membranous gap. Pelvic fin is elongated in adults. Anal fin has three spines and 12 or 13 rays. Caudal fin is rounded. Body and base of caudal fin are covered with scales, and scales also occur on nape, upper part of operculum, and cheek. Lateral line has 19 to 22 scales in anterior section and 5 in posterior section

on caudal peduncle. Diagonal row of scales on cheek extends anteriorly to posterior margin of eye. Scales between origin of first dorsal fin and lateral line number 4. Color is pale bluish gray, with pale blue vertical line or spot through each scale. Head has alternating brownish orange and pale blue vertical lines below eye. Large males are green, with a vertical blue line on each scale, a black spot within yellow area on middle of flank, and purplish blue and orange vertical bands on cheek. This species occurs between 5 and 30 m in the western Atlantic from southern Florida and Bermuda to Brazil, including the eastern Gulf of Mexico off the Dry Tortugas, the Bahamas, and the Greater and Lesser Antilles. It is associated with sand and coral rubble and sea-grass beds. Individuals avoid predation by hiding in algae and sea grass or by diving into sandy bottoms. Food consists of mollusks and crustaceans. Maximum known size is 119 mm SL. references: Longley and Hildebrand 1941 (as Hemipteronotus martinicensis and H. ventralis); Randall 1965b, 1968a, 1996; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997; SmithVaniz et al. 1999; Westneat 2002a.

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527

SCARIDAE

Parrotfishes

Parrotfishes are oblong and moderately compressed, with a nonprotrusible upper jaw, jaw teeth usually fused to form beak, and a single lower pharyngeal plate bearing conical, nodular, or molariform teeth. Snout is usually bluntly rounded. Nostril is paired, and anterior and posterior nares are generally close set. Mouth is small to moderate in size and terminal. Posterior end of maxilla is covered by lachrymal bone. Jaw teeth of some species are fused only at base, with individual teeth visible over length, or teeth are partially fused, with individual teeth visible only at margin of tooth plates. Small incisor-like teeth often occur outside dental plate, and recurved canine teeth often occur in corners of jaws. Pharyngeal dentition consists of upper and lower convex plates bearing molariform teeth. Posterior margin of preoperculum is serrated or entire. Opercular margin lacks spine on margin. Gills are arranged on three and one-half arches, and slit behind fourth gill arch is restricted or absent. Gill membranes are connected and joined to isthmus. Branchiostegal rays number 5 or 6. Pseudobranch on inner surface of opercular bones is well developed. Pectoral fin has oblique base, is greatly expanded distally, and has 2 unbranched and 11 to 14 branched rays. Dorsal fin is continuous and consists of nine slender spines and 10 rays. Pelvic fin inserts under pectoral fin base and has one spine and 5 rays. Anal fin consists of three spines and 9 rays. Caudal fin is moderately convex, truncate, or double convex and has 13 principal rays. Body, cheeks, and operculum are covered with large cycloid scales. Lateral line is discontinuous, and lateral line scales number 22 to 24. Vertebrae number 24 to 26. Most species are brightly colored, and color patterns often vary with age and sex. Maximum known size is 120 cm TL. Parrotfishes occur worldwide in tropical to warm temperate seas to depths from 1 to 30 m and rarely to 90 m. Many species are associated with coral reefs, where they are often one of the most abundant groups of fishes. Some species also occur in lagoons, creeks, and sea-grass beds near reefs. Adults are grazers that feed on filamentous algae, bacterial mats, sea grasses, and live coral polyps. Beaklike jaw teeth are used to crop their food, which is then processed with their pharyngeal teeth. Juveniles feed on small invertebrates. Most species are inactive at night and wedge themselves in crevices or secrete a mucus envelope around their bodies. Most species are hermaphrodites and generally function as females before changing to males. However, individuals of some species are primary males and do not change sexes but may change color patterns during growth. There are about 88 species in nine genera, and 14 species in four genera occur in the western Central Atlantic and the Gulf of Mexico.

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528 PERCIFORMES

references: Randall 1963c, 1986b, 1996; Nelson 1994; Smith 1997; Smith-Vaniz et al. 1999; Bellwood 2001; Westneat 2002b.

Key to the Species of the Gulf of Mexico (adults only for Scarus and Sparisoma species) (Adapted from Böhlke and Chaplin 1968; Randall 1965a,b; Smith-Vaniz et al. 1999) 1a. Teeth united at base only; body depth 22% to 33% of SL; snout rather pointed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Teeth coalesced into beaklike plates; body depth 33% to 40% of SL; snout oblique. . . . . . . . . . . . . . . . . . . . . . . . . 3 2a. Body depth 22% to 25% of SL; no membranous flap on anterior naris; maximum SL 100 mm or less; body pale to reddish, often with alternating white to pink and brown to green stripes. . . . . . . . . . . . . . . Cryptotomus roseus p. 531 2b. Body depth 31% to 33% of SL; membranous flap on anterior naris; maximum SL 290 mm or greater; body green . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nicholsina usta p. 538 3a. Single row of cheek scales; median predorsal scales four; upper dental tooth plate included in lower when mouth closed; gill membranes broadly joined to isthmus. . . . . . . 4 3b. Three or four rows of cheek scales; median predorsal scales six or seven; lower dental tooth plate included in upper when mouth closed; gill membranes narrowly joined to isthmus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 4a. Black saddle or black spot above pectoral fin base . . . . . . 5 4b. Black saddle or spot above pectoral fin base lacking or base entirely dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5a. Fleshy tab on rim of anterior naris greatly subdivided and complex (consisting of 12 to 20 cirri); interorbital region smoothly convex; medial three-fourths to four-fifths of pectoral fin darkly pigmented . . . . . . . . . . . . . Sparisoma rubripinne (supermales) p. 543 5b. Fleshy tab on rim of anterior naris little subdivided, simple, or ribbonlike (consisting of 1 to 6 cirri); interorbital region slightly concave; pectoral fin pale colored . . . . . . . . . . . . . . . . . . . . . Sparisoma chrysopterum p. 541 6a. Fleshy tab on anterior naris simple, ribbonlike; usually two to four canine teeth on side of upper dental plate (in specimens 60 cm SL, except canines may be present in S. radians at smaller sizes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6b. Fleshy tab on anterior naris composite, deeply incised; zero or one canine tooth on side of upper dental plate (in specimens 60 cm SL). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7a. Single midventral scale posterior to origin of pelvic fins; gill rakers on first arch 12 to 16; terminal males lack broad black margin on caudal fin . . Sparisoma atomarium p. 539 7b. Two midventral scales posterior to origin of pelvic fins; gill

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SCARIDAE 529

rakers on first arch 10 to 13; terminal males with broad black margin on caudal fin . . . . . Sparisoma radians p. 542 8a. White saddle or spot behind last dorsal fin ray . . . . . . . . . . . . . . . . . . . . . Sparisoma aurofrenatum p. 540 8b. No white saddle or spot behind last dorsal fin ray . . . . . . 9 9a. Posterior margin of opercular flap with round yellow (in life) or white (in preservative) spot above pale margin . . . . . . . . . . . . . . . . . . . . . . . . . . . Sparisoma viride p. 544 9b. Posterior margin of opercular flap without pale spot or pale margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10a. Shoulder region, above pectoral fin, with yellow or orange (in life) or pale (in preservative) blotch surrounding two or more black spots; gill rakers on first arch usually 11 to 16 . . . . . . . . . . . . . . . . . . . . . Sparisoma aurofrenatum p. 540 10b. Shoulder region, above pectoral fin, without pale blotch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11a. Broad white bar on base of caudal fin; three rows of widely spaced white spots on flank; gill rakers on first arch 17 to 21 . . . . . . . . . . . . . . . . . . . . Sparisoma viride p. 544 11b. No white bar on base of caudal fin; no rows of white spots on flank; gill rakers on first arch 12 to 16 . . . . . . . . . . . . . . . . . . . . . . . Sparisoma rubripinne p. 543 12a. Four horizontal rows of scales on cheek (below eye and above lower preopercular margin); females with a single broad white stripe on flank; pectoral fin of terminal males with dark upper margin adjacent pale stripe extending to tip of fin . . . . . . . . . . . . . . . . . . . . . . . Scarus vetula p. 538 12b. Three horizontal rows of scales on cheek (below eye and above lower preopercular margin); females without a single broad white stripe on flank; pectoral fin of terminal males without dark upper margin adjacent pale stripe . . . . . . . 13 13a. Pectoral fin rays 14 (including two short, unbranched rays); median predorsal scales seven or rarely eight . . . . 14 13b. Pectoral fin rays 15 or 16 (including two short, unbranched rays); median predorsal scales six. . . . . . . . . . . . . . . . . . 15 14a. Usually seven cheek scales in series immediately below eye (last scale of series is behind center of eye); primary phase individuals with dark upper and lower caudal fin margins and snout not yellowish; terminal males with upper and lower margins of caudal fin with narrow dark edges and submarginal orange (in life) or pale (in preservative) stripe . . . . . . . . . . . . . . . . . . . . . . . . . Scarus taeniopterus p. 537 14b. Usually six cheek scales in series immediately below eye (last scale of series is behind center of eye); primary phase individuals without dark upper and lower caudal fin margins and with snout yellowish; terminal males with dark green or blue caudal fin patterned with pink, yellow, or orangish linear markings along center . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scarus iseri p. 536

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530 PERCIFORMES

15a. Pectoral fin rays usually 15 (including two short, unbranched rays); snout broadly rounded or with distinct hump in large juvenile males and adult males; small to medium-sized individuals light blue, with basal section of scales pale salmon and upper part of head yellowish; large adults deep blue or greenish blue, with broad dark gray area on cheek and postorbital region of head; dental plates white in adults . . . . . . . . . . . . . . . . Scarus coeruleus p. 534 15b. Pectoral fin rays usually 16 (including two short, unbranched rays); snout without hump in large juvenile males and adult males; small to medium-sized individuals not light blue, with basal section of scales not pale salmon and upper part of head not yellowish . . . . . . . . . . . . . . . 16 16a. Usually one (occasionally two) scale(s) in lowest cheek scale row; green and bronze in life, without well-defined chin-strap marking below mouth. Scarus guacamaia p. 535 16b. Usually two (occasionally one) scale(s) in lowest cheek row; dark violet and light blue in life, with well-defined dark blue chin-strap marking below mouth (light in preserved specimens) . . . . . . . . . . . . Scarus coelestinus p. 533

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SCARIDAE 531

Cryptotomus roseus Bluelip parrotfish

Cope, 1871

Cryptotomus roseus is elongate, with a pointed snout, teeth fused only basally, and anterior naris slightly raised. Dorsal profile of head to level of eyes is straight. Nares are close set and of similar size, with anterior naris lacking membranous flap. Upper and lower jaws have an outer row of long, flattened, and pointed teeth near symphysis that slant obliquely forward and an inner row of small teeth fused into a band, with tips forming irregular cutting edge. Preopercular margin is membranous. Gill rakers on first arch are short and number 10 or 11. Measurements are expressed as percent of SL: head length 31%–35%, snout length 8%– 11%, eye diameter about 7%, upper jaw length 7%–9%, pectoral fin length 19%–21%, body depth 22%–25%. Pectoral fin has 13 rays. Caudal fin is truncate. Body and part of head are covered with scales. A single row of scales

occurs on cheek, and median predorsal scales number four. Color is olivaceous dorsally and light green ventrally, with small pink spots and a salmon stripe along flank and a black spot on upper part of pectoral fin base. Head is iridescent green with two salmon stripes running to eye and to operculum. This species occurs between 8 and 58 m in the western Atlantic from Florida and Bermuda to Brazil, including the eastern Gulf of Mexico along the Florida coast, the Bahamas, and the Antilles. It is associated with weedy or sandy bottoms. Maximum known size is about 120 mm TL. references: Longley and Hildebrand 1941; Schultz 1958a; Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002b.

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532 PERCIFORMES

Nicholsina usta Valenciennes, 1840 Emerald parrotfish

Nicholsina usta has a slightly acute snout, teeth fused only basally, and anterior naris possessing a simple dermal flap. Dorsal profile of head is evenly convex. Nares are close set and of similar size. Upper and lower jaws have an outer row of large, flattened, and pointed teeth near symphysis that slant obliquely forward and an inner row of incisor-like teeth that are fused only at bases and are bluntly pointed. Adult and large juvenile specimens may have canine teeth on sides of upper jaw. Preopercular margin is membranous. Gill membranes have a narrow free fold across isthmus. Gill rakers on first arch are short and number 12 or 13. Measurements are expressed as percent of SL: head length 32%–36%, snout length 10%–17%, eye diameter 6%–7%, upper jaw length 8%–10%, pectoral fin length 19%– 23%, body depth 31%–33%. Pectoral fin has 13 rays. Caudal fin is truncate to slightly convex. Body and part of head are covered with scales. A single row of scales occurs on cheek,

and median predorsal scales number four or five. Color is mottled olive green dorsally. Scales on flank have reddish margins and bluish white centers. Head has two diagonal reddish orange lines on cheek and yellow below mouth. Median fins are reddish, and dorsal fin has black blotch anteriorly. This species occurs in the tropical and warm temperate Atlantic. In the western Atlantic it occurs to a depth of 73 m from New Jersey and Bermuda to Brazil, including the eastern and southern Gulf of Mexico, the Florida Keys, Cuba, and Hispaniola. It is associated with both turtle-grass beds and the open bottom. Maximum known size is 300 mm TL.

references: Longley and Hildebrand 1941 (as Cryptotomus auropunctatus); Schultz 1958a; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Hastings 1979; Robins and Ray 1986; Boschung 1992; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Westneat 2002b.

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SCARIDAE 533

Scarus coelestinus Valenciennes, 1840 Midnight parrotfish

Scarus coelestinus has two scales in third row on cheek and jaw teeth fused to form beaklike plates. Dorsal and ventral profiles are rounded. Snout is moderately steep. Nares are close set, and anterior naris has a short flap on posterior margin. Tooth plate of upper jaw slightly overlaps that of lower jaw. Gill rakers on first arch number 12 or 13. Measurements are expressed as percent of SL: head length 34%–35%, snout length 11%–13%, eye diameter 7%– 8%, upper jaw length 8%–9%, pectoral fin length 22%–24%, body depth 37%– 40%. Pectoral fin has 16 rays. Caudal fin is slightly convex in juveniles and double emarginate in adults, with upper and lower lobes very long in large specimens. Body; much of head; and bases of dorsal, anal, and caudal fins are covered with scales. Cheek has three rows of scales, with two scales in lower row. Predorsal midrow scales number six. Color in life is bright blue, with edges of scales blackish and centers bright blue.

Scaled section of head is blackish. Light blue bar runs below lower jaw and joins blue scaleless area on cheek. Dental plates are green. Juveniles and all size of adults are similarly colored. This species occurs in the western Atlantic from southern Florida and Bermuda to Brazil, including the eastern and southern Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it occurs off southwestern Florida and in Campeche Bay. This species is associated with coral reefs at depths less than 20 m. Food consists of benthic algae and coral polyps. Maximum known size is 760 mm TL.

references: Longley and Hildebrand 1941 (as Pseudoscarus coelestinus); Schultz 1958a; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hastings 1979; Robins and Ray 1986; Boschung 1992; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002b.

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534 PERCIFORMES

Scarus coeruleus Blue parrotfish

(Bloch, 1786)

Scarus coeruleus has jaw teeth fused to form beaklike plates, two scales in third row on cheek, and a distinct hump on forehead of large juveniles and adults. Dorsal and ventral profiles are rounded. Snout is moderately steep in small specimens and very steep in large specimens. Nares are close set, and anterior naris has short flap on posterior margin. Tooth plate of upper jaw slightly overlaps that of lower jaw. Gill rakers on first arch number 31 to 50. Measurements are expressed as percent of SL: head length 33%–37%, snout length 10%–13%, eye diameter 5%– 6%, upper jaw length 6%– 7%, pectoral fin length 21%–25%, body depth 33%–38%. Pectoral fin has 15 rays. Caudal fin is slightly convex in juveniles and double emarginate in adults, with upper and lower lobes very long in large specimens. Body; much of head; and bases of dorsal, anal, and caudal fins are covered with scales. Cheek has three rows

of scales. Predorsal midrow scales number six. Color in life is light to dark blue, with basal part of scales salmon. Terminal males are deep blue or greenish blue, with broad dark gray region on cheek. Small juveniles are striped and have yellow heads to adulthood. This species occurs in the western Atlantic from Maryland and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs off Florida and Veracruz. This species is associated with coral reefs in shallow water. Food consists of benthic algae. Maximum known size is 900 mm TL. references: Longley and Hildebrand 1941 (as S. caeruleus); Schultz 1958a; Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999, Westneat 2002b.

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SCARIDAE 535

Scarus guacamaia Cuvier, 1829 Rainbow parrotfish

Scarus guacamaia has a single scale or occasionally two scales in third row on cheek and jaw teeth fused to form beaklike plates. Dorsal and ventral profiles are rounded. Snout is moderately steep in small specimens and slightly steeper in large specimens. Nares are close set, and anterior naris has a short flap on posterior margin. Tooth plate of upper jaw slightly overlaps that of lower jaw. Gill rakers on first arch number 51 to 64. Measurements are expressed as percent of SL: head length 35%–36%, snout length 10%–14%, eye diameter 5%–7%, upper jaw length 8%–10%, pectoral fin length 20%–24%, body depth 36%– 40%. Snout length and depth increase with body size. Pectoral fin has 16 rays. Caudal fin is truncate to slightly rounded in juveniles and small adults but is double emarginate in large adults. Body; much of head; and bases of dorsal, anal, and caudal fins are covered with scales. Cheek has three rows of scales. Predorsal midrow scales number six. Color in life is greenish dorsally and greenish orange ventrally, with scales greenish in center and orange on margins. Un-

scaled parts of head and chest are dull orange, and scaled parts are orangish brown with short green lines around eye. In terminal males, green coloration is brighter and more restricted to upper posterior section of body. Tooth plates are green. There is little sexual dichromism. This species occurs in the western Atlantic from southern Florida and Bermuda to Argentina, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it occurs off the southwestern coast of Florida, including the Dry Tortugas and Veracruz. It is associated with coral reefs in shallow water, and juveniles occur in mangrove regions. Food consists of benthic algae, sea grasses, and coral polyps. Maximum known size is 120 cm TL.

references: Longley and Hildebrand 1941 (as Pseudoscarus guacamaia); Schultz 1958a; Böhlke and Chaplin 1968; Randall 1968a, 1996; CastroAguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002b.

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536 PERCIFORMES

Scarus iseri Bloch, 1789 Striped parrotfish

Scarus iseri has six scales in series immediately below eye and jaw teeth fused to form beaklike plates. Dorsal and ventral profiles are rounded. Snout is moderately steep in small specimens and slightly steeper in large specimens. Nares are close set, and anterior naris has a short flap on posterior margin. Tooth plate of upper jaw slightly overlaps that of lower jaw. Outer gill rakers on first arch number 40 to 51, and inner gill rakers number 62 to 78. Measurements are expressed as percent of SL: head length 34%– 35%, snout length 11%–12%, eye diameter about 6%, upper jaw length 8%–11%, pectoral fin length 21%–24%, body depth 34%– 38%. Pectoral fin has 13 or 14 (usually 14) rays. Caudal fin is truncate to slightly convex. Body; much of head; and bases of dorsal, anal, and caudal fins are covered with scales. Cheek has three rows of scales, with five to seven in uppermost row and one to five (usually two or three) in lowermost row. Predorsal midrow scales number seven or (rarely) eight. Color in life is whitish, with three dark brown stripes: from nape to caudal peduncle, from eye or margin of operculum to caudal peduncle, and from pectoral fin base to caudal peduncle. Stripes are

indistinct on head, and upper part of snout is yellowish. Large terminal males are blue green to orange, with a green stripe on head below eye and pink below stripe. Median fins are orange with blue green reticulations or spots and blue medial and distal borders. Caudal fin is orange, with blue green upper and lower margins. This species occurs in the western Atlantic from southern Florida and Bermuda to Brazil, including the eastern and southern Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it occurs off the southwestern coast of Florida and Veracruz. It is associated with coral reefs in shallow water. Food consists of benthic algae. Maximum known size is 270 mm TL.

references: Longley and Hildebrand 1941; Schultz 1958a; Böhlke and Chaplin 1968; Randall 1968a, 1996; Sonnier et al. 1976; Colin 1978; Hastings 1979; Robins and Ray 1986; Boschung 1992 (all previous as S. croicensis); Randall and Nelson 1979; Castro-Aguirre and MárquezEspinoza 1981; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002b.

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SCARIDAE 537

Scarus taeniopterus Princess parrotfish

Desmarest, 1831

Scarus taeniopterus has seven scales in series immediately below eye and jaw teeth fused to form beaklike plates. Dorsal and ventral profiles are rounded. Snout is moderately steep in small specimens and slightly steeper in large specimens. Nares are close set, and anterior naris has a short flap on posterior margin. Tooth plate of upper jaw slightly overlaps that of lower jaw. Gill membranes are connected and joined to isthmus. Outer gill rakers on first arch number 40 to 52, and inner gill rakers number 54 to 67. Measurements are expressed as percent of SL: head length 32%–34%, snout length 12%–13%, eye diameter 5%– 6%, upper jaw length 7%– 8%, pectoral fin length 22%–23%, body depth 34%–38%. Pectoral fin has 13 or 14 rays. Caudal fin is truncate to slightly convex. Body; much of head; and bases of dorsal, anal, and caudal fins are covered with scales. Cheek has three rows of scales, usually seven (rarely eight) in uppermost row and usually two or three in lowermost row. Predorsal midrow scales number seven or (rarely) six. Color in life is tan, with dark brown stripes from nape to caudal peduncle, from eye or margin of operculum to caudal peduncle, and from pectoral fin base to caudal peduncle. Stripes are distinct on head, especially anterior

to eye, and pale stripes between upper two dark stripes meet on snout to form V. Upper and lower margins of caudal fin are dark. Large terminal males are blue green to orange, with two blue green stripes running diagonally from anterior flank through upper and lower margins of eye and to anterior margin of snout. Median fins are solid orange, with blue medial and distal borders. Caudal fin is blue, with orange upper and lower margins. This species occurs in the western Atlantic from southern Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. It is known from the northern and southern Gulf of Mexico. This species is associated with coral reefs, and it schools with S. iseri. Food consists of algae and coral polyps. Maximum known size is 330 mm TL.

references: Schultz 1958a (as S. croicensis, in part); Böhlke and Chaplin 1968; Randall 1963a, 1968a, 1996; Bright and Cashman 1974; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Robins and Ray 1986; Castro-Aguirre and MárquezEspinoza 1981; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002b.

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538 PERCIFORMES

Scarus vetula Bloch and Schneider, 1801 Queen parrotfish

Scarus vetula has four rows of cheek scales and jaw teeth fused to form beaklike plates. Dorsal and ventral profiles are rounded. Snout is moderately steep in small specimens and slightly steeper in large specimens. Nares are close set, and anterior naris has a short flap on posterior margin. Tooth plate of upper jaw slightly overlaps that of lower jaw. Outer gill rakers on first arch number 50 to 62, and inner gill rakers number 71 to 84. Measurements are expressed as percent of SL: head length 34%– 37%, snout length 12%–15%, eye diameter 5%–7%, upper jaw length 8%–9%, pectoral fin length 22%–27%, body depth 33%–38%. Pectoral fin has 14 or (rarely) 15 rays. Caudal fin is truncate in juveniles and small adults but double emarginate in large adults. Body; much of head; and bases of dorsal, anal, and caudal fins are covered with scales. Predorsal midrow scales number seven. Color in life is reddish to purplish brown, with a broad white stripe on

lower flank. Terminal males are bluish green, with reddish orange on edges of scales, alternating bands of orange and blue green on snout and chin, and submarginal orange bands on caudal fin. This species occurs in the western Atlantic from southern Florida and Bermuda to Venezuela, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs along the southwestern coast of Florida, the Flower Garden Banks, and Veracruz. It is associated with coral reefs in shallow water. Food consists of benthic algae and sea grasses. Maximum known size is 550 mm TL. references: Schultz 1958a; Böhlke and Chaplin 1968; Randall 1968a, 1996; Bright and Cashman 1974; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Hoese and Moore 1998; SmithVaniz et al. 1999; Westneat 2002b.

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SCARIDAE 539

Sparisoma atomarium (Poey, 1861) Greenblotch parrotfish

Sparisoma atomarium has jaw teeth fused to form beaklike plates, one to several canine teeth in upper jaw, and a single row of cheek scales. Dorsal and ventral profiles are rounded. Snout is moderately steep, and interorbital space is rather flat to slightly convex. Nares are close set, and anterior naris has a fleshy flap on posterior margin that is simple and ribbonlike. Tooth plate of lower jaw slightly overlaps that of upper jaw, and one to four curved canine teeth are located on dental plate of upper jaw in specimens greater than 25 mm SL. Gill rakers on first arch number 12 to 16 in adults. Measurements are expressed as percent of SL: head length 33%–35%, snout length 9%– 11%, eye diameter 9%–10%, pectoral fin length 23%–26%, body depth 33%–37%. Pectoral fin has 13 rays. Membranes behind dorsal fin spines are extended into cirri. Caudal fin is slightly convex. Body; much of head; and bases of dorsal, anal, and caudal fins are covered with scales. Predorsal midrow scales number four. One midventral scale is located posterior to origin of pelvic fins. In life, upper two-thirds of body is salmon, and lower onethird is pale bluish with two irregular salmon stripes. A large, squarish green blotch with two

deep blue spots occurs below lateral line immediately behind operculum. Dorsal fin has a small dark spot between first two spines, and remainder of dorsal and anal fins are mottled yellowish orange. Caudal fin is pink, with irregular yellow markings. In preservative, large males are light yellowish tan and slightly darker dorsally, with a dark blotch behind opercular opening, followed by five or six faint dark spots. Females and smaller males are pale yellowish tan, with a large dark blotch above and slightly behind pectoral fin base. Snout may be patterned with small dark spots or lines. This species occurs between 17 and 77 m in the western Atlantic from the Florida Keys and Bermuda to Honduras, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It occurs in the northern Gulf of Mexico and in the Dry Tortugas and is associated with coral reefs and sea-grass beds. Maximum known size is 81 mm SL. references: Longley and Hildebrand 1941 (as S. radians, in part); Schultz 1958a (as S. radians, in part); Randall 1965a, 1996; Williams and Shipp 1980; Robins and Ray 1986; Dennis and Bright 1988a; Smith-Vaniz et al. 1999; Westneat 2002b.

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540 PERCIFORMES

Sparisoma aurofrenatum (Valenciennes, 1840) Redband parrotfish

Sparisoma aurofrenatum has a white saddle behind last dorsal fin ray or a cluster of two or three inky black markings on shoulder, and its jaw teeth are fused to form beaklike plates. Dorsal and ventral profiles are rounded. Snout is moderately steep, and interorbital space is slightly concave to flat. Nares are close set, and anterior naris has a fleshy flap on posterior margin that is deeply incised and has four to eight cirri in adults. Upper and lower tooth plates have scalloped biting surfaces, and lower tooth plate slightly overlaps that of upper jaw. Curved canine teeth occur on dental plate of upper jaw in specimens greater than 150 mm SL. Gill rakers on first arch number 11 to 16 in adults. Measurements are expressed as percent of SL: head length 35%–36%, snout length 10%–14%, eye diameter 7%– 8%, upper jaw length 9%–10%, pectoral fin length 23%– 27%, body depth 36%– 42%. Relative snout length increases with size. Pectoral fin has 12 rays. Membranes behind dorsal fin spines are extended into filaments. Caudal fin is truncate in juveniles, emarginate in small adults, and lunate in large adults. Body; much of head; and bases of dorsal, anal, and caudal fins are covered with scales. Cheek has a single row of scales. Predorsal midrow scales number

four. Color in life is mottled brown to greenish brown dorsally, with mottled light red spots on upper part of caudal peduncle. Terminal males are green, with a diagonal orange band extending from corner of mouth to upper end of opercular opening, an orange spot above pectoral fin and enclosing one to several black spots, a white spot on caudal peduncle, and black tips on caudal fin. This species occurs in the western Atlantic from southern Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs off Texas, Louisiana, the southwestern coast of Florida, the Dry Tortugas, and Veracruz. Adults are associated with coral reefs, and juveniles are found in sea-grass beds. Food consists of benthic algae and sea grasses. Maximum known size is 280 mm TL.

references: Longley and Hildebrand 1941 (as Scarus festivus); Schultz 1958a; Böhlke and Chaplin 1968; Randall 1968a, 1996; Bright and Cashman 1974; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Hastings 1979; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002b.

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SCARIDAE 541

Sparisoma chrysopterum Redtail parrotfish

(Bloch and Schneider, 1801)

Sparisoma chrysopterum has jaw teeth fused to form beaklike plates, a single row of cheek scales, and a rather simple fleshy flap on rim of anterior naris. Dorsal and ventral profiles are rounded. Snout is moderately steep, and interorbital space is flat. Nares are close set, and anterior naris has a fleshy flap on posterior margin that is simple and ribbonlike in juveniles, but with a flange around margin or subdivided into a maximum of six cirri in adults. Tooth plates are scalloped, tooth plate of lower jaw slightly overlaps that of upper jaw, and canine teeth are absent. Gill rakers on first arch number 15 to 20 in adults. Measurements are expressed as percent of SL: head length 29%– 34%, snout length 8%–14%, eye diameter 6%– 8%, upper jaw length 7%–9%, pectoral fin length 20%–23%, body depth 34%–37%. Relative snout length and body depth increase with size. Pectoral fin has 13 rays. Membranes behind dorsal fin spines are extended as simple filaments. Caudal fin is rounded in juveniles, truncate in large juveniles and small adults, and lunate in large adults. Body; much of head; and bases of dorsal, anal, and caudal fins are covered with scales. Predorsal midrow scales

number four. Color in life is olivaceous dorsally and mottled light reddish on flanks and ventrally, with edges of scales darker than remainder of scales, head covered with small pale spots, and a saddle-shaped dark spot on upper part of pectoral fin base. Terminal males are green, with edges of scales lavender brown, ventral part of head turquoise, and a deep purple spot on upper part of pectoral fin base. This species occurs in the western Atlantic from southern Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs off Alabama, the southwestern coast of Florida, and Veracruz. It is associated with coral reefs and sea-grass beds in shallow water. Juveniles are found in sea-grass beds. Food consists of benthic algae and sea grasses. Maximum known size is 460 mm TL. references: Longley and Hildebrand 1941; Schultz 1958a; Hastings 1979; Matsuura 1983n; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002b.

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542 PERCIFORMES

Sparisoma radians (Valenciennes, 1840) Bucktooth parrotfish

Sparisoma radians has jaw teeth fused to form beaklike plates, one to several canine teeth in upper jaw, and a single row of cheek scales. Dorsal and ventral profiles are rounded. Snout is moderately steep, and interorbital space is rather flat to slightly convex. Nares are close set, and anterior naris has a fleshy flap on posterior margin that is simple and ribbonlike. Tooth plates are scalloped, tooth plate of lower jaw slightly overlaps that of upper jaw, and one to four curved canine teeth are located on dental plate of upper jaw in specimens greater than 25 mm SL. Gill rakers on first arch number 10 to 13 in adults. Measurements are expressed as percent of SL: head length 32%–36%, snout length 9%–13%, eye diameter 6%– 8%, upper jaw length 8%–10%, pectoral fin length 19%– 21%, body depth 33%– 42%. Relative snout length increases with size. Pectoral fin has 13 rays. Membranes behind dorsal fin spines are extended into cirri. Caudal fin is slightly convex. Body; much of head; and bases of dorsal, anal, and caudal fins are covered with scales. Predorsal midrow scales number four. Two midventral scales are located posterior to origin of pelvic fins. Color in life is olivaceous

to yellowish brown dorsally, with many pale spots, a bluish green opercular margin, and a bluish green bar on pectoral fin base. Terminal males are greenish brown, with faint pale dots or a reticular pattern, a diagonal blue and orange band from corner of mouth to behind eye, irregular orangish red markings on operculum, a blackish bar on pectoral fin base, and a blackish posterior margin on caudal fin. This species occurs in the western Atlantic from southern Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it occurs off Alabama, the southwestern coast of Florida, Veracruz, and Campeche. It is associated with sea-grass beds. Maximum known size is 200 mm TL.

references: Longley and Hildebrand 1941; Schultz 1958a; Randall 1965a, 1968a, 1996; Böhlke and Chaplin 1968; Hastings 1979; CastroAguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Vega-Cendejas 1997; Smith-Vaniz et al. 1999; Westneat 2002b.

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SCARIDAE 543

Sparisoma rubripinne (Valenciennes, 1840) Redfin parrotfish

Sparisoma rubripinne has a composite nasal flap and a single row of cheek scales, and jaw teeth are fused to form beaklike plates. Dorsal and ventral profiles are rounded. Snout is moderately steep, and interorbital space is smoothly convex. Nares are close set, and anterior naris has a palmate flap on posterior margin that is subdivided into 12 to 20 cirri. Upper and lower tooth plates have scalloped biting surfaces, and lower tooth plate slightly overlaps that of upper jaw. Specimens greater than 100 mm SL have curved canine teeth on dental plate of upper jaw. Gill rakers on first arch number 12 to 16 in adults. Measurements are expressed as percent of SL: head length 33%–37%, snout length 10%–13%, eye diameter 7%– 8%, upper jaw length 9%–10%, pectoral fin length 23%–26%, body depth 37%– 40%. Relative snout length increases with size. Pectoral fin has 13 rays. Membranes behind dorsal fin spines are extended as cirri. Caudal fin is slightly convex in juveniles, truncate to emarginate in small adults, and deeply emarginate in large adults. Body; much of head; and bases of dorsal, anal, and caudal fins are covered with scales. Predorsal midrow scales

number four. Color in life is light grayish brown dorsally and nearly white ventrally, with edges of scales darker, alternate light and dark bars on chin, and caudal peduncle and caudal fin yellow. Terminal males are dull green to bluish green, with a large black spot on upper half of pectoral fin base and pectoral fin dark olive except for pale margin. This species occurs in the tropical and warm temperate Atlantic Ocean. In the western Atlantic it occurs in shallow water from Massachusetts and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs off the coasts of Alabama and Florida, the Dry Tortugas, and Veracruz. It is associated with coral reefs and sea-grass beds. Food consists of benthic vegetation. Maximum known size is 450 mm TL.

references: Longley and Hildebrand 1941 (as Scarus brachyvarius); Schultz 1958a (as S. axillaris, in part); Böhlke and Chaplin 1968; Randall 1968a, 1996; Hastings 1979; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Westneat 2002b.

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544 PERCIFORMES

Sparisoma viride (Bonnaterre, 1788) Stoplight parrotfish

Sparisoma viride has jaw teeth fused to form beaklike plates and an opercular margin that is either white or dark. Dorsal and ventral profiles are rounded. Snout is moderately steep, and interorbital space is slightly convex. Nares are close set, and anterior naris has a membranous flap on posterior margin that is longer than it is wide and subdivided into four to seven cirri. Upper and lower tooth plates have scalloped biting surfaces, and lower tooth plate slightly overlaps that of upper jaw. Curved canine teeth occur on dental plate of upper jaw in large specimens. Gill rakers on first arch number 17 to 21 in adults. Measurements are expressed as percent of SL: head length 34%– 37%, snout length 11%–17%, eye diameter 6%–9%, upper jaw length 8%–12%, pectoral fin length 24%–28%, body depth 38%– 43%. Relative snout length increases with size. Pectoral fin has 13 rays. Membranes behind dorsal fin spines are extended into filaments. Caudal fin is truncate in juveniles, emarginate in small adults, and lunate in large adults. Body; much of head; and bases of dorsal, anal, and caudal fins are covered with scales. Cheek has a single row of scales. Predorsal midrow scales

number four. Color in life is dark brown dorsally and bright red ventrally, with three rows of white spots on flank and a broad white bar on base of caudal fin. Terminal males are green, with three diagonal yellowish orange bands on head, a yellowish orange margin and a yellow spot on operculum, a large yellow spot on base of caudal fin, and a narrow crescent near margin of caudal fin. This species occurs in shallow water in the western Atlantic from southern Florida and Bermuda to Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is associated with coral reefs, and juveniles are found in sea-grass beds. Food consists of benthic vegetation. Maximum known size is 640 mm TL.

references: Longley and Hildebrand 1941; Schultz 1958a; Böhlke and Chaplin 1968; Randall 1968a, 1996; Bright and Cashman 1974; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Hastings 1979; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; SmithVaniz et al. 1999; Westneat 2002b.

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545

ZOARCIDAE

Eelpouts

Eelpouts are elongate, rounded or ovoid in cross section anteriorly, and compressed posteriorly, with dorsal fin and anal fin continuous with caudal fin. Head is ovoid or depressed, and snout varies from blunt and short to relatively long and flat. Nostril is single and tubular. Mouth is terminal or inferior, and upper jaw is slightly protrusible. Jaw teeth are variable but are generally small, conical, and arranged in one to several rows. Head, including opercular bones, lacks spines. Gill opening varies from porelike to large, and when large, it extends ventrally to ventral margin of pectoral fin base or is continuous with gill opening on other side. Four gill arches are present, with slit behind fourth. Pseudobranch on medial surface of operculum is usually present. Branchiostegal rays usually number 6. Pectoral fin ranges from minute to well developed and fanlike, except it is absent in one species. Dorsal fin is long and low and lacks spines, although first and (usually) second rays are unsegmented, unbranched, and bilaterally divided at base only. Pelvic fin is usually thoracic and consists of two or three rays in most species; however, it is jugular in one species and absent in 9% of the species. Anal fin is long and low and consists of segmented rays. Body is naked or covered with minute, embedded cycloid scales. Head sensory pores are usually small and rounded or (rarely) large and ovoid, and number and location are often species specific. Lateral line is present or absent; when present, it consists of single neuromasts not enclosed in canals. Skin varies from firm in scaled species to gelatinous in unscaled species. In unscaled species, skin is continuous over head (including eyes), body, and fins. Gas bladder is absent. Ovary is single lobed. Maximum known size is about 118 cm TL. Eelpouts are found worldwide from the poles to the equator along the bottom from coastal regions to the abyss. All species are benthic, except for two genera that are benthopelagic or mesopelagic. Benthic species feed on epifaunal and infaunal organisms, such as polychaetes, bivalves, and gastropods. Pelagic species feed on small planktonic crustaceans. There are about 220 species in 46 genera worldwide, 4 species in separate genera in the western Central Atlantic, and 3 species in separate genera in the Gulf of Mexico. references: Anderson 1984, 1990, 1994, 2001; Nelson 1994.

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546 PERCIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Anderson 1994, in part) 1a. Scales absent on body . . . . Exechodontes daidaleus p. 547 1b. Scales present on body . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Pelvic fin consists of a single short, fleshy ray; upper jaw not extending beyond middle of eye; 17 or 18 pectoral fin rays . . . . . . . . . . . . . . . . . . . . . . Lycenchelys bullisi p. 548 2b. Pelvic fin present or absent, and when present, consists of 3 rays; upper jaw extending to middle of eye in females and to posterior margin of eye in males; 16 or 17 pectoral fin rays. . . . . . . . . . . . . . . . . . . . . . Pachycara sulaki p. 549

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ZOARCIDAE 547

Exechodontes daidaleus

DeWitt, 1977

Exechodontes daidaleus is elongate and compressed posteriorly, with pores of lateral line canal absent on head and body, and pelvic fins long but partially covered by skin. Head is small, and snout is very bluntly rounded. Eye is relatively large but does not enter dorsal profile of head. Mouth is small, slightly oblique, and terminal to subterminal, with lower jaw included in upper, and upper jaw extending to below anterior margin of pupil. Grooves behind lips are interrupted at symphyses of jaws, and upper jaw is not protrusible. Teeth in upper jaw are small and in a single row, and teeth in lower jaw are small and in two rows. Vomer has two teeth, and palatine lacks teeth. Gill slit extends ventrally almost to ventral margin of pectoral fin base. Gill rakers on first arch are absent on upper limb and number 9 to 12 on lower limb. Branchiostegal rays number 6. Measurements are expressed as percent of SL: head length about 14%, head width about 7%,

snout length about 4%, eye length about 3%, interorbital width about 4%, upper jaw length about 6%, body depth about 8%. Pectoral fin has a rounded margin and 13 to 15 rays. Dorsal fin originates behind pectoral fin base and has 86 rays, and anal fin has 79 rays. Pelvic fin has 2 or 3 rays. Vertebrae number 91 to 97: 18 to 21 precaudal and 72 to 78 caudal. Color is very pale yellowish brown, with large scattered brown melanophores more concentrated on ventral one-third than on dorsal two-thirds of body. This species occurs between 219 and 1,004 m in the western Atlantic from northeastern Florida and Cuba to the northern Gulf of Mexico. In the Gulf of Mexico it occurs from the Dry Tortugas to Louisiana. Maximum known size is 113 mm SL.

references: DeWitt 1977; Anderson 1994.

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548 PERCIFORMES

Lycenchelys bullisi

Cohen, 1964

Lycenchelys bullisi is moderately elongate and compressed posteriorly, with a short head, prominent head pores, and a single rayed pelvic fin. Dorsal profile of head and snout is moderately blunt. Nostril is located just above upper lip. Mouth is slightly subterminal. Upper lip is fleshy but does not form fleshy flap. Maxilla does not extend beyond middle of eye. Jaw teeth are short, conical, and arranged in a patch at symphysis and in a single row laterally. Vomer has a circular patch of teeth, and palatine has a single irregular row of four teeth. Gill slit extends from above to below pectoral fin base. Measurements are expressed as percent of SL: head length 15%–16%, snout length about 4%, eye length about 4%, preanal-fin distance 36%– 40%, body depth 8%– 9%. Pectoral fin is broad, with a vertical base and 17 or 18 rays. Dorsal fin is confluent with

caudal fin and anal fin and has 100 rays. Pelvic fin is jugular and consists of a single short, fleshy ray. Anal fin has 83 rays. Vertical fins are of uniform height. Head pores consist of seven maxillary pores, two postorbital pores, one occipital pore near upper angle of operculum, one preopercular pore about midway between posterior margin of eye and posterior margin of operculum, and seven mandibular pores. Color is uniformly light brown. This species occurs between 625 and 1,247 m in the western Atlantic from northeastern Florida to the northern Gulf of Mexico. In the Gulf of Mexico it is recorded from west of Key West and from Mississippi and Texas. Maximum known size is 180 mm TL.

references: Cohen 1964; Retzer 1990.

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ZOARCIDAE 549

Pachycara sulaki Anderson, 1989

Pachycara sulaki is robust and elongate, with pelvic fins, scales, and a lateral line. Head is large and ovoid, and snout is bluntly rounded. Eye is circular and in dorsal profile of head. Mouth is subterminal and oblique, with upper jaw extending to middle of eye in females and to posterior margin of eye in males. Teeth are present in jaws, vomer, and palatine. Lips are fleshy. Gill opening extends ventrally to ventral margin of pectoral fin base. Gill rakers on first arch number 2 or 3 on upper limb and 9 to 13 on lower limb. Measurements are expressed as percent of TL: head length 15%–18%, head width 7%–9%, pre-dorsal-fin length 21%– 24%, pre-anal-fin length 39%– 42%, pectoral fin length 10%–11%, body depth 8%–10%; and as percent of head length: snout length 16%–20%, eye length 18%–24%, upper jaw length 36%–50%, interorbital width 6%– 11%. Pectoral fin is fan shaped and has 16 or

17 rays. Dorsal fin has 99 to 107 rays, anal fin has 81 to 89 rays, and caudal fin has 9 to 11 rays. Pelvic fin is present or absent, and when present, consists of 3 rays. Scales are present but are absent on head and on body anterior to pectoral fin. Lateral line has mediolateral and ventral branches, and mediolateral branch originates just behind posteriormost postorbital pore. Vertebrae number 107 to 113: 27 to 30 precaudal and 80 to 85 caudal. Color is dark brown, dark blue, or sandy gray, with unpaired fins black or translucent. This species occurs between 2,000 and 3,510 m in the western Atlantic in the northeastern Gulf of Mexico and the Caribbean Sea. In the Gulf of Mexico it is known from the Florida Escarpment. Maximum known size is 189 mm SL. reference: Anderson 1989.

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550

CHIASMODONTIDAE

Chiasmodontids are elongate and moderately compressed, with a long and slender premaxilla, dorsal fin divided into two fins, and a very distensible stomach. Dorsal surface of head is rugose and pitted by sensory pores. Snout is moderately short to elongate, and acute to rounded. Nostril is paired, and anterior and posterior nares are close set and porelike. Mouth is terminal, large, and nearly horizontal. Premaxilla and maxilla are slender, nonprotractile, and firmly joined distally. Jaw teeth are large and arranged in one or two rows or in three to five bands. Teeth are present in palatine and present or absent in vomer. Gill rakers are absent or replaced by gill teeth fused to bony plates. Branchiostegal rays number 6 or 7. Gill membranes are separate and free of isthmus. Pectoral fin has a nearly vertical base and 9 to 15 rays. First dorsal fin has 5 to 13 flexible spines, and second dorsal fin has 0 to 1 spine and 18 to 29 rays. Pelvic fin is located beneath pectoral fin base and has 1 spine and 5 rays. Anal fin usually resembles second dorsal fin in size and shape and has 0 to 1 spine and 17 to 29 rays. Caudal peduncle is narrow, and caudal fin is forked and has 17 primary rays. Body is naked, or is covered with small projecting spinules, or has two or more rows of stout, projecting prickles. Lateral line is complete and consists of series of pores along side of body. Photophores are present in one genus. Vertebrae number 33 to 48. Right and left pelvic girdles are separate and not connected to pectoral girdle. Maximum known size is 262 mm SL. Chiasmodontids occur worldwide in the tropical and subtropical seas at mesopelagic and bathypelagic depths. Food consists of ray-finned fishes, and prey sometimes approach or exceed the size of the predator. There are about 35 species in four genera, 12 or 13 species in four genera occur in the western Central Atlantic, and 9 or 10 species in four genera occur in the Gulf of Mexico. Several of the genera are in need of revision, and the accounts given herein may not accurately reflect the species in the Gulf of Mexico. references: Johnson and Keene 1986a,b; Nelson 1994; Mooi and Paxton 2001; McEachran and Sutton 2002.

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CHIASMODONTIDAE 551

Key to the Species of the Gulf of Mexico (Adapted from Norman 1929; Johnson and Cohen 1974; Johnson and Keene 1986b; Robert Lavenberg, pers. comm., January 2002) 1a. Head and body with rows of small photophores arranged into definite rows and patterns; premaxilla greatly broadened; snout short and convex . . . . . . . . . . . . . . . . . . . . . . 2 1b. Head and body without photophores; premaxilla not broadened; snout moderately long and pointed . . . . . . . . 4 2a. Premaxilla with three uniserial rows of needlelike teeth; dorsal pectoral fin ray with short row of photophores; mandibular photophores in three groups . . . . . . . . . . . . . . . . . . . Pseudoscopelus obtusifrons p. 561 2b. Premaxilla with one uniserial row and two multiple rows of sharp, conical teeth; dorsal pectoral fin ray without row of photophores (a single light organ may be present in some individuals); mandibular photophores in two groups, anterior group absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. Single row of photophores between pelvic fin insertions . . . . . . . . . . . . . . . . . . . . Pseudoscopelus altipinnis p. 560 3b. Broad band of photophores between pelvic fin insertions . . . . . . . . . . . . . . . . . . . . . Pseudoscopelus scriptus p. 562 4a. Prickles along side of body in adults; jaw teeth small to moderate in size, straight, and arranged in three or four rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4b. Without prickles along side of body in adults; jaw teeth moderate to large, decurved, and arranged in one or two rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5a. Single row of prickles above and below lateral line; vomer with teeth . . . . . . . . . . . . . . . Dysalotus oligoscolus p. 556 5b. More than one row of prickles above and below lateral line; vomer without teeth. . . . . . . Dysalotus alcocki p. 555 6a. Pseudobranch absent or rudimentary; upper and lower jaws markedly arched, mouth not completely closing; teeth in front of jaws smaller than remainder; pectoral fin length much greater than body depth . . . . . . . . . . . . . . . . . . . . . 7 6b. Pseudobranch well developed; upper and lower jaws not markedly arched and meet when mouth closed; teeth in front of jaws largest; pectoral fin length equal to or slightly greater than body depth . . . . . . . . . . . . . . . . . . . . . . . . . . 9 7a. Upper jaw generally with 7 or more teeth in inner row; teeth in inner row relatively short and moderately curved . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kali macrodon p. 558 7b. Upper jaw with 6 or fewer teeth in inner row; teeth in inner rows relatively long and curved . . . . . . . . . . . . . . . . . . . . 8 8a. Upper and lower jaws with 14 to 38 teeth in outer rows; more than 1 small tooth in outer row preceding first tooth

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552 PERCIFORMES

in inner row of each jaw; well-developed teeth in gill arches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kali normani p. 559 8b. Upper and lower jaws with 6 to 12 teeth in outer rows; 1 small tooth in outer row preceding first tooth in inner row of each jaw; no teeth in gill arches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kali indica p. 557 9a. Skin beset with fine spinuloid scales in specimens less than 35 mm SL; single series of teeth in lower jaw; pectoral fin rays 15 . . . . . . . . . . . . . . . . . Chiasmodon subniger p. 554 9b. Skin naked at all stages; two series of teeth in lower jaw; pectoral fin rays 12 to 15 (usually 13) . . . . . . . . . . . . . . . . . . . . . . . . . . Chiasmodon niger p. 553

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CHIASMODONTIDAE 553

Chiasmodon niger

Johnson, 1864

Chiasmodon niger has two rows of jaw teeth, with anteriormost teeth larger than remainder of teeth. Snout is strongly rounded. Lower jaw projects beyond upper jaw. Upper jaw has three canine teeth on side of symphysis: first is fixed and strongly curved; second is moderately long, depressible, and directed posteriorly; and third is long, depressible, and directed posteromedially. Lower jaw has three canine teeth on side of symphysis: first is fixed and curved, second and third are depressible, and third is very long. Palatine teeth are small and in a single series. Preoperculum has a small spine at lower angle. First three gill arches lack gill teeth and tooth plates. Pseudobranch is well developed. Branchiostegal rays number 7. Measurements are expressed as percent of SL: head length 24%–30%, snout length 6%–7%, eye diameter 4%–5%, interorbital width 5%– 6%, body

depth 13%–16%. Pectoral fin is about as long as, or slightly longer than, body depth and has 12 to 15 (usually 13) rays. First dorsal fin has 10 to 12 spines, and second dorsal fin has 1 spine and 26 to 29 rays. Anal fin has 1 spine and 26 to 29 rays. Lateral line consists of two pores per body segment. Vertebrae number 43 to 46. Specimens of all sizes are naked. Color is brownish black. This species occurs worldwide in tropical seas. In the western Atlantic it occurs generally below 750 m from New England and Bermuda to Brazil, including the entire Gulf of Mexico. Maximum known size is 250 mm SL. references: Norman 1929; Parr 1933; Uyeno 1984; Johnson and Keene 1986a,b; McEachran and Sutton 2002.

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554 PERCIFORMES

Chiasmodon subniger

Garman, 1899

Chiasmodon subniger has a single row of jaw teeth in lower jaw, with anteriormost teeth larger than remainder of teeth. Snout is strongly rounded. Lower jaw extends beyond upper jaw. Upper jaw has three canine teeth on side of symphysis: first is moderately long, depressible, and directed posteriorly; second is fixed and strongly curved; and third is long, depressible, and directed posteromedially. Lower jaw has three canine teeth on side of symphysis: first is fixed and curved, and second and third are depressible. Palatine teeth are small and in a single series. Lower margin of preoperculum has a weak spine. First three gill arches lack gill teeth and tooth plates. Pseudobranch is well developed. Branchiostegal rays number 7. Measurements are expressed as percent of SL: head length 28%–29%, snout length 7%– 8%, eye diameter 5%– 6%, interorbital width 5%–

6%, body depth 14%–19%. Pectoral fin is about as long as, to slightly longer than, body depth and has 15 rays. First dorsal fin has 10 to 12 spines, and second dorsal fin has 1 spine and 26 to 29 rays. Anal fin has 1 spine and 26 to 29 rays. Lateral line consists of two pores per segment. Vertebrae number 43 to 46. Specimens up to 35 mm SL are covered with small spinuloid scales, and those larger than 35 mm SL are naked. Color is brownish black. This species occurs in the eastern Pacific and western Atlantic. In the western Atlantic it occurs in the Gulf of Mexico off Louisiana, Mississippi, and Florida. The western Atlantic form may represent a new species. references: Garman 1899; Norman 1929; Parr 1933 (as Chiasmodon niger pluriradiatus); McEachran and Sutton 2002.

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CHIASMODONTIDAE 555

Dysalotus alcocki

MacGilchrist, 1905

Dysalotus alcocki has prickles on sides of body and jaw teeth that are straight and of moderate to small size. Dorsal profile of head is slightly convex. Snout is acute, with lower jaw possessing symphyseal knob and projecting beyond upper jaw. Supramaxilla is present. Jaw teeth are straight, needlelike, depressible, and arranged in three or four rows. Jaw teeth in inner row are largest, and larger teeth in row have lanceolate tips. Vomerine teeth are absent. Palatine teeth are in a single row and number 3 to 14. Gill teeth occur on first three arches, and those on first arch number 4 to 14 on upper limb and 9 to 24 on lower limb. Branchiostegal rays number 7. Pseudobranch is poorly developed. Measurements are expressed as percent of SL: head length 27%–35%, snout length 10%–14%, eye diameter 3%–5%, interorbital width 6%– 8%, upper jaw length 16%–22%, pre-dorsal-fin length 32%– 40%,

pre-anal-fin length 50%–57%, body depth 11%–17%, caudal peduncle depth 3%–5%. Pectoral fin has 11 to 13 rays. First dorsal fin has 10 or 11 spines. Second dorsal fin has 1 spine and 25 to 27 rays. Anal fin has 1 spine and 26 to 28 rays. Lateral line consists of 42 to 45 pored scales. Side of body is sparsely covered with prickles. Vertebrae number 39 or 40. Color is black. This species occurs at mesopelagic depths beneath tropical seas except for the eastern Pacific. In the western Atlantic it is known from Bermuda, the northeastern Gulf of Mexico, and the Antilles. In the Gulf of Mexico it occurs between the surface and 1,000 m. Maximum known size is 223 mm SL. references: Norman 1929; Parr 1933; Johnson and Cohen 1974; Johnson and Keene 1986b; McEachran and Sutton 2002.

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556 PERCIFORMES

Dysalotus oligoscolus

Johnson and Cohen, 1974

Dysalotus oligoscolus has prickles on sides of body and jaw teeth that are straight and of moderate to small size. Dorsal profile of head is slightly convex. Snout is acute, and lower jaw either lacks or has very small symphyseal knob. Supramaxilla is present. Jaw teeth are straight, needlelike, depressible, and arranged in several rows. Jaw teeth in inner row are largest, and larger teeth in row have lanceolate tips. Vomerine teeth are present. Palatine teeth are in a single row and number 5 to 19. Gill teeth occur on first three arches, and those on first arch number 5 to 15 on upper limb and 14 to 27 on lower limb. Branchiostegal rays number 7. Pseudobranch is poorly developed. Measurements are expressed as percent of SL: head length 26%–32%, snout length 10%–13%, eye diameter 3%– 4%, interorbital width 4%– 8%, upper jaw length 18%–22%,

pre-dorsal-fin length 32%– 40%, pre-anal-fin length 51%–57%, body depth 8%–13%, caudal peduncle depth 3%– 4%. Pectoral fin has 11 or 12 rays. First dorsal fin has 10 to 12 spines. Second dorsal fin has 1 spine and 24 to 26 rays. Anal fin has 1 spine and 24 to 27 rays. Lateral line consists of 42 or 43 pored scales. Side of body has a single row of prickles above lateral line. Vertebrae number 38 to 40. Color is black. This species occurs beneath tropical, subtropical, and southern warm temperate seas at mesopelagic depths. It has been captured in the northeastern Gulf of Mexico between 1,000 and 1,500 m. Maximum known size is 227 mm SL. references: Johnson and Cohen 1974; McEachran and Sutton 2002.

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CHIASMODONTIDAE 557

Kali indica Lloyd, 1909

Kali indica has strongly arched jaws, two rows of jaw teeth, and no tooth plates on gill arches. Dorsal profile of head is evenly convex and has large cavities. Snout is strongly rounded. Middle sections of jaws are separated when mouth is closed, and lower jaw extends beyond upper jaw. Supramaxilla is absent. Jaw teeth are needlelike, moderately to strongly recurved, and usually depressible, with those of inner row very large, recurved, barbed, and considerably larger than those in outer row. Upper jaw has 6 to 12 teeth in outer row and 3 or 4 in inner row. Lower jaw has 6 to 10 teeth in outer row and 3 or 4 in inner row. Palatine teeth are long, needlelike, and arranged in a single row of 3 to 7 teeth. Vomerine teeth are absent. Branchiostegal rays number 6. Measurements are expressed as percent of SL: head length 23%–28%, snout length 7%–10%, eye diameter 3%– 6%, interorbital length 5%–7%, upper jaw length 18%–23%, pre-dorsal-fin length 26%–31%, pre-anal-fin

length 50%–58%, pectoral fin length 13%– 22%, body depth 12%–18%, caudal peduncle depth about 4%. Pectoral fin has 12 or (rarely) 13 rays. First dorsal fin has 10 to 12 spines, and second dorsal fin has 1 spine and 22 to 24 rays. Anal fin has 1 spine and 21 to 25 rays. Lateral line consists of about 40 pairs of pores, with five or more palps between each pair. Vertebrae number 37 to 41. Color is black. This species occurs in tropical and subtropical waters of the Atlantic, Pacific, and Indian Oceans, based on capture of relatively few specimens. In the western Atlantic it occurs off Bermuda and in the Yucatán Channel. Although it has not been reported from the Gulf of Mexico, it may occur there. It is usually captured at depths greater than 1,500 m. Maximum known size is 262 mm SL. references: Johnson 1969; Johnson and Cohen 1974; Johnson and Keene 1986a,b; McEachran and Sutton 2002.

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558 PERCIFORMES

Kali macrodon

(Norman, 1929)

Kali macrodon has strongly arched jaws, two rows of jaw teeth, and tooth plates on first three gill arches. Dorsal profile of head is slightly convex and has large cavities. Snout is very acute. Supramaxilla is absent. Middle sections of jaws are widely separated when mouth is closed. Lower jaw extends beyond upper jaw. Jaw teeth are needlelike, moderately recurved, and usually depressible, with those of inner row barbed and considerably larger and more curved than those in outer row. Upper jaw has 20 to 24 teeth in outer row and 6 to 9 in inner row. Lower jaw has 18 to 31 teeth in outer row and 7 to 10 in inner row. Palatine teeth are long and needlelike, are arranged in a single row, and number 5 to 12. Vomerine teeth are absent. First gill arch has 15 gill teeth. Branchiostegal rays number 6. Measurements are expressed as percent of SL: head length 20%–26%, snout length 8%–10%, eye diameter 3%–5%, interorbital width 5%–7%, upper jaw length 15%–16%, pre-dorsal-fin

length 27%–33%, pre-anal-fin length 50%– 56%, pectoral fin length 16%–24%, body depth 9%–13%, caudal peduncle depth 3%– 5%. Pectoral fins have 10 or 11 (usually 11) rays. First dorsal fin has 10 to 12 spines, and second dorsal fin has 1 spine and 22 to 25 rays. Anal fin has 1 spine and 22 to 25 rays. Vertebrae number 35 to 40. Color is black. This species occurs in tropical and subtropical waters of the Atlantic, Pacific, and Indian Oceans, based on capture of relatively few specimens. In the western Atlantic it occurs off North Carolina, in the northern and southern Gulf of Mexico, and off Suriname. It is usually captured at depths greater than 1,500 m. Maximum known size is 260 mm SL. references: Parr 1931 (as Dolichodon macrodon), 1933 (as Hemicyclodon macrodon); Johnson 1969; Johnson and Cohen 1974; Johnson and Keene 1986a,b; McEachran and Sutton 2002.

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CHIASMODONTIDAE 559

Kali normani

(Parr, 1931)

Kali normani has strongly arched jaws, two rows of jaw teeth, and tooth plates on first three gill arches. Dorsal profile of head is slightly convex. Snout is moderately acute. Supramaxilla is present. Middle sections of jaws are widely separated when mouth is closed. Lower jaw extends beyond upper jaw. Jaw teeth are needlelike, recurved, and usually depressible, with those of inner row barbed and considerably larger and more curved than those in outer row. Upper jaw has 16 to 38 depressible teeth in outer row and 4 or 6 long, depressible fangs in inner row. Lower jaw has 14 to 27 straight teeth in outer row and 4 or 5 long, depressible fangs in inner row. Palatine teeth are very small and needlelike and number 1 to 6. Vomerine teeth are absent. First gill arch has 12 gill teeth. Branchiostegal rays number 6. Measurements are expressed as percent of SL: head length 21%–28%, snout length 7%–10%, eye diameter 3%–5%, interorbital width 5%– 6%, upper jaw length 15%–22%, pre-dorsal-fin length 27%–33%, pre-anal-fin length 50%– 55%, pectoral fin length 15%–26%, body

depth 10%–15%, caudal peduncle depth 3%– 7%. Pectoral fins have 12 or 13 (usually 13 in the western Atlantic) rays. First dorsal fin has 11 or 12 spines, and second dorsal fin has 1 spine and 24 to 26 rays. Anal fin has 1 spine and 23 to 26 rays. Vertebrae number 36 to 40. Color is black. This species occurs in tropical and subtropical waters of the Atlantic, Pacific, and Indian Oceans, based on capture of relatively few specimens. In the western Atlantic it occurs from the east coast of Florida to Brazil, including the Gulf of Mexico and the Caribbean Sea. It occurs in the northern and eastern Gulf of Mexico. It is usually captured at depths greater than 1,000 m but occasionally is captured between 500 and 1,000 m. Maximum known size is 150 mm SL. references: Parr 1931 (as Dolichodon normani), 1933 (as Hemicyclodon normani); Johnson 1969; Johnson and Cohen 1974; Johnson and Keene 1986a,b; McEachran and Sutton 2002.

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560 PERCIFORMES

Pseudoscopelus altipinnis

Parr, 1933

Pseudoscopelus altipinnis has a short, rounded snout, elongated anterior rays of second dorsal fin and anal fin, and photophores on body and head. Upper jaw has several short, recurved, depressible teeth on side of symphysis. Side of upper jaw has outer single series of teeth and two inner bands of teeth, with teeth increasing in size both posteriorly and medially. Teeth on side of upper jaw have lanceolate crowns, with those of outermost row curved and fixed, and remainder depressible. Palatine teeth are small, needlelike, and arranged in a single series. Preoperculum has a stout, ventrally directed spine at corner. First three gill arches bear gill teeth. Branchiostegal rays number 6. Measurements are expressed as percent of SL: head length 30%–32%, snout length 8%–9%, eye diameter 5%– 6%, upper jaw length 21%–23%, interorbital width 9%–11%, body depth 17%– 20%. Pectoral fin has 13 to 15 rays. First dorsal fin has five to eight spines, and second dor-

sal fin has one spine and 23 to 26 rays. Anal fin has one spine and 24 to 26 rays. Caudal fin is deeply forked. Lateral line consists of 78 to 81 pores. Photophores occur in patches on maxilla, in two groups on lower jaw, under chin and thorax, on axil of pectoral fin, in a single row between pelvic fin insertions, along abdomen, and along anal fin base. Vertebrae number 37 or 38. Color is brownish black. Juveniles to 20 mm SL are banded. This species occurs in the tropical and subtropical North Atlantic and Pacific. In the western Atlantic it occurs in the Caribbean Sea and in the northern Gulf of Mexico off Mississippi at 914 m, and off Florida between 0 and 200 m and between 850 and 950 m. Maximum known size is 200 mm SL. references: Parr 1933; Uyeno and Aizawa 1983; Johnson and Keene 1986a; McEachran and Sutton 2002.

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CHIASMODONTIDAE 561

Pseudoscopelus obtusifrons

(Fowler, 1934)

Pseudoscopelus obtusifrons has a short, rounded snout, a greatly broadened premaxilla, and mandibular photophores arranged in three groups. Premaxilla has three uniserial rows of needlelike teeth. Lower jaw has two rows of teeth, with those of inner row larger. Teeth in both jaws have lanceolate crowns. Palatine teeth are small, needlelike, and arranged in a single series. Preoperculum has a stout, ventrally directed spine at corner. First three gill arches bear gill teeth. Branchiostegal rays number 6. Measurements are expressed as percent of SL: head length about 27%, snout length about 6%, eye diameter about 5%, upper jaw length about 20%, interorbital width about 10%, body depth about 18%. Pectoral fin has 14 rays. First dorsal fin has eight spines, and second dorsal fin has one spine and 21 to 25 rays. Anal fin has one spine and 19 to 23

rays. Caudal fin is deeply forked. Lateral line consists of 69 to 76 pores. Photophores occur in patches on maxilla, on lower jaw, under chin and thorax, on axil of pectoral fin, along uppermost pectoral fin ray, on abdomen, and on anal fin base. Photophores on tip of snout are in a single row, mandibular photophores are arranged in three groups, and those between pelvic fin origins are in a single row. Vertebrae number 37 or 38. Color is black. This species occurs in the western Atlantic and western Pacific in tropical seas between 914 and 1,097 m. In the western Atlantic it occurs in the Gulf of Mexico off Texas and Florida. Maximum known size is about 115 mm SL. references: Fowler 1934; McEachran and Sutton 2002.

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562 PERCIFORMES

Pseudoscopelus scriptus

Lütken, 1892

Pseudoscopelus scriptus has a relatively short, rounded snout; relatively short anterior rays of second dorsal and anal fins; and patches of photophores on body and head. Lower jaw extends little if any beyond upper jaw. Upper jaw has several short, recurved, depressible teeth on side of symphysis. Side of upper jaw has outer row of teeth and two inner bands of teeth, with teeth increasing in size posteriorly and medially. Teeth on side of upper jaw have lanceolate crowns, with those of outermost row curved and fixed and the remainder depressible. Palatine teeth are small, needlelike, and arranged in a single series. Preoperculum has a stout, ventrally directed spine at corner. First three gill arches bear gill teeth. Branchiostegal rays number 6. Measurements are expressed as percent of SL: head length 26%–31%, snout length 6%–7%, eye diameter 3%–5%, upper jaw length 21%–26%, interorbital width 7%–

11%, body depth 18%–23%. Pectoral fin has 13 rays. First dorsal fin has six or seven spines, and second dorsal fin has one spine and 22 to 24 rays. Anal fin has one spine and 22 to 24 rays. Caudal fin is forked. Lateral line consists of 74 pores. Photophores occur in patches on maxilla, in two groups on lower jaw, under chin and thorax, on axil of pectoral fin, along lower pectoral fin ray, on abdomen, in a broad band between pelvic fin insertions, and along anal fin base. Color is dark brown. This species occurs in the tropical and subtropical North Atlantic and Pacific. In the western Atlantic it occurs at 914 m in the northern Gulf of Mexico off Mississippi and off Campeche Bank. Maximum known size is 200 mm SL.

references: Parr 1933; Uyeno and Aizawa 1983; Bañón et al. 1999; McEachran and Sutton 2002.

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563

PERCOPHIDAE

Flatheads

Flatheads are elongate, depressed anteriorly, and compressed posteriorly, with large eyes and a narrow interorbital space. Mouth is large and terminal, with either upper jaw extending beyond lower jaw or lower jaw extending beyond upper jaw. Jaw teeth are villiform, and teeth are present or absent from vomer and palatine. Nostril is paired, and anterior and posterior nares are close set. Eyes are located on dorsal or dorsolateral aspect of head. Branchiostegal rays number 6 or 7. Gill membranes are separate and connected to isthmus. Pectoral fin has 16 to 30 rays. Dorsal fin is divided into separate spinous and rayed fins; first has 2 to 6 spines, and second dorsal fin has 13 to 23 rays. Pelvic fin consists of one spine and 5 rays and inserts anterior to pectoral fin base. Pelvic fin bases are widely separated. Anal fin is similar in size and shape to rayed dorsal fin and has 15 to 29 rays. Caudal fin is truncate or rounded and has 7 to 13 rays. Scales are ctenoid. Lateral line runs along midflank and consists of 31 to 87 scales. Gas bladder is absent. Maximum known size is 30 cm TL. Flatheads occur in the Atlantic, Indo-West Pacific, and eastern Pacific Oceans. They are benthic along the outer continental shelf to the upper slope between 80 and 900 m. Food consists of shrimps and small ray-finned fishes. There are 40 species in 13 genera, 8 species in 2 genera in the western Central Atlantic, and 2 or 3 species in a single genus in the Gulf of Mexico. references: Ginsburg 1955; Heemstra and Nelson 1986; Nelson 1994, 2001; Thompson 2002b.

Key to the Species of the Gulf of Mexico (Adapted from Ginsburg 1955) 1a. Lateral scale rows 55 to 59; pectoral fin rays 25; dorsal fin rays 14; four spots on sides . . . . . . . . . . . . . . . . . . . . . . . Bembrops macromma p. 566 1b. Lateral scale rows 61 to 71; pectoral fin rays 26 to 28; spots on sides present or absent . . . . . . . . . . . . . . . . . . . . 2 2a. Dorsal fin rays usually 15 (rarely 14); second dorsal spine filamentous in large males; 10 spots on sides, best developed in small specimens . . . . Bembrops anatirostris p. 564 2b. Dorsal fin rays usually 17 (rarely 16 or 18); second dorsal fin spine not filamentous in large males; no distinct spots on sides . . . . . . . . . . . . . . . . . . Bembrops gobioides p. 565

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564 PERCIFORMES

Bembrops anatirostris Duckbill flathead

Ginsburg, 1955

Bembrops anatirostris has a tentacle on posterior margin of maxilla and 10 spots on side. Lower jaw projects beyond upper jaw. Maxilla is exposed posteriorly and is deeply notched on posterior margin. Jaw, vomerine, and palatine teeth are villiform and arranged in bands, with those in lower jaw exposed when mouth is closed. Angle of preoperculum has two broad spinous processes. Lower limb of suboperculum has a single spine. Operculum has two diverging spines on posterior margin, a third spine near ventral margin, and an elongate flap extending from posterior margin. Posttemporal bone has a well-developed spine on posterior margin, just medial to upper margin of operculum. Branchiostegal rays number 7. Gill rakers on first arch number 4 to 6 on upper limb and 12 to 15 on lower limb. Measurements are expressed as percent of SL: head length 35%– 42%, snout length 12%–16%, eye diameter 8%–9%, upper jaw length 13%–15%, pectoral fin length 21%–24%, body depth 11%–

16%. Pectoral fin has 23 to 26 rays. First dorsal fin has six slender spines, with second spine extended as a filament in large males, and second dorsal fin has 14 or 15 rays. Anal fin has 17 or 18 rays. Caudal fin is truncate. Body and much of head are covered with ctenoid scales. Lateral line slopes ventrally and reaches lowest point at level of pectoral fin base. Lateral line scales number 60 to 68. Color is tan dorsally and light yellowish tan to white ventrally, with a series of irregularly shaped dark brown spots or blotches on midflank. This species occurs between 82 and 538 m in the western North Atlantic in the Gulf of Mexico and off Puerto Rico to northern South America. It is found throughout the Gulf of Mexico within its depth range. Maximum known size is 350 mm TL. references: Ginsburg 1955; Grey 1959; Robins and Ray 1986; Boschung 1992; Das and Nelson 1996; Thompson 2002b.

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PERCOPHIDAE 565

Bembrops gobioides Goby flathead

(Goode, 1880)

Bembrops gobioides has a narrow flap or tentacle on posterior margin of maxilla and lacks distinct spots on side. Lower jaw projects beyond upper jaw. Maxilla is exposed posteriorly and is deeply notched on posterior margin. Jaw, vomerine, and palatine teeth are villiform and arranged in bands, with those in lower jaw exposed when mouth is closed. Angle of preoperculum has two broad spinous processes. Lower limb of suboperculum has a single spine. Operculum has two diverging spines on posterior margin, a third spine near ventral margin, and an elongate flap extending from posterior margin. Posttemporal bone has a well-developed spine on posterior margin, just medial to upper margin of operculum. Branchiostegal rays number 7. Gill rakers on first arch number 5 or 6 on upper limb and 12 to 14 on lower limb. Measurements are expressed as percent of SL: head length 34%–38%, snout length 10%– 12%, eye diameter 7%–9%, upper jaw length 12%–15%, pectoral fin length 23%–26%, body depth 9%–15%. Pectoral fin has 22 to 26 rays. First dorsal fin has six slender spines,

and second dorsal fin has 16 or 17 rays. Anal fin has 17 or 18 rays. Caudal fin is truncate. Body and much of head are covered with ctenoid scales. Lateral line slopes ventrally and reaches lowest point at level of pectoral fin base. Lateral line scales number 60 to 66. Color is tan dorsally and light yellowish tan to white ventrally, with a dark reticulate pattern formed by dark pigment outlining scale pockets, a dark dorsal fin, and a dark blotch at base of caudal fin. Dark check-shaped marks may be present in juvenile specimens. This species occurs between 82 and 567 m in the western North Atlantic from New York to the Gulf of Mexico, including the Bahamas and the Antilles. It occurs throughout the Gulf of Mexico along the outer continental shelf and upper slope. Maximum known size is 220 mm SL.

references: Ginsburg 1955; Grey 1959; Robins and Ray 1986; Boschung 1992; Das and Nelson 1996; Thompson 2002b.

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566 PERCIFORMES

Bembrops macromma

Ginsburg, 1955

Bembrops macromma has a narrow flap on posterior margin of maxilla and four spots on side. Lower jaw projects beyond upper jaw. Maxilla is exposed posteriorly and bears a large, triangular tentacle on posterior tip. Jaw, vomerine, and palatine teeth are villiform and arranged in bands, with those in lower jaw exposed when mouth is closed. Angle of preoperculum has two broad spinous processes. Lower limb of suboperculum has a single spine. Operculum has two diverging spines on posterior margin, a third spine near ventral margin, and an elongate flap extending from posterior margin. Posttemporal bone has a welldeveloped spine on posterior margin, just medial to upper margin of operculum. Branchiostegal rays number 7. Gill rakers on first arch number 4 to 6 on upper limb and 12 to 15 on lower limb. Measurements are expressed as percent of SL: head length 37%– 43%, snout length 11%–15%, eye diameter 9%–10%, upper jaw length 14%–15%, pectoral fin length 22%–24%, body depth 10%–14%. Pectoral fin has 22 to 24 rays. First dorsal fin has six slender spines, and second dorsal fin has 14 or 15 rays. Anal fin has 17 or 18 rays. Caudal fin

is truncate. Body and much of head are covered with ctenoid scales. Lateral line slopes ventrally and reaches lowest point at level of pectoral fin base. Lateral line scales number 54 to 64. Color is yellowish tan, with four more or less evenly spaced elongate dark blotches on midflank, and three short and narrow dark bars between blotches located nearer to anterior than to posterior blotch. Anterior section of first dorsal fin is black, and margins of second dorsal fin and anal fin are black. This species occurs between 220 and 412 m in the western North Atlantic from the Bahamas, the Greater and Lesser Antilles, and Honduras. It has been recorded from the northern Gulf of Mexico, but according to Bruce Thompson (pers. comm., June 2002), it does not occur in the Gulf of Mexico. It is included herein because it has been captured in the Straits of Yucatán, suggesting that it may occasionally occur in the Gulf. Maximum known size is 202 mm TL. references: Ginsburg 1955; Grey 1959; Robins and Ray 1986; Das and Nelson 1996; Thompson 2002b.

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567

URANOSCOPIDAE

Stargazers

Stargazers are moderately elongate, depressed or compressed anteriorly, and moderately compressed posteriorly, with a massive and dorsally flattened head and fringed lips. Dorsal and lateral surfaces of head are encased in sculptured bone. Nostril is paired, with anterior and posterior nares close set and anterior naris forming a short tube. Eye is generally dorsally but occasionally laterally located. Interorbital space is broad. Mouth is large and strongly oblique to nearly vertical. Lips have fleshy ridges or fimbriae. Upper jaw is somewhat protrusible, and lower jaw is heavy and projects beyond upper jaw. Villiform teeth occur in jaws, prevomer, and palatine. Some species have a wormlike filament attached to floor of mouth near symphysis. Gill membranes are nearly separate and are free of isthmus. Gill arches number three and one-half, with a small slit behind last arch. Pseudobranch is present. Gill rakers are rudimentary. Branchiostegal rays number 6. Cleithral bone of shoulder girdle, above pectoral fin, possesses a spine that is either short, blunt, and covered with skin or long, sharp, and free of skin. Cleithral spine possesses two large, double-grooved poison spines in most species. Pectoral fin is broad based and fanlike and consists of 13 to 25 rays, with lower rays branched. Dorsal fin is either single, with zero to four rudimentary spines and 12 to 20 rays, or double, with first fin consisting of three to five spines and second fin consisting of 12 to 15 rays. Pelvic fin is jugular in position, inserts on isthmus, and consists of one short, occasionally obscure spine and five rays. Pelvic fin bases are close together but not connected. Anal fin is elongate and consists of zero to one spine and 12 to 19 rays. Caudal fin is fan shaped and consists of 11 to 14 principal rays. Body is naked or covered with cycloid scales. Lateral line runs along upper part of side or is obscure. One genus has internal nares and electric organs derived from eye muscles. Scales are small or absent except for pored lateral line scales. Vertebrae number 24 to 29. Maximum known size is 700 mm TL. Stargazers occur worldwide in tropical to temperate seas between the shoreline and 900 m on continental and insular shelves and upper slopes. All species are benthic, and many bury themselves in sand and muddy substrates with only their eyes protruding. They are ambush predators. The wormlike appendage of some species is apparently used to lure prey to within range of capture. Food consists of small polychaetes, crustaceans, and small benthic ray-finned fishes. There are about 50 species in eight genera, 4 species in three genera in the western Central Atlantic, and 3 species in separate genera in the Gulf of Mexico.

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568 PERCIFORMES

references: Watson et al. 1984; Heemstra 1986m; Hureau 1986; Pietsch 1989; Nelson 1994; Smith 1997; Smith-Vaniz et al. 1999; Kistimoto 2001; Carpenter 2002d.

Key to the Species of the Gulf of Mexico (Adapted from Carpenter 2002d) 1a. Spinous dorsal fin present . . . . . . . . . . . . . . . . . . . . . . Astroscopus y-graecum p. 569 1b. Spinous dorsal fin absent . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Cleithral spine, above pectoral fin base, flattened and blunt; pair of converging bony ridges at symphysis of lower jaw . . . . . . . . . . . . . . . . . . . . . Gnathagnus egregius p. 570 2b. Cleithral spine, above pectoral fin base, conical and pointed; no pair of body ridges at symphysis of lower jaw . . . . . . . . . . . . . . . . . . . . . . Kathetostoma albigutta p. 571

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URANOSCOPIDAE 569

Astroscopus y-graecum Southern stargazer

(Cuvier, 1829)

Astroscopus y-graecum has a short spinous dorsal fin and large white spots on upper part of body. Upper and lower lips bear long, slender folds (fimbriae). Anterior naris is surrounded by fimbriae. Posterior naris is extended as a fringed groove that curves behind eye, with posterior end of groove much closer to margin of eye than to anterior margin of electric organ. Subtrapezoidal depressions, lacking bone, are located posterior to groove behind each eye and cover electric organs derived from eye muscles. A Y-shaped bony ridge separates depressions. Preopercular margin is membranous. Gill slit extends from above pectoral fin base to isthmus and is connected to antimere by a fold of skin. Cleithral spine is short, blunt, and covered with skin. Measurements are expressed as percent of SL: head length 37%– 40%, snout length 6%–7%, orbit diameter 2%–3%, interorbital distance 8%–9%, upper jaw length 15%–17%, pectoral fin length 27%–30%, body depth 23%–27%. Pectoral fin has 19 to 22 rays. First dorsal fin has three to five stout spines, and second dorsal fin has

13 to 15 rays. Anal fin has 12 to 14 rays. Body is covered with small, embedded scales. Lateral line runs along dorsal surface, curves ventrally at caudal peduncle, and bends anteroventrally to unite with antimere at caudal fin base. Color is dark dorsally and light ventrally, with large, irregular, dark-margined white spots on tip of lower jaw, on dorsal section of head, and on body to insertion of dorsal fin. A black blotch is located on side of chin, and a dark stripe runs along caudal peduncle. First dorsal fin is black, and second dorsal fin is striped with one to three diagonal bars. This species occurs from near shore to about 70 m in the western Atlantic from North Carolina to Santos, Brazil, including the entire Gulf of Mexico. It is associated with softbottom substrates. Maximum known size is 440 mm TL. references: Berry and Anderson 1961; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Carpenter 2002d.

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570 PERCIFORMES

Gnathagnus egregius Freckled stargazer

(Jordan and Thompson, 1905)

Gnathagnus egregius has a pair of bony ridges at symphysis of lower jaw and lacks a spinous dorsal fin. Upper and lower lips lack fimbriae. Anterior naris is at end of a short tube and has a stout tentacle on posterior margin. Posterior naris consists of a pore medial to eye. Depressions are lacking behind each eye, but shallow recess of frontal bones extends between eyes. Preopercular margin is membranous, with a broad, triangular process at angle. Operculum has a striated and ridged surface and a spine on posterior margin. Gill slit extends from above pectoral fin base to isthmus and is connected to antimere by a fold of skin. Cleithral spine is prominent, laterally compressed, and curved posterodorsally. Measurements are expressed as percent of SL: head length 35%–39%, snout length 7%– 8%, orbit diameter 5%– 6%, interorbital distance 8%–9%, upper jaw length 14%–15%, pectoral fin length 23%–27%, body depth 19%–24%. Pectoral fin has 20 to 24 rays. Dorsal fin has 12 to 14 rays. Anal fin

has 16 or 17 rays. Body and fleshy section of caudal fin are covered with small, deeply embedded scales. Lateral line runs along dorsal surface, curves ventrally at caudal peduncle, and runs to near end of caudal fin. Color is dark gray to brown dorsally, with darker lines and spots, and light ventrally. Dorsal and pectoral fins are dark brown except for margins, which are transparent. Caudal fin is blackish except along margin, where it is light. This species occurs between 20 and 538 m in the western North Atlantic from Georgia and Bermuda to the southern Gulf of Mexico. In the northwestern Gulf of Mexico it is generally limited to the edge of the continental shelf; however, juveniles occur in shallow water to 57 m. Maximum known size is 330 mm SL. references: Berry and Anderson 1961; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Smith-Vaniz et al. 1999; Carpenter 2002d.

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URANOSCOPIDAE 571

Kathetostoma albigutta Lancer stargazer

Bean, 1892

Kathetostoma albigutta has three spines on ventral margin of preoperculum and lacks a spinous dorsal fin. Upper and lower lips are sparsely covered with short fimbriae. Anterior naris is at end of a short tube and has a multilobed tentacle on posterior margin. Posterior naris consists of a pore located anterior to eye. Depressions are lacking behind eye, but shallow recess in frontal bones extends between eyes. Dentary bones of lower jaw lack pair of prominent ridges. Preopercular margin is membranous, with three stout spines on lower margin. Operculum lacks spines and has a crenate membranous margin. Gill slit extends from above pectoral fin base to isthmus and is connected to antimere by a fold of skin. Cleithral spine is prominent, conical, sharply pointed, and directed posterodorsally. Measurements are expressed as percent of SL: head length 38%– 42%, snout length 6%– 8%, orbit diameter 4%–5%, interorbital distance 8%– 9%, upper jaw length 15%–18%, pectoral fin length 24%–29%, body depth 23%–27%. Pectoral fin has 14 to 16 rays. Dorsal fin has

13 to 15 rays. Pelvic fin base has a forwardly directed spine projecting through skin. Anal fin has 12 to 15 rays. Body lacks scales except for scalelike structures associated with lateral line. Lateral line runs along dorsal surface, curves ventrally at caudal peduncle, and runs to near end of caudal fin. Color is reddish brown dorsally, with round white spots, and white ventrally. Pectoral fin is brownish black. Dorsal fin has two to five elongate black spots, and caudal fin has three or more. This species occurs between 27 and 436 m in the western North Atlantic from Cape Lookout, North Carolina, to the southern Gulf of Mexico. It is found throughout the Gulf of Mexico within its depth range. Juvenile specimens occur in shallow water to 57 m. Maximum known size is 280 mm TL. references: Berry and Anderson 1961; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997c; Carpenter 2002.

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572

TRIPTERYGIIDAE

Tripterygiids are rather elongate, robust anteriorly, and moderately compressed posteriorly, with dorsal fin divided into three distinct segments, pelvic fin consisting of a hidden spine and several rays, and first gill arch attached to operculum by membrane. Dorsal profile of head is moderately to strongly elevated, and snout is shorter to slightly longer than eye length. Nostril is paired, with anterior and posterior nares close set, and anterior naris forming a simple pore or tube that may bear a tentacle or cirri. Orbital tentacle is present or absent and, when present, varies from simple to branched. Eye is large and located on dorsolateral aspect of head. Mouth is terminal and moderately oblique. Jaw teeth are conical and arranged in bands. Teeth are present or absent in vomer and palatine. Branchiostegal rays number 6 or 7. Pectoral fin is broad based and relatively long and has 10 to 19 rays. First dorsal fin consists of 3 to 10 spines, second consists of 8 to 26 spines, and third consists of 7 to 17 segmented rays, the last of which is separated at base. Pelvic fin consists of 1 obscure spine and 2 or 3 rays. Anal fin is long and consists of 0 to 2 spines followed by 14 to 32 rays. Caudal fin is truncate to convex and has 13 principal rays, 9 of which are branched. Body is covered with small, usually ctenoid scales. Head and belly lack scales in most but not all species. Lateral line terminates at midbody and consists of pored scales or of pored scales anteriorly and notched scales posteriorly. Males and females of the same species generally differ in coloration, but both sexes are cryptically colored. Maximum known size is 350 mm TL. All species are benthic and cryptic and range from the shoreline to about 40 m, and occasionally to 550 m. They are associated with coral reefs and rocky outcroppings. There are about 150 species in 30 genera worldwide, 5 species in a single genus in the western Central Atlantic, and 4 species in the Gulf of Mexico. references: Fritzsche and Fuiman 1982; Matarese et al. 1984; Holleman 1986; Nelson 1994; Smith 1997; Williams and Fricke 2001; Williams 2002a.

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TRIPTERYGIIDAE 573

Key to the Species of the Gulf of Mexico (Adapted from Rosenblatt 1960; Williams 2002a) 1a. Pectoral fin base and belly covered with scales; scales in first row above lateral line distinctly larger than scales above this row; segmented anal fin rays usually 15. . . . . . 2 1b. Pectoral fin base and belly naked; scales in first two rows above anterior section of lateral line subequal; segmented anal fin rays usually 16 . . . . . Enneanectes boehlkei p. 575 2a. Pectoral fin rays usually 14; pored lateral line scales usually 11 . . . . . . . . . . . . . . . . . . . Enneanectes altivelis p. 574 2b. Pectoral fin rays usually 15; pored lateral line scales usually 13 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. Anal fin uniformly pigmented; cheek behind eye naked or with 1 or 2 small cycloid scales . . . . . . . . . . . . . . . . . . . . . . . . Enneanectes jordani p. 576 3b. Anal fin with 6 or 7 bars; cheek behind eye with 3 to 8 small ctenoid scales . . . . . . . Enneanectes pectoralis p. 577

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574 PERCIFORMES

Enneanectes altivelis Lofty triplefin

Rosenblatt, 1960

Enneanectes altivelis has scales on pectoral fin base and belly and a relatively long first spine of first dorsal fin. Top of head is spinose, and lateral ethmoid and sphenotic bones are expanded into anterior and posterior flanges located above orbit. Anterior flange and nasal bone are smooth, and posterior flange is spinose. Snout is rather pointed. Anterior naris is a short tube with a lobate tentacle on posterior margin, and posterior naris is located on anterior margin of orbit and has a slightly raised rim. Orbital tentacle is longer than it is broad and slightly longer than interorbital width. Maxilla extends beyond middle of eye. Jaw teeth are conical, recurved, pointed, and arranged in bands, with outer row more widely spaced and slightly enlarged to enlarged. Vomerine teeth are arranged in a V-shaped patch, and palatine lacks teeth. Upper margin of operculum is slightly concave and bears spines; lower margin is strongly rounded and smooth. First gill arch has six or seven short gill rakers on lower limb. Measurements are expressed as percent of SL: head length 31.9%–35.4%, snout length 8.4%–11.2%, eye diameter 9.8%– 12.9%, upper jaw length 14.2%–15.8%, pectoral fin length 32%–39.8%, first spine of first dorsal fin 17.2%–22.7%, third spine of second dorsal fin 15.7%–18.9%, head depth 18.8%– 22.7%, caudal peduncle depth 9%–11.4%.

Pectoral fin rays number 13 to 15 (usually 14). First dorsal fin has 3 spines, second has 10 to 12 spines, and third has 7 or 8 rays. First dorsal fin is higher than anterior section of second dorsal fin. Pelvic fin has 1 embedded spine and 2 rays. Pelvic fins are nearly completely separate. Anal fin has 2 spines and 14 to 16 (usually 15) rays. Body and much of head are covered with ctenoid scales, and scales on belly and pectoral fin base are cycloid. Scales in first row above anterior section of lateral line are larger than those in second row. Lateral line is discontinuous and consists of an anterior section of 10 to 12 (usually 11) pored scales and a posterior section of 20 to 22 notched scales. Color in preservative is almost white to light tan, with five rather strong brown bars on side. Last bar on base of caudal fin is most distinct. This species occurs in the western Atlantic from southern Florida and the Bahamas to Nicaragua, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from the Dry Tortugas and Campeche Bay. Maximum known size is 22.6 mm SL. references: Rosenblatt 1960; Birdsong and Emery 1968; Böhlke and Chaplin 1968; Greenfield and Johnson 1981; Robins and Ray 1986; Smith 1997; Williams 2002a.

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TRIPTERYGIIDAE 575

Enneanectes boehlkei Roughhead triplefin

Rosenblatt, 1960

Enneanectes boehlkei has a naked pectoral fin base and belly and a relatively short first spine of first dorsal fin. Top of head is spinose, and lateral ethmoid and sphenotic bones are expanded into anterior and posterior flanges located above orbit. Anterior flange, posterior flange, and nasal bone are spinose. Snout is rather blunt. Anterior naris is a short tube with a lobate tentacle on posterior margin, and posterior naris is located on anterior margin of orbit and has a slightly raised rim. Orbital tentacle is longer than it is broad and slightly longer than interorbital width. Maxilla extends beyond middle of eye. Jaw teeth are conical, recurved, pointed, and arranged in bands, with teeth of outer row widely spaced and slightly to moderately enlarged. Vomerine teeth are arranged in a V-shaped patch, and palatine teeth are absent. Upper margin of operculum is slightly concave and bears spines; lower margin is strongly rounded and smooth. First gill arch has six or seven short gill rakers on lower limb. Measurements are expressed as percent of SL: head length 29.5%–32.8%, snout length 9.8%–12.3%, eye diameter 8%–11.9%, upper jaw length 12.3%–15%, pectoral fin length 35.4%– 40%, first spine of first dorsal fin 12.7%–15.9%, third spine of second dorsal fin 14%–19.3%, head depth 17.7%–21.5%, cau-

dal peduncle depth 8.9%–11.2%. Pectoral fin rays number 14 to 16. First dorsal fin has 3 spines, second has 11 to 13 spines, and third has 7 to 9 rays. Pelvic fins are nearly completely separate. Anal fin has 2 spines and 16 or 17 (usually 16) rays. Body and much of head are covered with ctenoid scales. Scales in first two scale rows above anterior part of lateral line are about the same size. Lateral line is discontinuous and consists of an anterior section of 14 to 17 (usually 15) pored scales and a posterior section of 18 to 22 notched scales. Color in preservative is light tan, with five dark, diagonal bars on side. Last bar is most distinct. Head has bar extending from eye to corner of mouth. This species occurs off southern Florida and the Bahamas to Venezuela, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from the Dry Tortugas and Veracruz. Maximum known size is 30.5 mm SL.

references: Longley and Hildebrand 1941 (as Enneapterygius jordani); Rosenblatt 1960; Caldwell 1963; Birdsong and Emery 1968; Böhlke and Chaplin 1968; Greenfield and Johnson 1981; Robins and Ray 1986; Smith 1997; Williams 2002a.

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576 PERCIFORMES

Enneanectes jordani Mimic triplefin

(Evermann and Marsh, 1899)

Enneanectes jordani has scales on pectoral fin base and belly, a relatively long and narrow orbital tentacle, and a relatively short first spine of first dorsal fin. Top of head is spinose, and lateral ethmoid and sphenotic bones are expanded into anterior and posterior flanges located above orbit. Anterior and posterior flanges are spinose. Snout is very blunt. Anterior naris is a short tube with a lobate tentacle on posterior margin, and posterior naris is located on anterior margin of orbit and has a slightly raised rim. Orbital tentacle is longer than it is broad and slightly longer than interorbital width, with tip pointed or occasionally rounded or ragged. Maxilla extends from beyond middle of eye to near end of pupil. Jaw teeth are conical, recurved, pointed, and arranged in bands, with outer row widely spaced and slightly to moderately enlarged. Vomerine teeth are arranged in a V-shaped patch, and palatine teeth are absent. Upper margin of operculum is concave, with spines on margin; lower margin is rounded and smooth. First gill arch has six or seven short gill rakers. Measurements are expressed as percent of SL: head length 31.5%–33.9%, snout length 8.9%– 11.6%, eye diameter 11.2%–12.7%, upper jaw length 13.6%–14.8%, pectoral fin length 33.7%–38%, first spine of first dorsal fin 11.6%–13.9%, third spine of second dorsal fin 14%–17.7%, head depth 20.5%–22%, caudal

peduncle depth 9.8%–11.6%. Pectoral fin rays number 14 or 15 (usually 15). First dorsal fin has 3 spines, second dorsal fin has 11 or 12 spines, and third dorsal fin has 7 or 8 rays. Pelvic fins are nearly completely separate. Anal fin has 2 spines and 15 rays. Body and much of head are covered with ctenoid or cycloid scales. Cheek either lacks scales or has 1 or 2 thin scales on upper anterior corner. Scales in first row above anterior section of lateral line are larger than those in second row. Lateral line is discontinuous and consists of an anterior section of 12 or 13 (usually 13) pored scales and a posterior section of 19 or 20 notched scales. Color in preservative is light yellowish tan, with five regular dark brown bars on side. Last bar, on base of caudal fin, is most distinct. Cheek lacks pigment except for two dark bars. Anal fin is marked with six or seven bars. This species occurs in the western Atlantic from the Florida Keys, the Gulf of Mexico, the Bahamas, Puerto Rico, and Antigua. In the Gulf of Mexico it occurs in the Dry Tortugas and off Veracruz, at Isla de Lobos. Maximum known size is 24 mm SL.

references: Longley and Hildebrand 1941 (as Enneapterygius jordani); Rosenblatt 1960; Böhlke and Chaplin 1968; Smith 1997c; Williams 2002a.

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TRIPTERYGIIDAE 577

Enneanectes pectoralis Redeye triplefin

(Fowler, 1941)

Enneanectes pectoralis has scales on pectoral fin base and belly, a relatively short and broad orbital tentacle, and a relatively short first spine of first dorsal fin. Top of head is spinose, and lateral ethmoid and sphenotic bones are expanded into anterior and posterior flanges located above orbit. Anterior flange is smooth to irregular, and posterior flange is spinose. Snout is rather blunt. Anterior naris is a short tube with a lobate tentacle on posterior margin, and posterior naris is located on anterior margin of orbit and has a slightly raised rim. Orbital tentacle is broader than it is long and as long as interorbital width, with tip truncate or broadly rounded. Maxilla extends beyond middle of eye. Jaw teeth are conical, recurved, pointed, and arranged in bands, with outer row widely spaced and slightly enlarged to enlarged. Vomerine teeth are arranged in a V-shaped patch, and palatine teeth are absent. Upper margin of operculum is slightly concave and bears spines; lower margin is strongly rounded and smooth. First gill arch has six or seven short gill rakers. Measurements are expressed as percent of SL: head length 30.9%–34.8%, snout length 9.4%–11.9%, eye diameter 9.3%–12.8%, upper jaw length 14.2%–16.5%, pectoral fin length 32.4%–39.4%, first spine of first dorsal fin 10.4%–15.7%, third spine of second dorsal fin 15.6%–19%, head depth 18.2%–23.7%, caudal peduncle depth 9.5%–11.6%. Pectoral fin rays number 15 or 16 (usually 15). First dor-

sal fin has 3 spines, second dorsal fin has 11 or 12 spines, and third dorsal fin has 6 to 8 rays. Pelvic fins are nearly completely separate. Anal fin has 2 spines and 14 to 16 (usually 15) rays. Body and much of head are covered with ctenoid or cycloid scales. Cheek has three to eight ctenoid scales on upper corner behind eye. Scales in first row above anterior section of lateral line are larger than those in second row. Lateral line is discontinuous and consists of an anterior section of 12 to 14 (usually 13) pored scales and a posterior section of 18 to 21 notched scales. Color is light tan, with five regular and vertical dark bars on side. Last bar is broader and darker than remainder and has a red posterior margin. Head has bar extending from eye to corner of mouth, and iris is red. Caudal fin is dusky, with red at base; anal fin lacks bars. Red coloration is lost in preservative. This species occurs in the western Atlantic from southern Florida, the Bahamas, the Lesser Antilles, and the eastern Gulf of Mexico. In the Gulf of Mexico it is known from southwestern Florida, including Sanibel Island. It is associated with coral reefs to a depth of 10.6 m. Maximum known size is 32.4 mm SL. references: Rosenblatt 1960; Caldwell and Caldwell 1964; Birdsong and Emery 1968; Böhlke and Chaplin 1968; Greenfield and Johnson 1981; Robins and Ray 1986; Smith 1997; Williams 2002a.

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578

DACTYLOSCOPIDAE

Sand stargazers

Sand stargazers are moderately elongate, tapering and compressed distally, with superior eyes and an oblique to subvertical mouth. Head is usually broad and deep. Dorsal profile of head is moderately convex, and ventral profile is moderately to strongly convex. Snout is short and moderately acute to blunt. Nostril is paired, with anterior naris tubular and posterior naris porelike. Eye is located on top of head and is often stalked or protractile. Lips are usually lined with fimbriae, and lower jaw projects beyond upper jaw. Jaw teeth are small, conical, and arranged into several rows. Vomer and palatine lack teeth. Upper half of operculum is fringed with 4 to 25 fleshy projections or fimbriae, and lower half of opercular apparatus is greatly expanded, with antimeres overlapping ventrally below isthmus. Gill membranes are separate and free of isthmus. Fleshy projections or fimbriae often line lips and gill opening. Pectoral fin is large, especially in mature males, and has a broad, nearly vertical base and 12 to 16 rays. Dorsal fin is long and continuous, or deeply notched between spinous and rayed sections, and consists of 5 to 18 spines and 14 to 36 rays. Some species have up to 7 free spines preceding membranous section of dorsal fin. Pelvic fin is jugular, is directed outwardly from ventral midline, and consists of 1 spine and 3 thickened, unbranched rays. Anal fin is long and consists of 2 spines and 22 to 41 rays. Dorsal and anal fins are either free of or connected to caudal fin by thin membranes. Caudal fin is truncate to strongly convex. Body is covered with large cycloid scales. Head lacks scales except in one genus. Lateral line is complete and is highly elevated over anterior section of body. Maximum known size is 150 mm TL. Sand stargazers occur in tropical and warm temperate seas of the eastern Pacific and the western Atlantic. Species are benthic and associated with sand and reef habitats from the shoreline to 137 m on continental and insular shelves. Some species inhabit estuaries. Eggs are benthic, and males of some genera carry clusters of developing eggs beneath their pectoral fins. There are about 41 species in nine genera, 16 species in six genera in the western Central Atlantic, and 4 to 6 species in three genera in the Gulf of Mexico. references: Matarese et al. 1984; Nelson 1994; Smith 1997; SmithVaniz et al. 1999.

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DACTYLOSCOPIDAE 579

Key to the Species of the Gulf of Mexico (Adapted from Dawson 1982) 1a. Dorsal fin lacking distinct anterior finlet; first preopercular canal branched, with two or more distal pores. . . . . . . . . 2 1b. Dorsal fin with isolated or semi-isolated anterior finlet; first preopercular pore unbranched, with a single distal pore .............................................3 2a. Expanded eyestalk not markedly long and slender; last lateral line scale, penultimate scale on lower part of caudal fin base, not overlying distinct notchlike depression in ventral margin of caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . Dactyloscopus moorei p. 580 2b. Expanded eyestalk relatively long and slender; last lateral line scale, terminal scale on lower part of caudal fin base, overlying distinct notchlike depression in ventral margin of caudal peduncle . . . . . . . Dactyloscopus tridigitatus p. 581 3a. Upper lip without fimbriae; pectoral fin rays modally 13; venter naked. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3b. Upper lip with fimbriae; pectoral fin rays modally 14; venter scaled. . . . . . . . . . . . . Platygillellus rubrocinctus p. 585 4a. Dorsal fin spines 11 to 15. . . . . . . . . . . . . . . . . . . . . . . . . 5 4b. Dorsal fin spines 17 to 20 . . . . . . . . Gillellus greyae p. 582 5a. Segmented dorsal fin rays 14 to 17; total lateral line scales 36 to 41. . . . . . . . . . . . . . . . . . . . Gillellus uranidea p. 584 5b. Segmented dorsal fin rays 27 to 29; total lateral line scales 47 or 48 . . . . . . . . . . . . . . . . . . . . . Gillellus healae p. 583

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580 PERCIFORMES

Dactyloscopus moorei Speckled stargazer

(Fowler, 1906)

Dactyloscopus moorei has stalked eyes, a continuous dorsal fin originating on nape, and a relatively short arched anterior section of lateral line. Dorsal profile of head is slightly concave posterior to eye, and ventral profile is broadly rounded. Eyestalk is about as long as orbit diameter. Anterior naris is tubular, and posterior naris is a single pore located on anterior rim of preorbital bone and adjacent to base of anterior naris. Mouth is strongly oblique. Pedicel of premaxilla extends beyond posterior margin of eye. Jaw teeth are small, conical, and arranged in two rows. Fimbriae are present on both lips and operculum. Opercular fimbriae are located on dorsal aspect of posterior margin, number 9 to 22, and increase in number with growth. Principal preopercular sensory pores number 3, and each is usually branched to total from 3 to 18. Measurements are expressed as percent of SL: head length about 19.7%, head width about 13.6%, orbit diameter about 2.8%, upper jaw length about 8%, pre-dorsal-fin length about 18.3%, body depth about 12.7%, caudal peduncle depth about 4.9%. Pectoral fin has 13 rays. Dorsal fin has

9 to 11 spines and 26 to 30 rays, with a notch between first 3 or 4 spines and remainder of spines. Anal fin has 2 spines and 30 to 34 rays. Caudal fin has 10 rays, and 6 to 8 of these are branched. Scales are present on body but absent on head, pectoral fin base, and vent region. Scales between dorsal fin origin and lateral line usually number 3 or 4 and number 1.5 or 2 between dorsal fin base and posterior part of lateral line arch. Lateral line scales number 43 to 47, and the pore of last lateral line scale on lower part of caudal fin base is angled posteroventrally. Caudal vertebrae number 31 to 34. Upper surface of head and body is blotched or irregularly barred with brown, and upper flank is flecked with brown, with a subtle indication of stripes along scale rows. This species occurs between 3 and 13.7 m in the western Atlantic from North Carolina to Texas. Maximum known size is 75.3 mm SL. references: Hoese and Moore 1977 (as D. tridigitatus), 1998; Dawson 1982; Robins and Ray 1986; Boschung 1992; Smith 1997; Williams 2002b.

09A-T3324 9/13/05 11:18 AM Page 581

DACTYLOSCOPIDAE 581

Dactyloscopus tridigitatus Sand stargazer

Gill, 1859

Dactyloscopus tridigitatus has stalked eyes, a discontinuous dorsal fin originating on nape, and a relatively short arched anterior section of lateral line. Dorsal profile of head is more or less straight posterior to eye, and ventral profile is broadly rounded. Eyestalk is longer than orbit diameter. Anterior naris is tubular, and posterior naris is a single pore located on anterior rim of preorbital bone and adjacent to base of anterior naris. Mouth is strongly oblique. Pedicel of premaxilla extends beyond posterior margin of eye. Jaw teeth are small, conical, and arranged in two rows. Fimbriae are present on both lips and operculum. Opercular fimbriae are located on dorsal aspect of posterior margin, number 8 to 19, and increase in number with growth. Principal preopercular sensory pores number 3, with first branched to produce 2 to 11 pores. Number of pores increases with growth. Dorsal fin has 10 to 13 (usually 11 or 12) spines and 26 to 31 (usually 27 to 29) rays, with first 3 or 4 spines separated from remainder of fin. Anal fin has 2 spines and 30 to 34 (usually 31 to 33) rays. Caudal fin has 10 rays, and 6 to 8 of these are branched. Scales are present on body but absent on head, pectoral

fin base, and vent region. Scales between dorsal fin origin and lateral line usually number 2 or 3 and number 1.5 or 2 between dorsal fin base and posterior part of lateral line arch. Lateral line scales number 43 to 47 (usually 45 or 46), and pore of last lateral line scale on lower part of caudal fin base is angled ventrally and overlies distinct notch in posterior section of caudal peduncle. Caudal vertebrae number 31 to 35. Color is tan or pale, with dorsal and dorsolateral sections of head and body blotched or mottled with tan, and 11 to 14 brownish bars between nape and caudal fin base. This species occurs from the shoreline to 29 m in the western Atlantic from southeastern Florida and Bermuda to Venezuela, including the Bahamas and the Antilles. It has been reported from the Gulf of Mexico (Smith 1997), but this record could not be confirmed. Maximum known size is 74.5 mm SL. references: Kanazawa 1952; Böhlke and Chaplin 1968; Dawson 1982; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Williams 2002b.

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582 PERCIFORMES

Gillellus greyae Kanazawa, 1952 Arrow stargazer

Gillellus greyae has a subdivided dorsal fin and a relatively long arched anterior section of lateral line. Profile of head is angular. Eye is slightly elevated but not stalked. Anterior naris is tubular, and posterior naris is a single pore located on anterior rim of preorbital bone and adjacent to base of anterior naris. Mouth is strongly oblique. Pedicel of premaxilla extends beyond posterior margin of eye. Jaw teeth are small, conical, and arranged in two rows. Fimbriae are present on lower lip and operculum and number 4 to 16 on lower lip. Principal preopercular sensory pores number three, and first is unbranched. Measurements are expressed as percent of SL: head length about 20.7%, head width about 15.1%, orbit diameter about 4.3%, upper jaw length about 9.7%, pre-dorsal-fin length about 19.4%, body depth about 15.5%, caudal peduncle depth about 4.5%. Pectoral fin has 12 to 15 (usually 13) rays. Dorsal fin has 17 to 20 spines, with first 3 in a separate finlet, and 20 to 24 rays. Anal fin has 2 spines and 31 to 35 rays. Caudal fin has 10 rays, and 7 or 8 of these are branched. Scales are present on body but absent on head, pectoral fin base, and vent region. Scales between dorsal fin origin and lateral line usually

number 2 or 3 and number 1 between dorsal fin base and posterior part of lateral line arch. Lateral line scales number 50 to 58, with 27 to 33 in arched section and 22 to 25 in straight section, and pore of last lateral line scale on lower part of caudal fin base is angled posteroventrally. Caudal vertebrae number 31 to 36. Color in preservative is pale, with six dark bars that run from near dorsal midline. Bars are reddish dark brown in life. This species occurs in the western Atlantic from southeastern Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it occurs at the Dry Tortugas and Alacrán Reef. This species has been captured between the shoreline and 27.4 m and is associated with sandy areas around coral reefs, sea-grass beds, boulders, and pilings. Maximum known size is 77.7 mm SL. references: Longley and Hildebrand 1941 (as G. semicinctus, in part); Kanazawa 1952; Böhlke and Chaplin 1968; Dawson 1982; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Williams 2002b.

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DACTYLOSCOPIDAE 583

Gillellus healae Dawson, 1982 Masked stargazer Gillellus healae has a relatively long arched anterior section of lateral line. Head is angular in profile. Eye is elevated but not stalked. Anterior naris is tubular, and posterior naris is a single pore located on anterior rim of preorbital bone and adjacent to base of anterior naris. Mouth is strongly oblique. Pedicel of premaxilla extends to posterior margin of eye. Jaw teeth are small, conical, and arranged in two rows. Fimbriae are present on lower lip and operculum and number four on lower lip. Principal preopercular sensory pores number three, and first is unbranched. Measurements are expressed as percent of SL: head length about 22.3%, head width about 12.2%, orbit diameter about 3.6%, upper jaw length about 7.3%, pre-dorsal-fin length about 19.1%, body depth about 15.4%, caudal peduncle depth about 4.7%. Pectoral fin has 13 or 14 rays. Dorsal fin has 11 to 15 spines, with first 3 in a separate finlet, and 27 to 29 rays. Anal fin has 2 spines and 31 to 33 rays. Caudal fin has 10 rays, and 7 or 8 of these are branched. Scales

are present on body but absent on head, pectoral fin base, and vent region. Scales between dorsal fin origin and lateral line are lacking. Lateral line scales number 47 or 48, with 30 or 31 in arched section and 16 or 17 in straight section. Caudal vertebrae number 32 to 34. Color in preservative is pale, with a broad, dusky mask across top and side of head above eye, and seven dark saddles on body that extend to lateral midline and run from nape to caudal fin base. Saddles are widest at lateral midline and may have pale centers and dark margins. This species occurs between 21.3 and 75 m in the western Atlantic from South Carolina to Belize, including the eastern Gulf of Mexico, the Florida Keys, Puerto Rico, and Aruba in the Lesser Antilles. In the Gulf of Mexico it is known from the Dry Tortugas to Alabama. Maximum known size is 55 mm SL. references: Dawson 1982; Robins and Ray 1986; Boschung 1992; Smith 1997; Williams 2002b.

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584 PERCIFORMES

Gillellus uranidea Warteye stargazer

Böhlke, 1968

Gillellus uranidea has a subdivided dorsal fin and a relatively long arched anterior section of lateral line. Profile of head is angular. Eye is elevated but not stalked and is surrounded by short, fleshy papillae. Anterior naris is tubular, and posterior naris is a single pore located on anterior rim of preorbital bone and adjacent to base of anterior naris. Mouth is strongly oblique. Pedicel of premaxilla extends to posterior margin of eye. Jaw teeth are small, conical, and arranged in two rows. Fimbriae are present on lower lip and operculum and number two to four (usually four) on lower lip. Principal preopercular sensory pores number three, and first is unbranched. Measurements are expressed as percent of SL: head length about 28.6%, head width about 20.4%, orbit diameter about 5.3%, upper jaw length 11.2%, pre-dorsal-fin length 21.8%, body depth 17.5%, caudal peduncle depth 5.8%. Pectoral fin has 12 to 14 rays. Dorsal fin has 11 to 15 spines, with first 3 in a separate finlet, and 14 to 17 rays. Anal fin has 2 spines and 21 to 24 rays. Caudal fin has 10 rays, and 7

or 8 of these are branched. Scales are present on body but absent on head, pectoral fin base, and vent region. Scales between dorsal fin origin and lateral line are lacking. Lateral line scales number 36 to 41, with 22 to 27 in arched section and 12 to 15 in straight section. Caudal vertebrae number 23 to 26. Color in preservative is pale, with a dark bar above pectoral fin base and generally four or five dark saddles that extend to lateral midline and run from nape to caudal fin base. Dark saddles may be pallid, with or without black flecks. This species occurs in the western Atlantic from southeastern Florida and the Bahamas to Panama. It has not been reported from the Gulf of Mexico but has been taken from Looe Key in the Florida Keys, thus it is possible that it also occurs in the eastern and possibly the southern Gulf of Mexico. Maximum known size is 37.4 mm SL. references: Böhlke 1968; Böhlke and Chaplin 1968; Dawson 1982; Robins and Ray 1986; Smith 1997; Williams 2002b.

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DACTYLOSCOPIDAE 585

Platygillellus rubrocinctus Saddle stargazer

(Longley, 1934)

Platygillellus rubrocinctus has fimbriae on lips, a subdivided dorsal fin, and a relatively long arched anterior section of lateral line. Profile of head is angular to slightly rounded. Eye is slightly elevated but not stalked. Anterior naris is tubular, and posterior naris is a single pore located on anterior rim of preorbital bone and adjacent to base of anterior naris. Mouth is moderately oblique. Pedicel of premaxilla extends to posterior margin of eye. Jaw teeth are small, conical, and arranged in two rows. Fimbriae are present on upper and lower lips and operculum, and number 7 to 21 (usually 11 to 19) on lower lip. Principal preopercular sensory pores number three, and first is unbranched. Measurements are expressed as percent of SL: head length about 24.3%, head width about 17.4%, orbit diameter about 3.9%, upper jaw length about 9.6%, predorsal-fin length 20.7%, body depth 18.6%, caudal peduncle depth 6.3%. Pectoral fin has 13 to 15 rays. Dorsal fin has 15 to 17 spines, with first 3 in a separate finlet, and 14 to 17 (usually 15 or 16) rays. Anal fin has 2 spines and 23 to 27 (usually 24 to 26) rays. Caudal

fin has 11 segmented rays. Scales are present on body, including pectoral fin base and vent region, but absent on head. Scales between dorsal fin origin and lateral line number three. Lateral line scales number 38 to 43, with 21 to 25 in arched section and 15 to 19 in straight section. Caudal vertebrae number 25 to 28. Color in preservative is pale, with four dark saddles that extend to near ventral midline and begin at nape and end at caudal fin base. First saddle may be indistinct from markings on head. Other saddles are variable in shape, and interspaces range from immaculate to spotted. This species occurs between the shoreline and 30.5 m in the western Atlantic from southeastern Florida and the Bahamas to Venezuela, including the Dry Tortugas in the eastern Gulf of Mexico and the Antilles. It is associated with rubble areas around coral reefs. Maximum known size is 47.1 mm SL. references: Longley and Hildebrand 1941 (as Gillellus rubrocinctus); Böhlke and Chaplin 1968; Dawson 1982; Robins and Ray 1986; Smith 1997; Williams 2002b.

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586

LABRISOMIDAE

Scaly blennies

Scaly blennies are elongate, moderately slender, and moderately compressed, with scales on body and cirri usually on head. Profile of head is moderately elevated. Eye is large and located near dorsal margin of head. Nostril is paired, anterior naris is tubelike and often possesses a cirrus on posterior rim, and posterior naris possesses a raised rim. Supraorbital cirrus is usually present above eye, nuchal cirrus is often present on side of nape, and cirri vary considerably in complexity. Mouth is terminal, small to moderate in size, horizontal to slightly oblique, and often projecting. Lips are fleshy and have free margins. Jaw teeth are caninelike, variable in size, and arranged in one to several rows or in bands. Teeth are usually present in vomer and palatine. First gill arch has gill rakers. Opercular membranes are connected beneath head. Pectoral fin is minute to well developed, with a narrow to broad, nearly vertical base and 4 to 14 rays. Dorsal fin is long and continuous, with 17 to 31 spines and 0 to 13 rays. Dorsal fin spines are more numerous than rays. Pelvic fin is in jugular location and consists of 1 embedded spine and 2 or 3 rays. Anal fin is long and consists of 2 usually flexible spines and 18 to 38 rays. All fin rays are unbranched. Caudal fin is truncate to convex. Cycloid or ctenoid scales are present on body. Lateral line is complete. Maximum known size is 250 mm TL. Scaly blennies occur in the tropical to warm temperate seas of the western Atlantic and eastern Pacific. They are benthic fishes generally associated with coral and rocky reefs in shallow water. There are about 97 species in 16 genera, 41 species in 6 genera in the western Central Atlantic, and 19 or 20 species in 5 genera in the Gulf of Mexico. references: Matarese et al. 1984; Springer 1993; Nelson 1994; Smith 1997; Smith-Vaniz et al. 1999; Williams 2002c.

Key to the Species of the Gulf of Mexico (Adapted from Springer 1955b, 1959c; Springer and Gomon 1975; Williams 2002c) 1a. Dorsal fin elements consisting only of spines or spines and 1 ray. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Dorsal fin elements consisting of spines and 7 to 37 segmented rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2a. Pelvic fins with 1 spine and 3 rays; last dorsal fin element is a segmented ray (except rarely in P. nigripinnis last element is spinous) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2b. Pelvic fins with 1 spine and 2 rays; last dorsal fin element spinous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

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LABRISOMIDAE 587

3a. Opercular spine ending in one point and extending beyond vertical line from base of third dorsal fin spine . . . . . . . . . . . . . . . . . . . . . . Paraclinus marmoratus p. 605 3b. Opercular spine ending in two to eight points and not extending to vertical line from base of third dorsal fin spine . . . . . . . . . . . . . . . . . . . . . . . Paraclinus nigripinnis p. 606 4a. Pectoral fin base without scales; anal fin with 2 spines and 15 or 16 rays; pectoral fin rays usually 12 . . . . . . . . . . . . . . . . . . . . . . . . Paraclinus cingulatus p. 603 4b. Pectoral fin base with two or three scale rows; anal fin with 2 spines and 17 to 20 rays; pectoral fin rays usually 13 . . . . . . . . . . . . . . . . . . . . . . . . . Paraclinus fasciatus p. 604 5a. Two or more cirri on nape just anterior to dorsal fin origin (except in Malacoctenus macropus, which usually has 15 pectoral fin rays and lacks palatine teeth); pelvic fin rays 3 (innermost ray may be short and folded over middle); scales in lateral line series 40 to 69 (some pores in posterior section may lack sensory tubes but are included in count); maximum size 29 to 176 mm SL . . . . . . . . . . . . . 6 5b. Either no cirri or single cirrus on each side of nape just anterior to dorsal fin origin; pelvic fin rays 2 (3 pelvic rays in Starksia, but third ray is minute); scales in lateral series 35 to 41 (tubed scales may be present only in anterior part of lateral line; where tubes are present in posterior portion, some scales may lack tubes; counts given here refer only to sum of tubed scales in both portions of lateral line); maximum size 45 mm SL . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 6a. Posterior end of maxilla exposed when mouth closed; patches of small teeth behind outer row of large teeth in upper jaw and occasionally in lower jaw; teeth present or absent in palatine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 6b. Posterior end of maxilla sheathed when mouth closed; small teeth behind outer row of large teeth absent in most species; teeth absent in palatine . . . . . . . . . . . . . . . . . . . 18 7a. Palatine teeth present, and several considerably larger than vomerine teeth (in some specimens of L. haitiensis teeth may be about equal in size, but these specimens have 14 pectoral fin rays) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7b. Palatine teeth absent or, if present, same size or smaller than those of vomer . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 8a. Pectoral fin rays usually 14; shortest pelvic fin ray less than half the length of longest . . . . Labrisomus haitiensis p. 594 8b. Pectoral fin rays usually 13; shortest pelvic fin ray longer than half the length of longest . . . . . . . . . . . . . . . . . . . . . 9 9a. Dorsal fin spines usually 20; anal fin rays usually 20; lateral line scales 45 to 48; first dorsal fin spine longer than fifth; peritoneum uniformly dusky . . . . . . . . . . . . . . . . . . . . . . Labrisomus bucciferus p. 591

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588 PERCIFORMES

9b. Dorsal fin spines usually 19; anal fin rays usually 19; lateral line scales 48 to 53; first dorsal fin spine shorter than fifth; peritoneum white, with scattered melanophores. . . . . . . 10 10a. Specimens longer than 40 mm SL. . . . . . . . . . . . . . . . . . 11 10b. Specimens 28 to 40 mm SL . . . . . . . . . . . . . . . . . . . . . . 13 11a. Pores at symphysis of lower jaw two . . . . . . . . . . . . . . . . . . . . . . . Labrisomus kalisherae p. 595 11b. Pores at symphysis of lower jaw more than two . . . . . . . 12 12a. Operculum with ocellus; maximum known size 88 mm SL . . . . . . . . . . . . . . . . . . . . . . . . . Labrisomus guppyi p. 593 12b. Operculum without ocellus; maximum known size 49 mm SL . . . . . . . . . . . . . . . . . . Labrisomus gobio p. 592 13a. Opercular ocellus absent . . . . . . . . . . . . . . . . . . . . . . . . 14 13b. Opercular ocellus present . . . . . . . . . . . . . . . . . . . . . . . . 15 14a. Dorsal and anal fins pale or with very faint markings . . . . . . . . . . . . . . . . . . . . . . . . . . Labrisomus gobio p. 592 14b. Dorsal and anal fins usually heavily spotted . . . . . . . . . . . . . . . . . . . . . . . Labrisomus kalisherae p. 595 15a. Gill rakers on first arch usually 11 . . . . . . . . . . . . . . . . . . . . . . . Labrisomus kalisherae p. 595 15b. Gill rakers on first arch usually 13 or 14 . . . . . . . . . . . . . . . . . . . . . . . . . Labrisomus guppyi p. 593 16a. Opercular ocellus absent . . . . Labrisomus albigenys p. 590 16b. Opercular ocellus present . . . . . . . . . . . . . . . . . . . . . . . . 17 17a. Palatine teeth absent . . . . . Labrisomus nigricinctus p. 596 17b. Palatine teeth present . . . . . Labrisomus nuchipinnis p. 597 18a. Pectoral fin rays 15 to 17; jaws with small teeth behind large teeth in outer row; scales present on pectoral fin base and scales same size as body scales . . . . . . . . . . . . . . . . . . . . . Malacoctenus macropus p. 600 18b. Pectoral fin rays usually 14; scales present or absent on pectoral fin base, and if present, smaller than those on body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 19a. Side of body with three to five dark inverted-triangular bars; at least one mid-dorsal scale anterior to dorsal fin origin; small scales on flank anterior to pectoral fin base . . . . . . . . . . . . . . . . . . . . Malacoctenus triangulatus p. 601 19b. Side of body lacking three to five dark inverted-triangular bars; mid-dorsal scale anterior to dorsal fin origin lacking; no scales on flank anterior to pectoral fin base . . . . . . . . 20 20a. Dark ocellus on anterior aspect of dorsal fin and on posterior aspect of spinous section of dorsal fin and extending onto dorsal aspect of flank; scales on breast same size as those on side; last dorsal fin rays joined to caudal fin base by membrane . . . . . . . . . . . . . . . Malacoctenus gilli p. 599 20b. Dark ocelli absent on dorsal fin and flank; scales absent on breast; last dorsal fin ray not connected to caudal fin base by membrane. . . . . . . . . Malacoctenus aurolineatus p. 598

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LABRISOMIDAE 589

21a. Pectoral fin rays 11 or 12 (usually 12); no cirrus on anterior naris; central pectoral fin rays elongated and filamentous; first anal fin spine of males shorter than second spine . . . . . . . . . . . . . . . . . . . . . . . . Nemaclinus atelestos p. 602 21b. Pectoral fin rays 13 to 15 (usually 14); cirrus present on anterior naris; pectoral fin rays not elongated or filamentous; first anal fin spine of males longer than second. . . . 22 22a. Belly entirely naked . . . . . . . . . . . . Starksia ocellata p. 608 22b. Belly entirely scaled . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23a. Body tan, with eight or nine irregular, widely spaced dark bands; stripe extending from third band to caudal fin; anal fin with two spines and 18 or 19 rays; total dorsal fin elements 29 or 30 . . . . . . . . . . . . . . . . Starksia starcki p. 609 23b. Body plain colored or bands present but narrowly spaced and not markedly in contrast with ground color; no stripe along posterior section of body; anal fin with two spines and 16 to 19 (usually 17) rays; total dorsal fin elements 27 to 29 (usually 28) . . . . . . . . . . . . Starksia lepicoelia p. 607

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590 PERCIFORMES

Labrisomus albigenys Whitecheek blenny

Beebe and Tee-Van, 1928

Labrisomus albigenys has an exposed maxilla, widely separated nuchal cirri, and anterior pores of posterior lateral line scales exposed. Dorsal profile of head is slightly and evenly curved to dorsal fin origin. Snout is moderately acute. Anterior naris is tubular and has a simple to multifid flat cirrus on posterior margin. Cirrus becomes branched with growth. Supraorbital cirrus is flattened and multifid. Nuchal cirrus is flattened, oriented perpendicular to body axis, and multifid. Maxilla extends to near pupil. Lips are fleshy and protruding, and upper one has a continuous fold. Jaws possess an outer row of canine teeth and an inner row of very small teeth. Teeth occur in vomer but not in palatine. Gill rakers on first arch number 13 or 14. Measurements are expressed as percent of SL: head length 33.8%–34.4%, snout length 7.8%– 8.7%, upper jaw length 11.5%–13.4%, pre-dorsal-fin length 26.9%–

29.9%, pre-anal-fin length 52.6%–53.7%, body depth at pelvic fin origin 24%–28.8%. Pectoral fin has 13 rays. Dorsal fin has 18 spines and 11 rays, with first spine less than 8% of SL and usually shorter than other spines. Pelvic fin has 1 spine and 3 rays, with third ray vestigial. Anal fin has 2 spines and 18 rays. Lateral line scales number 40 to 44. Color is dusky brown, with scattered darker spots and eight faint bars. Fins are darkly pigmented or spotted. This species occurs in the western Atlantic from the southern Gulf of Mexico and the Greater Antilles to Venezuela. In the Gulf of Mexico it occurs on Campeche Bank. Maximum known size is 52.4 mm SL.

references: Springer 1959c; Smith 1997; Williams 2002c.

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LABRISOMIDAE 591

Labrisomus bucciferus Puffcheek blenny

Poey, 1868

Labrisomus bucciferus has an exposed maxilla, widely separated nuchal cirri, and a relatively long first dorsal fin spine. Dorsal profile of head is convex to dorsal fin origin. Snout is moderately blunt. Anterior naris is tubular and has a simple to multifid flat cirrus on posterior margin. Cirrus becomes branched with growth. Supraorbital cirrus is flattened and multifid. Nuchal cirrus is flattened and oriented perpendicular to body axis and has a number of fine fimbriae. Maxilla extends to near posterior margin of orbit. Lips are fleshy, and upper lip has a continuous fold. Jaws possess an outer row of canines and an inner row of very small teeth. Teeth occur in vomer and palatine, and some of those in palatine are larger than those in vomer. Gill rakers on first arch number 11 to 14. Measurements are expressed as percent of SL: head length 30%–35%, snout length 6%– 8.2%, upper jaw length 14.4%–19%, pre-dorsal-fin length 23.7%–26.5%, pre-anal-fin length 47.5%– 54.4%, body depth at pelvic fin origin 25%– 28.6%. Pectoral fin has 12 to 14 rays. Dorsal fin has 19 to 21 (usually 20) spines and 10

to 12 rays, with first spine usually more than 10.5% of SL and longer than third, fourth, fifth, and last 2 spines. Pelvic fin has 1 spine and 3 rays, with third ray more than half the length of longest ray. Anal fin has 2 spines and 19 to 21 [usually 20] rays. Lateral line scales number 45 to 48. More than two pores occur on symphysis of lower jaw. Color is greenish to reddish brown, with four or five dark, irregular bands that are darker ventrally than dorsally. Males have red fins, and females have densely spotted fins. Peritoneum is dusky. This species occurs between 1 and 4 m in the western Atlantic from southern Florida and Bermuda to Nicaragua, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is known only from the Dry Tortugas in the Gulf of Mexico. Maximum known size is 90 mm TL. references: Longley and Hildebrand 1941; Hubbs and Springer 1954; Springer 1959c; Böhlke and Chaplin 1968; Greenfield and Johnson 1981; Robins and Ray 1986; Randall 1996; Smith 1997; Smith-Vaniz et al. 1999; Williams 2002c.

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592 PERCIFORMES

Labrisomus gobio Palehead blenny

(Valenciennes, 1836)

Labrisomus gobio has an exposed maxilla, widely separated nuchal cirri, and an unmarked operculum. Dorsal profile of head is slightly rounded to dorsal fin origin. Snout is moderately blunt. Anterior naris is tubular and has a simple to multifid flat cirrus on posterior margin. Cirrus becomes branched with growth. Supraorbital cirrus is flattened and multifid. Nuchal cirrus is multifid and has four or five branches. Maxilla extends to posterior margin of orbit. Lips are fleshy, and upper lip has a continuous fold. Jaws possess an outer row of canines and an inner row of very small teeth. Teeth occur in vomer and palatine, and some of those in palatine are larger than those in vomer. Gill rakers on first arch number 11 to 14 (usually 13 or 14). Measurements are expressed as percent of SL: head length 28.1%–33.5%, snout length 5.8%– 8.4%, upper jaw length 12.7%–16%, pre-dorsal-fin length 24.5%–28.5%, pre-anal-fin length 48.1%–51.2%, body depth at pelvic fin origin 22.4%–25.2%. Pectoral fin has 12 or 13 rays. Dorsal fin has 18 to 20 (usually 19) spines and

10 to 12 rays, with first spine usually about 10% of SL, shorter than fourth and fifth spines, and longer than penultimate spine. Pelvic fin has 1 spine and 3 rays, with third ray more than half the length of longest ray. Anal fin has 2 spines and 18 to 20 (usually 19) rays. Lateral line scales number 48 to 53 (usually 49 to 51). More than two pores occur on symphysis of lower jaw. Color is tan to light brown, with four or five darker bars on sides. Bars are broader and darker dorsally than ventrally. Ventral section of head and body is plain and light colored. Fins are colorless in males, but dorsal, anal, and caudal fins are lightly speckled in females. This species occurs in the western Atlantic from southeastern Florida and the Bahamas to Nicaragua, including the southern Gulf of Mexico and the Greater and Lesser Antilles. Maximum known size is 49 mm SL. references: Hubbs and Springer 1954; Springer 1959c; Robins and Ray 1986; Smith 1997; Williams 2002c.

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LABRISOMIDAE 593

Labrisomus guppyi Mimic blenny

(Norman, 1922)

Labrisomus guppyi has an exposed maxilla, widely separated nuchal cirri, and a black blotch on operculum. Dorsal profile of head is slightly rounded to dorsal fin origin. Snout is moderately blunt. Anterior naris is tubular and has a simple to multifid flat cirrus on posterior margin. Cirrus becomes branched with growth. Supraorbital cirrus is flattened and multifid. Nuchal cirri have four or five branches. Maxilla extends to near posterior margin of orbit. Lips are fleshy, and upper lip has a continuous fold. Jaws possess an outer row of canines and an inner row of very small teeth. Teeth occur in vomer and palatine, and some of those in palatine are larger than those in vomer. Gill rakers on first arch number 12 to 15 (usually 13 or 14). Measurements are expressed as percent of SL: head length 31%– 35.7%, snout length 7.8%–10.6%, upper jaw length 15.2%–17.5%, pre-dorsal-fin length 27.8%–29.7%, pre-anal-fin length 53.6%– 57.4%, body depth at pelvic fin origin 25%– 27.2%. Pectoral fin has 12 to 14 rays. Dorsal fin has 18 or 19 (usually 19) spines and 10 to 12 rays, with first spine rarely more than 10% of SL and usually shorter than all but third and penultimate spines. Pelvic fin has 1 spine and

3 rays, with third ray more than half the length of longest ray. Anal fin has 2 spines and 18 to 20 (usually 19) rays. Lateral line scales number 48 to 53 (usually 49 to 51). More than two pores occur on symphysis of lower jaw. Color of males is dark, with five to six darker bars on sides that are often obscured by dark background coloration, and fins are reddish, with few spots. Females are pale greenish, with fins heavily spotted. All specimens have an ocellus on operculum. Peritoneum is white, with scattered melanophores. This species occurs between 3 and 6 m in the western Atlantic from southern Florida and the Bahamas to Brazil, including the Gulf of Mexico, the Florida Keys, and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from the Dry Tortugas, Veracruz, and Campeche Bay. Maximum known size is 88 mm SL.

references: Longley and Hildebrand 1941; Springer 1959b,c; Cervigón 1966; Birdsong and Emery 1968; Böhlke and Chaplin 1968; CastroAguirre 1978; Greenfield and Johnson 1981; Robins and Ray 1986; Smith 1997; Castro-Aguirre et al. 1999; Williams 2002c.

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594 PERCIFORMES

Labrisomus haitiensis Longfin blenny

Beebe and Tee-Van, 1928

Labrisomus haitiensis has an exposed maxilla, widely separated nuchal cirri, and a relatively short third pelvic ray. Dorsal profile of head is slightly rounded to dorsal fin origin. Snout is moderately blunt. Anterior naris is tubular and has a simple, flat cirrus on posterior margin. Supraorbital cirrus is flattened and multifid. Nuchal cirrus is multifid and has four or five branches. Maxilla extends to near posterior margin of orbit. Lips are fleshy, and upper lip has a continuous fold. Jaws possess an outer row of canines and an inner row of very small teeth. Teeth occur in vomer and palatine, and some of those in palatine are larger than those in vomer. Gill rakers on first arch number 10 to 13. Measurements are expressed as percent of SL: head length 29.5%–31.7%, snout length 6.5%– 8.2%, upper jaw length 12.8%–15.9%, pre-dorsal-fin length 23.6%– 25.7%, pre-anal-fin length 47.9%–55%, body depth at pelvic fin origin 20.1%–21.5%. Pectoral fin has 13 to 15 (usually 14) rays. Dorsal fin has 20 spines and 10 to 12 rays, with first spine greater than 10% of SL and longer than

third, fourth, and penultimate spines. Pelvic fin has 1 spine and 3 rays, with third ray less than half the length of longest ray. Anal fin has 2 spines and 18 to 22 rays (usually 20 or 21). Lateral line scales number 43 to 46. Color is tan, with six or seven dark brown bars on body that extend from dorsal fin base to belly or anal fin base, and extend in less intensity from base to margin of dorsal fin. Dorsal and anal fins are spotted between bars. This species occurs to a depth of 15 m in the western Atlantic from southern Florida and the Bahamas to Honduras, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from southwestern Florida, the Dry Tortugas, and Campeche Bay. Maximum known size is 70 mm TL. references: Longley and Hildebrand 1941; Springer 1959b,c; Birdsong and Emery 1968; Böhlke and Chaplin 1968; Smith et al. 1975; Smith 1976; Robins and Ray 1986; Greenfield and Johnson 1981; Smith 1997; Williams 2002c.

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LABRISOMIDAE 595

Labrisomus kalisherae Downy blenny

(Jordan, 1904)

Labrisomus kalisherae has an exposed maxilla, widely separated nuchal cirri, and a relatively long first dorsal fin spine and a short penultimate one. Dorsal profile of head is slightly rounded to dorsal fin origin. Snout is moderately blunt. Anterior naris is tubular and has a simple to multifid flat cirrus on posterior margin. Cirrus becomes branched with growth. Supraorbital cirrus is flattened and multifid. Nuchal cirrus is multifid and has four or five branches. Maxilla extends to near posterior margin of orbit. Lips are fleshy, and upper lip has a continuous fold. Jaw possesses an outer row of canines and an inner row of very small teeth. Teeth occur in vomer and palatine, and some of those in palatine are larger than those in vomer. Gill rakers on first arch number 10 to 13 (usually 11). Measurements are expressed as percent of SL: head length 31.5%– 35.8%, snout length 7%–7.9%, upper jaw length 15.1%–18.9%, pre-dorsal-fin length 26.4%–27.8%, pre-anal-fin length 49.5%– 54.8%, body depth at pelvic fin origin 22.4%– 26.3%. Pectoral fin has 13 or 14 (usually 13) rays. Dorsal fin has 18 to 20 (usually 19) spines and 10 to 12 rays, with first spine greater than 10% of SL and longer than third, fourth, and penultimate spines. Pelvic fin has 1 spine and 3

rays, with third ray more than half the length of longest ray. Anal fin has 2 spines and 18 to 20 (usually 19) rays. Lateral line scales number 48 to 53. Two pores occur on symphysis of lower jaw. Color is tan, with five or six dark brown bars on body that extend from dorsal fin base to belly or anal fin base and continue, with less intensity, onto basal half of dorsal fin. Pectoral and anal fins are not heavily pigmented. Juveniles have a distinct blotch on operculum. Peritoneum is white, with scattered melanophores. This species occurs in the western Atlantic from southern Florida to southern Brazil, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs between 1.5 and 4.6 m in the Dry Tortugas, off Veracruz, and on Campeche Bank. It is associated with coarse coral rubble and thick algal mats. Maximum known size is 69 mm SL.

references: Longley and Hildebrand 1941 (as L. guppyi, in part); Springer 1955b, 1959c; Caldwell and Caldwell 1964; Cervigón 1966; Birdsong and Emery 1968; Greenfield and Johnson 1981; Robins and Ray 1986; Smith 1997; Williams 2002c.

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596 PERCIFORMES

Labrisomus nigricinctus Spotcheek blenny

Howell Rivero, 1936

Labrisomus nigricinctus has an exposed maxilla, widely separated nuchal cirri, and an ocellus on operculum. Dorsal profile of head is slightly and evenly curved to dorsal fin origin. Snout is moderately acute. Anterior naris is tubular and has a simple to multifid flat cirrus on posterior margin. Cirrus becomes branched with growth. Supraorbital cirrus is flattened and multifid. Nuchal cirrus is flattened, oriented perpendicular to body axis, and multifid. Maxilla extends to near pupil. Lips are fleshy and protruding, and upper lip has a continuous fold. Jaws possess an outer row of canines and an inner row of very small teeth. Teeth occur in vomer but not in palatine. Gill rakers on first arch number 9 to 12 (usually 10 or 11). Measurements are expressed as percent of SL: head length 30.3%–34.9%, snout length 6.9%– 8.9%, upper jaw length 9.7%–11.6%, pre-dorsal-fin length 28.2%–31.2%, pre-analfin length 53.5%–59.5%, body depth at pelvic fin origin 23.3%–26.9%. Pectoral fin has 13 rays. Dorsal fin has 17 to 19 spines, with first spine less than 8% of SL and usually shorter than other spines, and 10 to 12 rays. Pelvic fin

has 1 spine and 3 rays, with third ray vestigial. Anal fin has 2 spines and 17 to 20 (usually 18) rays. More than two pores occur on symphysis of lower jaw. Lateral line scales number 40 to 44 (usually 41 or 42). Females and immature males have eight or nine dark brown bands on side extending onto dorsal and anal fins, and bands are separated by narrow, light interspaces. Operculum has a large ocellus, and two dark lines radiate from posterior margin of eye. Adult males are uniformly lightly pigmented, with some dark markings on body, a large ocellus on operculum, and mottling on fins. This species occurs in shallow water in the western Atlantic from southern Florida and the Bahamas to Venezuela, including the Florida Keys, the Dry Tortugas, the southern Gulf of Mexico, and the Greater and Lesser Antilles. It is associated with clear water in tide pools and coral reefs. Maximum known size is 54 mm SL. references: Longley and Hildebrand 1941; Springer 1959c; Robins and Ray 1986; Smith 1997; Williams 2002c.

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LABRISOMIDAE 597

Labrisomus nuchipinnis Hairy blenny

(Quoy and Gaimard, 1824)

Labrisomus nuchipinnis has an exposed maxilla, widely separated nuchal cirri, and a black blotch on operculum. Dorsal profile of head is rather steep to orbit. Snout is moderately blunt. Anterior naris is tubular and has a simple to multifid flat cirrus on posterior margin. Cirrus becomes branched with growth. Supraorbital cirrus is flattened and multifid. Nuchal cirrus is multifid and has four or five branches. Maxilla extends to near pupil. Lips are fleshy, and upper lip has a continuous fold. Jaw teeth are arranged into an outer row of canines and an inner row of very small teeth. Teeth occur in vomer and palatine and are about equal in size. Gill rakers on first arch number 10 to 13 (usually 11 or 12). Measurements are expressed as percent of SL: head length 28%–31%, snout length 9.1%–10.2%, upper jaw length 12.7%– 17.4%, pre-dorsal-fin length 24.3%–26.2%, pre-anal-fin length 50.2%–53.5%, body depth at pelvic fin origin 24.4%–29.4%. Pectoral fin has 13 to 15 (usually 14) rays. Dorsal fin has 17 to 20 (usually 18) spines and 10 to 13 (usually 12) rays, with first spine less than 9% of SL and usually shorter than other spines in specimens over 79 mm SL. Pelvic fin has 1 spine and 3 rays, with third ray more than half the length

of longest ray. Anal fin has 2 spines and 17 to 19 (usually 18 or 19) rays. Lateral line scales number 64 to 69. Color is tan, with light brown bands, to nearly uniformly black, with red fins. Males tend to be darker and more uniformly colored than females. Both sexes have a dark opercular blotch surrounded by a paler ring and a dark spot between first and third or fourth dorsal fin spines except for juveniles less than 25 mm SL. This species occurs in the tropical Atlantic. In the western Atlantic it occurs in less than 3 m from Florida and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with rocky areas and coarse coral rubble. Maximum known size is 180 mm SL.

references: Longley and Hildebrand 1941; Baughman 1947; Springer 1959b,c; Birdsong and Emery 1968; Böhlke and Chaplin 1968; CastroAguirre 1978; Castro-Aguirre and MárquezEspinoza 1981; Greenfield and Johnson 1981; Robins and Ray 1986; Randall 1996; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Williams 2002c.

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598 PERCIFORMES

Malacoctenus aurolineatus Goldline blenny

Smith, 1957

Malacoctenus aurolineatus has nearly symmetrical dorsal and ventral profiles; maxilla covered posteriorly; and a short, bony point projecting from dorsolateral margin of cleithrum. Snout is slightly spatulate. Anterior naris is tubular and has a simple cirrus on posterior margin. Supraorbital cirrus is short, slender, and biramous. Nuchal cirrus is moderately long and multifid and has 11 branches. Maxilla extends to anterior margin of orbit. Lips are fleshy, and upper lip has a continuous fold. Jaw teeth are canines arranged in a single row. Teeth occur in vomer but not in palatine. Gill rakers on first arch number 10 to 12. Measurements are expressed as percent of SL: head length 24.4%–30.4%, snout length 7.6%– 11.3%, orbital diameter 7.5%–10.4%, upper jaw length 8.1%–10.6%, body depth at pelvic fin origin 22.3%–24.4%. Pectoral fin has 13 to 15 rays. Dorsal fin has 18 to 21 (usually 19 or 20) spines and 10 to 12 rays, with first spine usually more than 10% of SL and longer than second or third and last 4 spines. Pelvic fin has 1 spine and 3 rays, with third ray usually no more than half the length of longest ray. Anal fin has 2 spines and 17 to 21 (usually 19

or 20) rays. Body is covered with ctenoid scales except for dorsal midline anterior to dorsal fin, prepectoral region, and breast. Scales on belly are smaller than those on flank. Lateral line scales number 42 to 55 (usually 48 to 53). Color is dark tan, with six rectangular dark brown bands: first two bands often coalesce on dorsal part of body to form humeral blotch, first three bands extend to ventral midline, and last three extend below midline as dark subquadrate splotches. Yellow lines occur along lower side of body. Fins lack pigment except for dusky streak along dorsal fin base. This species occurs to a depth of 6 m in the western Atlantic from southern Florida and the Bahamas to Venezuela, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs at the Dry Tortugas and Campeche Bank. It is associated with rock-boring sea urchins along rocky shorelines. Maximum known size is 60 mm SL.

references: Springer 1959b,c; Böhlke and Chaplin 1968; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Williams 2002c.

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LABRISOMIDAE 599

Malacoctenus gilli Dusky blenny

(Steindachner, 1867)

Malacoctenus gilli has symmetrical dorsal and ventral profiles, maxilla covered posteriorly, and dark ocellated spots on dorsal fin and upper flank. Snout is slightly spatulate. Anterior naris is tubular and has a simple cirrus on posterior margin. Supraorbital cirrus is long, slender, and biramous. Nuchal cirrus is long and multifid and has three to seven branches. Maxilla extends to anterior margin of orbit. Lips are fleshy, and upper lip has a continuous fold. Jaw teeth consist of a row of canines. Teeth occur in vomer but not in palatine. Gill rakers on first arch number 9 to 12. Measurements are expressed as percent of SL: head length 26%–30%, snout length 8.1%–9.4%, upper jaw length 8.8%–10.7%, pre-dorsal-fin length 26.1%–28.8%, pre-anal-fin length 49.9%– 51.9%, body depth at pelvic fin origin 22.9%– 25.1%. Pectoral fin has 13 to 16 (usually 14) rays. Dorsal fin has 18 to 21 (usually 20) spines and 9 to 11 rays, with first spine usually more than 11% of SL and longer than third, fourth, and last 4 spines. Pelvic fin has 1 spine and 3 rays, with third ray usually no more than half the length of longest ray. Anal fin has 2 spines and 17 to 21 (usually 18 to 20) rays. Body,

including breast, is covered with ctenoid scales except for dorsal midline anterior to dorsal fin and prepectoral region. Lateral line scales number 42 to 47. Color is tan to dark brown, with five to seven bands on flank that range from moderately developed to obscure. One dark ocellus is located on anterior section of dorsal fin, and a second dark ocellus is located on flank at bases of last several dorsal fin spines and on dorsal fin. Males are reddish dorsally and greenish ventrally, with obscure bands, or very dark and without bands; fins are dusky to unpigmented. Females have better-defined bands and are spotted on both body and fins. This species occurs in the western Atlantic from the western Gulf of Mexico and Bermuda to Venezuela, including the Bahamas and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from Tampico. Maximum known size is 75 mm TL. references: Springer 1959b,c; Cervigón 1966; Birdsong and Emery 1968; Böhlke and Chaplin 1968; Greenfield and Johnson 1981; Smith 1997; Smith-Vaniz et al. 1999; Williams 2002c.

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600 PERCIFORMES

Malacoctenus macropus Rosy blenny

(Poey, 1868)

Malacoctenus macropus has symmetrical dorsal and ventral profiles, maxilla covered posteriorly, and normal-sized scales preceding pectoral fin. Snout is moderately spatulate. Anterior naris is tubular and has a simple cirrus on posterior margin. Supraorbital cirrus is slender and simple. Nuchal cirrus is oriented perpendicular to body axis; has a single long, slender filament; and is separated from its antimere by distance equal to half the base of cirrus. Maxilla extends to anterior margin of orbit. Lips are fleshy, and upper lip has a continuous fold. Jaw teeth are arranged into an outer row of canines and an inner row of very small teeth. Teeth occur in vomer but not in palatine. Gill rakers on first arch number 8 to 11. Measurements are expressed as percent of SL: head length 24.5%–28.6%, snout length 7%–10%, upper jaw length 8.5%–9.6%, predorsal-fin length 23.5%–27.9%, pre-anal-fin length 42.6%– 47.8%, body depth at pelvic fin origin 23.8%–27.8%. Pectoral fin has 14 to 17 (usually 15) rays. Dorsal fin has 21 to 23 spines and 8 to 11 rays, with first spine 11.3%–18.8% of SL and longer than second through fifth spines. Pelvic fin has 1 spine and 3 rays, with third ray no more than half the length of long-

est ray. Anal fin has 2 spines and 18 to 22 (usually 20 or 21) rays. Body is covered with ctenoid scales except for midline anterior to dorsal fin. Lateral line scales number 40 to 45. Females are tan, with irregular brown blotches on upper body, and reddish ventrally. Males are dark grayish, with dark gray stripe on upper third of body interrupted by white blotches along dorsal fin base, and red pigment on side and on underside of head. This species occurs in the western Atlantic from southern Florida and Bermuda to Venezuela, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs at the Dry Tortugas and Campeche Bank. It is associated with sponge beds and sea-grass beds to a depth of 7.5 m. Maximum known size is 43.2 mm SL.

references: Longley and Hildebrand 1941; Kanazawa 1952; Springer 1955b, 1959c; Caldwell and Caldwell 1964; Cervigón 1966; Birdsong and Emery 1968; Böhlke and Chaplin 1968; Greenfield and Johnson 1981; Robins and Ray 1986; Randall 1996; Smith 1997; Smith-Vaniz et al. 1999; Williams 2002c.

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LABRISOMIDAE 601

Malacoctenus triangulatus Saddled blenny

Springer, 1959

Malacoctenus triangulatus has nearly symmetrical dorsal and ventral profiles, maxilla covered posteriorly, and triangular saddle markings on side of body. Snout is slightly spatulate. Anterior naris is tubular and has two to four long, slender cirri on posterior margin. Supraorbital cirrus is long and multifid and has two to eight branches. Nuchal cirrus is oriented perpendicular to body axis, is multifid, and has 4 to 13 branches. Maxilla extends to anterior margin of orbit. Lips are fleshy, and upper lip has a continuous fold. Jaw teeth consist of a single row of canines. Teeth occur in vomer but not in palatine. Gill rakers on first arch number 11 to 14. Measurements are expressed as percent of SL: head length 26%–30%, snout length 7.3%–10.3%, upper jaw length 8.2%– 10.6%, pre-dorsal-fin length 25.1%–28.1%, pre-anal-fin length 46.1%– 49.2%, body depth at pelvic fin origin 23.3%–29.4%. Pectoral fin has 13 to 15 (usually 14) rays. Dorsal fin has 19 to 21 spines and 11 to 13 rays, with first spine usually more than 11% of SL and longer than second through fifth spines. Pelvic fin has 1 spine and 3 rays, with third ray no more than half the length of longest ray. Anal fin has 2 spines and 20 to 22 rays. Scales are present

on body, including predorsal and prepectoral areas. Breast is scaled in adult males. Lateral line scales number 52 to 61. Color is usually pale yellowish, with three to five dark brown or reddish triangular saddles on upper to midflank. Large spots on lower side are located midway between saddles. Saddles may be light or dark, uniform in color, or forming a reticulate pattern, or they may be absent. Fins of males are unpigmented, but fins of females are densely spotted. Color patterns vary over the geographic range of the species. This species occurs in the western Atlantic from southern Florida and the Bahamas to Brazil, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs at the Dry Tortugas, Veracruz, and Campeche Bank. It is associated with rocks and coral reefs in clear water to 14 m. Maximum known size is 48 mm SL.

references: Springer 1959b,c; Birdsong and Emery 1968; Böhlke and Chaplin 1968; Springer and Gomon 1975; Castro-Aguirre and MárquezEspinoza 1981; Greenfield and Johnson 1981; Robins and Ray 1986; Randall 1996; Smith 1997; Williams 2002c.

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602 PERCIFORMES

Nemaclinus atelestos Threadfin blenny

Böhlke and Springer, 1975

Nemaclinus atelestos has dorsal profile slightly more rounded than ventral profile, some pectoral fin rays elongated, and lateral line well developed only anteriorly. Snout is slightly spatulate. Anterior naris is tubular and lacks cirrus, and posterior naris is a simple pore. Supraorbital cirrus is about as long as eye, slender, and flattened. Nuchal cirrus is simple, located on side of nape, and very small. Maxilla extends to middle of eye. Lips are fleshy, and upper lip has a continuous fold. Upper jaw has an outer row of slightly compressed, pointed teeth and an inner row of one to five very small teeth. Lower jaw has a single row of large teeth. Vomer has four to seven moderately large teeth, and palatine lacks teeth. Measurements are expressed as percent of SL: head length 30%–35%, pectoral fin length 30%–35%, body depth 20%–22%; and as percent of head length: snout length 22%–30%, eye diameter 23%–30%, upper jaw length 36%– 42%. Pectoral fin has 11 or 12 rays, with central ray elongated and filamentous. Dorsal fin has 21 to 23 spines and 7 to 9 rays. Pelvic fin has

1 spine and 3 rays, with third ray vestigial and not visible externally. Anal fin has 2 spines and 18 or 19 rays, with first spine shorter than second spine in males. Body is completely covered with cycloid scales, including predorsal region, pectoral fin base, and area around vent. Head is naked. Lateral line consists of 8 to 10 pored scales behind head and occasionally pored scales along posterior section of trunk. Vertebrae number 33 to 35: 11 precaudal and 22 to 24 caudal. Color is brown, with an elongate black spot on anterior section of dorsal fin. This species occurs between 34 and 256 m in the western Atlantic from the northern Gulf of Mexico and Bermuda to Nicaragua, including the Bahamas and the Antilles. In the Gulf of Mexico it has been captured at the Flower Garden Banks and on the western side of the Yucatán Peninsula. Maximum known size is 28.6 mm SL.

references: Böhlke and Springer 1975; SmithVaniz et al. 1999; Williams 2002c.

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LABRISOMIDAE 603

Paraclinus cingulatus Coral blenny

(Evermann and Marsh, 1899)

Paraclinus cingulatus has nearly symmetrical dorsal and ventral profiles, a nuchal cirrus resembling a lappet, and a dorsal fin consisting only of spines. Snout is slightly acute and slightly projecting. Anterior naris is tubular and has a simple cirrus on posterior margin. Supraorbital cirrus consists of three or four fingerlike projections arising from a single base. Nuchal cirrus consists of several fingerlike projections and is oriented perpendicular to body axis. Both supraorbital and nuchal cirri are shorter than eye diameter. Maxilla extends to or beyond posterior margin of orbit (males) or to pupil (females). Lips are fleshy and have continuous folds. Jaw teeth are conical and arranged in bands that narrow toward corner of mouth. Small granular teeth occur in vomer. Measurements are expressed as percent of SL: head length 32.2%–33.7%, snout length 7.5%– 8.6%, eye diameter 6.4%–7.3%, upper jaw length 13.2%–20.5%, pre-dorsalfin length 26.3%–28.6%, pre-anal-fin length

57%– 60.2%, body depth 18.2%–20.1%. Pectoral fin has 11 or 12 (usually 12) rays. Dorsal fin has 27 or 28 spines, with first 3 spines connected to remainder of fin by low membrane. Pelvic fin has 1 spine and 2 rays. Anal fin has 2 spines and 15 or 16 rays. Lateral line scales number 27 to 31 (usually 29 or 30). Scales are absent at base of pectoral fin. Color is cream, with five dark chocolate bars along flanks. Bars may be fused dorsally and ventrally. Dorsal fin lacks ocellus. This species occurs in the western Atlantic from southern Florida and the Bahamas to Honduras, including the Florida Keys, the Gulf of Mexico, and the Greater Antilles. It is known from the Dry Tortugas in the Gulf of Mexico and is associated with coral rubble on reefs and in tide pools. Maximum known size is about 20 mm SL. references: Springer 1955b; Robins and Ray 1986; Smith 1997; Williams 2002c.

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604 PERCIFORMES

Paraclinus fasciatus Banded blenny

(Steindachner, 1876)

Paraclinus fasciatus has nearly symmetrical dorsal and ventral profiles, a nuchal cirrus resembling a lappet, and a dorsal fin consisting only of spines. Snout is slightly acute and slightly projecting. Anterior naris is tubular and has a simple cirrus on posterior margin. Supraorbital cirrus consists of one to four slender filaments arising from a single base. Nuchal cirrus is paddlelike and oriented perpendicular to body axis, with three short points on distal margin. Maxilla extends to or beyond posterior margin of orbit (males) or to pupil (females). Lips are fleshy and have continuous folds. Jaw teeth are conical and arranged in bands that narrow posteriorly. Small granular teeth occur in vomer. Measurements are expressed as percent of SL: head length 25.1%–30.2%, snout length 5.1%–7.7%, upper jaw length 10.5%–12.9%, pre-dorsalfin length 19.1%–25.3%, pre-anal-fin length 47%–54.6%, body depth 22.6%–25%. Pectoral fin has 12 to 14 (usually 13) rays. Dorsal

fin has 28 to 31 (usually 29) spines, with first 3 spines connected to remainder of fin by low membrane. Pelvic fin has 1 spine and 2 rays. Anal fin has 2 spines and 17 to 20 (usually 19) rays. Scales are present on base of pectoral fin. Lateral line scales number 32 to 38 (usually 35 or 36). Two or three scale rows occur at base of pectoral fin. Color varies from uniformly pale to dark, with darker bars along flanks. Zero to four ocelli are present on dorsal fin. Cirri are darkly pigmented, especially in males. This species occurs in the western Atlantic from southern Florida and the Bahamas to Venezuela, including the eastern and southern Gulf of Mexico and the Greater and Lesser Antilles. Maximum known size is 40.5 mm SL. references: Springer 1955b; Birdsong and Emery 1968; Böhlke and Chaplin 1968; Greenfield and Johnson 1981; Robins and Ray 1986; Randall 1996; Smith 1997; Williams 2002c.

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LABRISOMIDAE 605

Paraclinus marmoratus Marbled blenny

(Steindachner, 1876)

Paraclinus marmoratus has nearly symmetrical dorsal and ventral profiles, first several dorsal fin spines and membranes almost completely separated from remainder of fin, and ocelli located on posterior sections of dorsal and anal fins. Snout is very acute and projecting. Anterior naris is tubular and has a simple cirrus on posterior margin. Supraorbital cirrus consists of one to four slender filaments arising from a single base. Nuchal cirrus is flat, fringed distally, shorter than orbit diameter, and oriented perpendicular to body axis. Maxilla extends to near posterior margin of orbit. Lips are fleshy, upper lip has a continuous fold, and lower lip has a nearly continuous fold. Jaw teeth are small, conical, and arranged in a band that narrows toward corner of mouth. Small granular teeth occur in vomer. Opercular spine extends beyond base of third dorsal fin spine and ends in a single point. Measurements are expressed as percent of SL: head length 24.6%–32.1%, snout length 5.7%– 8.6%, upper jaw length 10.8%–16.8%, pre-dorsalfin length 17.4%–24.2%, pre-anal-fin length

45.8%–55.8%, body depth 21.6%–26.9%. Pectoral fin has 12 to 14 rays. Dorsal fin has 28 or 30 (usually 28) spines and 1 ray, with first 2 spines longer than remainder of spines and first 3 nearly free of remainder of fin. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 19 to 21 rays. Lateral line scales number 35 to 40. Base of pectoral fin has two to four scale rows. Color varies from light tan to dark brown, with about six irregular bands on flanks and continuing onto dorsal and anal fins. Membrane of anterior 3 dorsal fin spines often has narrow black and white bands. This species occurs in the western Atlantic from southern Florida and the Bahamas to Venezuela, including the eastern Gulf of Mexico off Florida and Cuba. It is an inquiline of sponges. Maximum known size is 63 mm SL. references: Springer 1955b; Dawson 1960; Springer and Woodburn 1960; Birdsong and Emery 1968; Böhlke and Chaplin 1968; Greenfield and Johnson 1981; Smith 1997; Williams 2002c.

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606 PERCIFORMES

Paraclinus nigripinnis (Steindachner, 1867) Blackfin blenny

Paraclinus nigripinnis has nearly symmetrical dorsal and ventral profiles, a minute opercular spine, and an ocellus located on posterior section of dorsal fin. Snout is acute and slightly projecting. Anterior naris is tubular and has a simple cirrus on posterior margin. Supraorbital cirrus consists of several fingerlike projections. Nuchal cirrus is flat, simple, or with several fingerlike projections; shorter than orbit diameter; and oriented perpendicular to body axis. Maxilla extends to near posterior margin of orbit. Lips are fleshy, upper lip has a continuous fold, and lower lip has a nearly continuous fold. Jaw teeth are small, conical, and arranged in bands that narrow toward corner of mouth. Small granular teeth occur in vomer. Opercular spine terminates as several triangular points. Measurements are expressed as percent of SL: head length 24.6%–36.6%, snout length 4.6%–7.3%, upper jaw length 9.8%–17.4%, pre-dorsal-fin length 21%– 24%, pre-anal-fin length 47.8%–59.3%, body depth 20.8%–25.7%. Pectoral fin has 12 to 14 rays. Dorsal fin has 29 to 31 spines and 0 or 1 ray, with first 2 spines slightly longer than re-

mainder of spines and first 3 more flexible than and nearly free of remainder of fin. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 17 or 18 rays. Lateral line scales number 31 to 35 (usually 33 to 35). Base of pectoral fin has a single row of scales. Color varies from light tan to dark brown. Specimens with intermediate levels of pigment have six or seven faint and irregular bands on flanks that continue onto dorsal and anal fins. A small ocellus is located on posterior aspect of dorsal fin, and a brown bar runs across base of caudal fin. This species occurs in the western Atlantic from southern Florida and Bermuda to Brazil, including the Bahamas, the Gulf of Mexico, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs in the Dry Tortugas. Maximum known size is 40.5 mm SL.

references: Longley and Hildebrand 1941; Springer 1955b; Caldwell 1963; Birdsong and Emery 1968; Böhlke and Chaplin 1968; Greenfield and Johnson 1981; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Williams 2002c.

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LABRISOMIDAE 607

Starksia lepicoelia Blackcheek blenny

Böhlke and Springer, 1961

Starksia lepicoelia has a simple cirrus on anterior naris, on eyeball, and on nape, and ventral side of body is covered with scales. Dorsal profile of head is gently rounded. Snout is slightly acute. Anterior naris is tubular, with cirrus on posterior margin extending one-third to one-half the length of eye diameter. Posterior naris is a simple pore. Supraorbital cirrus is equal to or longer than nasal cirrus. Nuchal cirrus is shorter than nasal and supraocular cirri. Maxilla extends beyond posterior margin of orbit. Lips are fleshy, upper lip has a continuous fold, and lower lip has a nearly complete fold. Jaws have a single row of canines and a patch of small teeth behind anterior canines. Teeth occur in vomer and palatine. Operculum lacks spine. Measurements are expressed as percent of SL: head length 31.9%–35.8%, snout length 6.7%–7.3%, upper jaw length 15.6%–18.3%, pre-dorsal-fin length 27.1%– 30%, pre-anal-fin length 56.4%– 60.2%, body depth at pelvic fin origin 19.9%–23.5%. Pectoral fin has 13 or 14 rays. Dorsal fin has 19 to 21 spines and 7 to 9 (usually 8) rays. Pelvic fin has 1 spine and 3 rays, but third ray is a mi-

nute basal element and generally is not evident. Anal fin has 2 spines and 16 to 19 (usually 17) rays. In males, genital papilla is free of first anal fin spine, and first anal fin spine is free of second spine. Caudal fin has 12 or 13 segmented rays. Scales are present on body, with exception of head. Lateral line is discontinuous and has 14 to 17 (usually 15 or 16) scales in anterior section and 19 to 22 in posterior section. Color is usually uniformly reddish brown, but occasionally ground color is interrupted with narrow light bands. Mature males have a dark blotch on cheek. This species occurs in the western Atlantic from the Bahamas to Nicaragua, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from the northern and northwestern coasts of Yucatán and on Alacrán Reef. It is associated with coral reefs between 8 and 20 m. Maximum known size is 29.2 mm SL.

references: Böhlke and Springer 1961; Smith 1997; Williams 2002c.

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608 PERCIFORMES

Starksia ocellata (Steindachner, 1876) Checkered blenny

Starksia ocellata has a long, simple cirrus on naris, on eyeball, and on nape, and ventral side of body is largely naked. Dorsal profile of head is gently rounded. Snout is slightly acute. Anterior naris is tubular and has a simple, slender cirrus on posterior margin. Supraorbital cirrus is slender and longer than other cirri. Nuchal cirrus is slender. Maxilla extends to or beyond posterior margin of orbit. Lips are fleshy, upper lip has a continuous fold, and lower lip has a nearly complete fold. Jaws have a single row of canines and a patch of small teeth behind anterior canines. Teeth occur in vomer and are present or absent in palatine. Measurements are expressed as percent of SL: head length 28%–31.4%, snout length 6.2%–7.2%, upper jaw length 13.3%–14.4%, pre-dorsalfin length 26.1%–27.9%, pre-anal-fin length 48.3%–54%, body depth at pelvic fin origin 17.8%–20.7%. Pectoral fin has 12 to 15 (usually 14) rays. Dorsal fin has 20 to 22 spines and 7 to 9 (usually 8 or 9) rays. Pelvic fin has 1 spine and 3 rays, but third ray is a minute basal element and generally not evident. Anal fin has 2 spines and 16 to 19 (usually 18 or 19) rays. In males, first anal spine is completely

separated from second spine and rays, is longer than second spine, and is connected to elongate genital papilla. Caudal fin has 12 or 13 segmented rays. Lateral line is discontinuous and has 15 to 19 scales in anterior section and 20 to 23 in posterior section. Scales are present on body, with exception of belly, and are absent on head. Color varies from plain brown to light, with three distinct rows of irregularly shaped blotches. Cheek, operculum, and pectoral fin base are usually covered with small ringlike markings. This species occurs in the western Atlantic from North Carolina and the Bahamas to the Gulf of Mexico. In the Gulf of Mexico it occurs along the west coast of Florida and Veracruz. It is associated with rock and coral habitats between 1 and 24 m. Occasionally it has been found as an inquiline in sponges. Maximum known size is 34 mm SL. references: Longley and Hildebrand 1941; Dawson 1960; Springer and Woodburn 1960; Böhlke and Springer 1961; Böhlke and Chaplin 1968; Greenfield 1979; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Williams 2002c.

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LABRISOMIDAE 609

Starksia starcki Gilbert, 1971 Key blenny Starksia starcki has a simple cirrus on anterior naris, on eyeball, and on nape, and ventral side of body is covered with scales. Dorsal profile of head is gently rounded. Snout is slightly acute. Anterior naris is tubular, with a cirrus on posterior margin. Posterior naris is a simple pore. Maxilla extends beyond posterior margin of orbit. Lips are fleshy, upper lip has a continuous fold, and lower lip has a nearly complete fold. Jaws have a single row of canines and a patch of small teeth behind anterior canines. Teeth occur in vomer and palatine. Operculum lacks spine. Measurements are expressed as percent of SL: head length 29.9%–32.2%, snout length 6.6%–7.4%, upper jaw length 12.3%–14.7%, pre-dorsal-fin length 25.9%– 28.1%, pre-anal-fin length 49.6%–53.1%, body depth at pelvic fin origin 17%–18.5%. Pectoral fin has 13 rays. Dorsal fin has 20 or 21 spines and 8 or 9 (usually 9) rays. Pelvic fin has 1 spine and 3 rays, but third ray is a minute basal element and generally not evident. Anal

fin has 2 spines and 18 or 19 (usually 19) rays. In males, genital papilla is free from first anal fin spine, and first anal spine is partially free of second spine. Caudal fin has 12 or 13 segmented rays. Lateral line is discontinuous and has 13 or 14 scales in anterior section and 22 or 23 in posterior section. Scales are present on body, with exception of head. Color is tan, with eight or nine irregular or broken, widely spaced dark brown bars, the first of which circles nape. Stripe extends from third band to caudal fin. This species occurs at less than 8 m in the western Atlantic from the Florida Keys and Honduras. It is associated with surge channels on reefs. It has not been recorded from the Gulf of Mexico, but it may occur there. Maximum known size is 27.3 mm SL.

references: Gilbert 1971a; Robins and Ray 1986; Smith 1997; Williams 2002c.

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610

CHAENOPSIDAE

Tube blennies

Chaenopsids are elongate, moderately slender, and moderately compressed, with or without cirri on head, with maxilla not visible externally, and without lateral line. Lateral profile of head is slightly to strongly elevated. Eye is large and located near dorsal margin of head. Nostril is paired, with anterior naris tubelike and often possessing a cirrus on posterior rim, and posterior naris possessing a raised rim. Supraorbital cirrus is usually present, nuchal cirrus is often present, and cirri vary considerably in complexity. Some species bear multiple spines on roofing bones of cranium and bones surrounding eyes. Mouth is terminal, small to moderate in size, and horizontal to slightly oblique. Lips are fleshy and have free margins. Jaw teeth are caninelike or incisor-like, variable in size, and arranged in one to several rows. Teeth are usually present in vomer and palatine. First gill arch has gill rakers. Gill membranes are connected beneath head. Pectoral fin is minute to well developed, with a narrow to broad, nearly vertical base and 4 to 15 rays. Dorsal fin is long and continuous and has 17 to 33 spines and 10 to 34 rays. Dorsal fin spines are usually more numerous than rays. Pelvic fin is jugular in location and consists of 1 embedded spine and 2 or 3 rays. Anal fin is long and consists of 2 flexible spines and 18 to 38 rays. All fin rays are unbranched. Caudal fin is truncate to convex. Scales are entirely lacking in all but one species. Maximum known size is less than 150 mm TL. Tube blennies occur in the western Atlantic, eastern Pacific, and western Pacific in tropical and subtropical seas. They are benthic and usually associated with rocky and coral reefs and rubble areas around reefs. Many species occupy tests of invertebrates. Males generally are more brightly colored and have a much larger dorsal fin than females. There are about 56 species in 11 genera, 40 species in 10 genera in the western Central Atlantic, and 11 or 12 species in 6 genera in the Gulf of Mexico. references: Matarese et al. 1984; Hastings and Springer 1994; Nelson 1994; Smith 1997; Smith-Vaniz et al. 1999; Williams 2002d.

Key to the Species of the Gulf of Mexico (Adapted from Stephens 1963, 1970; Williams 2002d) 1a. Dorsal fin elements consisting entirely of spines . . . . . . . . 2 1b. Dorsal fin elements consisting of spines and some segmented elements (rays) . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2a. Preopercular cirrus present (on ventralmost or second most ventral preopercular sensory pore); nuchal cirrus present; three infraorbital pores. . . . . . . Stathmonotus stahli p. 623 2b. Preopercular cirrus absent; nuchal cirrus absent; two infraorbital pores . . . . . . . . . . . Stathmonotus hemphillii p. 622

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CHAENOPSIDAE 611

3a. Total dorsal fin elements 44 to 56 . . . . . . . . . . . . . . . . . . 4 3b. Total dorsal fin elements 29 to 40 . . . . . . . . . . . . . . . . . . 6 4a. Dorsal fin elements (both spines and rays) 44 or 45 . . . . . . . . . . . . . . . . . . . . . . . . . Chaenopsis roseola p. 615 4b. Dorsal fin elements 51 or more . . . . . . . . . . . . . . . . . . . . 5 5a. Corona sensory pore (pore along midline of head near posterior margin of orbits) in line with or slightly posterior to nearest supraorbital pore . . . . . Chaenopsis ocellata p. 614 5b. Corona sensory pore (pore along midline of head near posterior margin of orbits) slightly anterior to nearest supraorbital pore. . . . . . . . . . . . . . Chaenopsis limbaughi p. 613 6a. Palatine teeth in two or more rows; top of head spiny . . . . . . . . . . . . . . . . . . . . Acanthemblemaria aspera p. 612 6b. Palatine with a single tooth row; top of head smooth . . . . 7 7a. Tip of lower jaw projecting beyond tip of upper jaw; broad stripe or series of dark blotches extending from eye to caudal fin base usually present; no supraorbital cirrus . . . . . . . . . . . . . . . . . . . . Hemiemblemaria simulus p. 621 7b. Tip of lower jaw not projecting beyond tip of upper jaw; no stripe or series of dark blotches on head and body; supraorbital cirrus present or absent . . . . . . . . . . . . . . . . 8 8a. Supraorbital cirrus longer than eye diameter in males and often in females; segmented dorsal fin rays 13 to 17. . . . . 9 8b. Supraorbital cirrus, when present, shorter than eye diameter; segmented dorsal fin rays 10 to 13 (rarely 14 in one species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 9a. Third pelvic ray vestigial . . . . . Emblemaria piratula p. 618 9b. Third pelvic ray one-fourth to one-half the length of longest pelvic ray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10a. Supraorbital cirrus banded; cirruslike papilla (one or two) on upper posterior section of orbit; pectoral rays usually 14 . . . . . . . . . . . . . . . . . . . . . . . Emblemaria atlantica p. 616 10b. Supraorbital cirrus not banded; no cirruslike papilla on upper posterior section of orbit; pectoral rays usually 13 . . . . . . . . . . . . . . . . . . . . . . Emblemaria pandionis p. 617 11a. First dorsal fin spine slightly longer than third; six (and one commissural) pores on frontal bone . . . . . . . . . . . . . . . . . . . . Emblemariopsis diaphana p. 620 11b. First dorsal fin spine slightly shorter than second and third; three (and one commissural) pore on frontal bone . . . . . . . . . . . . . . . . . . Emblemariopsis bahamensis p. 619

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612 PERCIFORMES

Acanthemblemaria aspera Roughhead blenny

(Longley, 1927)

Acanthemblemaria aspera has a spiny head and a branched supraorbital cirrus. Anterior naris has a multifid cirrus on posterior margin. Supraorbital cirrus is deeply branched. Cranial spines occur midway between posterior margin of orbit and origin of dorsal fin. Nasal bone has knobby spines. Frontal bone has 3 to 5 spines directed anterodorsally, with those on each side separated by narrow groove. Spines on posterior dorsal rim of orbit are separated from spines on frontal bone by narrow groove. Upper jaw has an outer row of 11 to 15 conical teeth and an inner band of small, conical teeth. Lower jaw has a single outer row of 17 to 22 conical teeth and an inner band of small, conical teeth. Vomer has 8 to 10 teeth arranged in a ring or a crescentic row. Palatine has two rows of teeth. Gill rakers on first arch number 10 or 11. Pectoral fin has 12 to 14 rays. Dorsal fin has 19 to 22 spines and 14 to 17 rays. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 22 to 25 rays. Segmented caudal rays number 11 to 14 (usually 12). Vertebrae

number 40 to 43: 12 or 13 precaudal and 28 to 31 caudal. Color is variable and differs in males and females. Males are dark anteriorly and have an ocellated black spot on anterior section of dorsal fin, unless this portion of fin is uniformly black. Females are light and lack ocellated spot, but have three broken lines across lower surface of head. This species occurs in the western Atlantic from southern Florida to Panama, including the Gulf of Mexico, the Florida Keys, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it has been captured at the Dry Tortugas and along the west coast of the Yucatán Peninsula. It is associated with coral reefs. Maximum known size is 40 cm TL. references: Longley 1927 (as Paremblemaria aspera); Longley and Hildebrand 1941; Stephens 1963; Böhlke and Chaplin 1968; Smith-Vaniz and Palacio 1974; Robins and Ray 1986; Smith 1997; Williams 2002d.

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CHAENOPSIDAE 613

Chaenopsis limbaughi Yellowface pikeblenny

Robins and Randall, 1965

Chaenopsis limbaughi has a U-shaped snout and a dorsal fin with more rays than spines. Head is low in profile and does not ascend appreciably to origin of dorsal fin. Snout is long and very acute. Anterior naris is a long tube lacking a cirrus. Lower jaw projects slightly beyond upper jaw, and upper jaw extends to eye in juveniles but beyond eye in adults. Labial fold of upper lip is not continuous across symphysis. Jaw teeth are arranged in bands that are broad near symphysis but narrow toward corner of mouth, with those in outer row enlarged. Vomer and palatine have tooth patches, and tip of tongue extends to vomerine tooth patch. Lower jaw has poorly developed medial dewlap. Medial dewlap on anterior section of throat is well developed in mature males. Measurements are expressed as percent of SL: head length 25%–30%, snout length 5.9%–9.6%, eye diameter 3.1%–5.8%, snout to dorsal fin origin 19%–26%, snout to anal fin origin 47%–56%, body depth at dorsal fin origin 5.4%– 8.2%. Pectoral fin has 12 or 13 rays, with lower rays progressively shorter than median rays. Dorsal fin has 18 to 21 spines and 31 to 36 rays. Spinous section of

dorsal fin is elevated in males. Pelvic fin has 1 spine and 3 rays, with first and second rays elongated and third short. Anal fin has 2 spines and 33 to 37 rays. Membranes of last dorsal and anal fin rays are continuous with caudal fin. Color is pale tan, with a yellow cast anteriorly, pale spots, and a narrow midlateral stripe. Midlateral band may be broken into blotches. Mature males have a pink eye, a lemon yellow anterior head, black branchiostegal membranes, an ocellus on anterior section of dorsal fin, and a yellow blotch above ocellus. This species occurs between 3 and 21 m in the western Atlantic from the Florida Keys, the Bahamas, and throughout the Caribbean Sea. It has not been reported from the Gulf of Mexico, but it may occur there. It lives in holes in coral reefs and in coral rubble and sand in the vicinity of reefs. Food consists of small polychaetes, crustaceans, and ray-finned fishes. Maximum known size is 68 mm SL.

references: Robins and Randall 1965; Robins and Ray 1986; Smith 1997; Williams 2002d.

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614 PERCIFORMES

Chaenopsis ocellata Poey, 1865 Bluethroat pikeblenny

Chaenopsis ocellata has a bluntly V-shaped snout and a dorsal fin with more rays than spines. Head is slightly elevated in profile and ascends to origin of dorsal fin. Snout is long and very acute. Anterior naris is a long tube lacking a cirrus. Lower jaw projects slightly beyond upper jaw, and upper jaw extends to eye in juveniles but beyond eye in adults. Labial fold of upper lip is not continuous across symphysis. Jaw teeth are arranged in bands that are broad near symphysis but narrow toward corner of mouth, with those in outer row enlarged. Vomer and palatine have tooth patches, and tip of tongue extends to vomerine tooth patch. Lower jaw has poorly developed medial dewlap. Measurements are expressed as percent of SL: head length 26%–28%, snout length 6.3%–7.8%, eye diameter 2.7%– 4.2%, snout to dorsal fin origin 19%–22%, snout to anal fin origin 48%–55%, and body depth at dorsal fin origin 5.4%–7.7%. Pectoral fin has 12 to 14 rays, with lower rays progressively shorter than median rays. Dorsal fin has 18 to 20 spines and 33 to 36 (usually 34 or 35) rays. Pelvic fin has 1 spine and 3 rays, with first and

second rays elongated and third short. Anal fin has 2 spines and 34 to 37 rays. Membranes of last dorsal and anal fin rays are continuous with caudal fin. Color is sandy brown, with dark bars on side and dark branchiostegal membranes. Mature males have a dusky head, with blue branchiostegal membranes, an orange iris, and a curved black marking enclosing an orange spot between first and second dorsal fin spines. This species occurs in the western Atlantic from southern Florida and the Bahamas to Cuba and the Virgin Islands, including the Gulf of Mexico. In the Gulf of Mexico it occurs at the Dry Tortugas and from Tampa to Cedar Key, FL. It is associated with rocky reefs and silty grass beds and lives solitarily in polychaete tubes from near shore to 12 m. Maximum known size is 111 mm SL. references: Longley and Hildebrand 1941; Böhlke 1957b; Springer and Woodburn 1960; Robins and Randall 1965; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997; Williams 2002d.

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CHAENOPSIDAE 615

Chaenopsis roseola Flecked pikeblenny

Hastings and Shipp, 1981

Chaenopsis roseola has an evenly rounded lower jaw, a dorsal fin with more rays than spines, and black blotches on flank. Head is slightly elevated and slightly convex in profile. Snout is long and very acute. Anterior naris is a tube lacking a cirrus, and posterior naris has a raised rim. Lower jaw projects slightly beyond upper jaw, and upper jaw extends to eye in juveniles but beyond eye in adults. Labial fold of upper lip is not continuous across symphysis. Jaw teeth are arranged in a narrow band, with those in outer row enlarged and decreasing in size toward corner of mouth. Vomer has several small teeth, and palatine has a single row of teeth. Tip of tongue extends beyond vomerine teeth. Lower jaw has a medial dewlap in throat region that extends to anterior margin of orbit. Measurements are expressed as percent of SL: head length 22.5%–25.8%, snout length 4.5%– 6%, eye diameter 3.9%–5%, snout to dorsal fin origin 19.2%–21.8%, snout to anal fin origin 43%–50%, body depth at dorsal fin origin 8%–10%. Pectoral fin has 12 to 14 rays, with lower rays progressively

shorter than median rays. Dorsal fin is low in both sexes and has 17 or 18 spines and 26 to 28 rays. Pelvic fin has 1 spine and 3 rays, with first and second rays elongated and third short. Anal fin has 2 close-set spines and 29 to 30 rays. Membranes of last dorsal and anal fin rays are continuous with caudal fin. Vertebrae number 48 or 49. Body is straw colored, with eight black blotches along flank and small pink flecks scattered on side of head, on dorsum, and along pectoral and anal fin bases. Mature males have a concentration of melanophores on membranes between first 4 dorsal fin spines. This species occurs between 33 and 64 m in the western Atlantic from North Carolina and the eastern Gulf of Mexico. In the Gulf of Mexico it occurs from Tampa Bay to De Soto Canyon. It is associated with rubble and shell fragments. Maximum known size is 43 mm SL. references: Springer and Woodburn 1960 (as Chaenopsis sp.?); Hastings and Shipp 1981; Robins and Ray 1986; Boschung 1992; Smith 1997; Williams 2002d.

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616 PERCIFORMES

Emblemaria atlantica Banner blenny

Jordan and Evermann, 1898

Emblemaria atlantica has a single supraorbital cirrus, a cirruslike papilla on eye, and bony ridges on snout. Head is elevated in profile and smoothly rounded to dorsal fin origin. Snout is short and slightly blunt. Groove extends from interorbital region to dorsal fin and is bordered by fleshy folds. Cirruslike papilla is located anterior to origin of folds. Anterior naris bears a simple flat cirrus about half of eye diameter. Supraorbital cirrus is simple, slender, and 3 times the length of eye diameter in males. Maxilla extends half of orbital length beyond eye. Jaw has an outer row of slender, recurved teeth and an inner band of small teeth. Vomer possesses several minute teeth, and palatine has a single row of blunt teeth. Measurements are expressed as percent of SL: head length 27.8%–30.9%, snout length 7%–7.8%, eye diameter 8.2%– 8.3%, upper jaw length 17.5%– 17.7%, pre-dorsal-fin length 20.9%–22.5%, pre-anal-fin length 47.1%– 47.4%. Pectoral fin is rounded to slightly pointed and has 13 or 14 (usually 14) rays. Dorsal fin originates closer to posterior rim of orbit than to posterior margin of operculum and has 21 or 22 spines and 14 to 16 (usually 14) rays, with spinous section of fin high in males and low in females. Pelvic

fin has 1 spine and 3 rays, with third ray onefourth to one-half the length of second-longest ray. Anal fin has 2 spines and 21 to 23 rays. Dorsal and anal fins are free of caudal fin. Caudal fin is broadly rounded and has 13 branched rays. Males have longer supraorbital cirrus and longer jaws than females. Color in preservative is dusky gray to light brown, and head is covered with small brown spots and ringlike mottling. Nasal and supraorbital cirri are banded. Females have six bars on flank that are outlined by narrow brown lines. This species occurs in the western Atlantic from Georgia and Bermuda and the northern Gulf of Mexico. In the Gulf of Mexico it is known from the west coast of Florida and off Texas. It is absent from southern Florida. It is associated with rocky reefs. Males apparently use their elongated dorsal fins to attract females. Maximum known size is 75 mm TL. references: Böhlke 1957a; Springer and Woodburn 1960; Stephens 1963, 1970; Robins and Ray 1986; Dennis and Bright 1988a; Smith 1997; Hoese and Moore 1998; Smith-Vaniz et al. 1999; Williams 2002d.

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CHAENOPSIDAE 617

Emblemaria pandionis Sailfin blenny

Evermann and Marsh, 1900

Emblemaria pandionis has a single supraorbital cirrus above eye and bony ridges on snout. Head is steep in profile and abruptly rounded above eye, and snout is short and blunt. Groove extends from interorbital region to dorsal fin and is bordered by fleshy folds. Anterior naris has a multifid flat cirrus on posterior margin that is about one-third of eye length. Supraorbital cirrus is trifid at tip and slender. Maxilla extends two-thirds of orbital length beyond eye. Jaw has an outer row of slender, recurved teeth and an inner band of small teeth. Vomer possesses several minute teeth, and palatine has a single row of blunt teeth. Measurements are expressed as percent of SL: head length 25%– 27%, snout length 5.2%–5.9%, eye diameter 6.6%–7%, upper jaw length 13.4%–14.6%, pre-dorsal-fin length 18.3%–19.5%, pre-analfin length 44.3%– 45.1%, and body depth 18%–21.4%. Pectoral fin is rounded and has 13 rays. Dorsal fin originates closer to posterior rim of orbit than to posterior margin of operculum and has 21 spines and 16 rays, with spinous section of fin high in males and low in females. Pelvic fin has 1 spine and 3 rays, with third ray one-fourth to one-half the length of second-longest ray. Anal fin has 2 spines and 23 or 24 rays. Dorsal and anal fins are separated from caudal fin. Caudal fin is broadly rounded and has 13 rays. Males have longer or-

bital cirri than females. Color of males in preservative is tan on head and uniformly brown on remainder of body, with 9 or 10 deep brown spots on flank and small brown dots on upper flank and dorsal fin base. Supraorbital cirrus is not banded. Females are light tan, with about 13 brown markings along midline and 13 or 14 large ringlike markings along ventral midline. This species occurs usually in shallow water in the western Atlantic from southern Florida and the Bahamas to Venezuela, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs from Texas to Florida, including the Dry Tortugas and along the coast of Veracruz. Males maintain territories in coral rubble, and throughout the day they rapidly raise and lower their elongated dorsal and anal fins, apparently to attract females. This species is associated with strong currents and lives in polychaete tubes or clam holes in sandy substrates with coral rubble. Maximum known size is 45.2 mm SL.

references: Longley and Hildebrand 1941; Böhlke 1957a; Stephens 1963, 1970; Böhlke and Chaplin 1968; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Greenfield and Johnson 1981; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Smith et al. 1998; Williams 2002d.

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618 PERCIFORMES

Emblemaria piratula Pirate blenny

Ginsburg and Reid, 1942

Emblemaria piratula has a single supraorbital cirrus above eye and bony ridges on snout. Head is moderately elevated in profile and moderately rounded to dorsal fin origin. Snout is short and acute. Groove extends from interorbital region to dorsal fin. Anterior naris has a simple cirrus about one-third the length of orbit. Supraorbital cirrus is simple and fleshy, about twice as long as orbit, and usually flattened in males. Maxilla extends slightly beyond orbit. Jaws have an outer row of slender, recurved teeth and an inner band of small teeth. Vomer possesses several minute teeth, and palatine has a single row of teeth. Measurements are expressed as percent of SL: head length 25.6%–27.4%, snout length 4.9%–5.6%, eye length 4.8%– 6.1%, upper jaw length 11.2%– 14.9%, pre-dorsal-fin length 20%–24%, preanal-fin length 48.1%–52%, body depth 13.4%–15.6%. Pectoral fin is rounded and has 12 or 13 rays. Dorsal fin originates closer to posterior margin of operculum than to posterior rim of orbit and has 17 to 20 spines and

13 to 15 (usually 15) rays, with spinous section of fin high in males and low in females. Pelvic fin has 1 spine and 3 rays, but third ray is vestigial. Anal fin has 2 spines and 20 or 21 rays. Dorsal fin is free of caudal fin, but anal fin is connected to caudal fin by membrane. Caudal fin is broadly rounded and has 13 branched rays. Males are blackish anteriorly, grading to ashy gray posteriorly, with basal half of anterior dorsal fin spines orange. Supraorbital cirrus is unpigmented. Females are mottled brown anteriorly, grading to pale posteriorly. This species occurs in the western Atlantic from North Carolina to the Gulf of Mexico. In the Gulf of Mexico it is known from the Dry Tortugas, along the west coast of Florida, and off Alabama. Maximum known size is 19.8 mm SL. references: Ginsburg 1942; Stephens 1963, 1970; Williams and Shipp 1980; Hastings and Shipp 1981; Robins and Ray 1986; Boschung 1992; Smith 1997; Williams 2002d.

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CHAENOPSIDAE 619

Emblemariopsis bahamensis Blackhead blenny

Stephens, 1961

Emblemariopsis bahamensis has bony ridges on snout and lacks a cirrus above eye. Head in profile is smoothly curved to origin of dorsal fin and rounded in dorsal view. Snout is spatulate. Anterior naris has a simple, short cirrus that may be rudimentary. Nasal ridges are smooth. Maxilla extends slightly beyond orbit. Jaws have an outer row of small, slightly recurved teeth and an inner band of small teeth. Vomer possesses about 10 to 12 minute teeth, and palatine has a single row of teeth. Measurements are expressed as percent of SL: head length 24.1%–30%, snout length 6.7%– 8.3%, orbit diameter 6.7%–10.9%, upper jaw length 11.7%–15%, pre-dorsal-fin length 17.5%–23.8%, pre-anal-fin length 42.9%– 47.6%, head depth 11.3%–15.3%. Pectoral fin is acutely rounded and has 13 rays. Dorsal fin originates closer to posterior margin of operculum than to posterior rim of orbit and has 20 to 21 spines and 12 or 13 rays. First dorsal fin spine is shorter than second, and first dor-

sal fin ray is more than twice as long as last dorsal fin spine. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 21 rays. Dorsal fin is free of caudal fin, but anal fin is connected to caudal fin by membrane. Caudal fin is broadly rounded and has 13 rays. Pores on frontal bone number three, in addition to commissural pore. Males are black anteriorly and light gray posteriorly, with spinous section of dorsal fin black. Females are translucent. This species occurs in the western Atlantic in the Florida Keys and from the Bahamas to northern South America, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from the Dry Tortugas and Veracruz. Maximum known size is 24.5 mm SL. references: Stephens 1963, 1970; Böhlke and Chaplin 1968; Robins and Ray 1986 (as Coralliozetus bahamensis); Smith 1997; Williams 2002d.

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620 PERCIFORMES

Emblemariopsis diaphana Longley, 1927 Glass blenny Emblemariopsis diaphana has bony ridges on snout and lacks a cirrus above eye. Head in profile is smoothly curved dorsally to origin of dorsal fin and triangular in dorsal view. Snout is blunt. Anterior naris has a simple, short cirrus. Nasal ridges are slightly rugose. Maxilla extends to postorbital margin. Jaws have an outer row of small, slightly recurved teeth and an inner band of small teeth. Vomer possesses about 10 to 12 minute teeth, and palatine has a single row of teeth. Measurements are expressed as percent of SL: head length 25.7%– 29.2%, snout length 3.9%–5.1%, orbit diameter 6.2%–7.8%, upper jaw length 10.7%– 12.8%, pre-dorsal-fin length 16%–16.9%, pre-anal-fin length 44.5%– 47.1%, head depth 15.1%–15.9%. Pectoral fin is acutely rounded and has 13 rays. Dorsal fin originates closer to posterior margin of operculum than to posterior rim of orbit and has 20 to 21 spines and 13 or 14 rays. First, and often second, dorsal fin spine is longer than remainder of spines, and first dorsal fin ray is more than twice as long as

last dorsal fin spine. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 22 rays. Dorsal and anal fins are free from caudal fin. Caudal fin is broadly rounded and has 13 branched rays. Pores on frontal bone number six, in addition to commissural pore. In life, males have a black head when in their burrows but resemble females when swimming. Females have a translucent brownish olive head and pectoral fin base, with minute white spots forming white lines. Dorsal fin is pigmented between first several spines, but remainder of fin and body are unpigmented. This species occurs in the western Atlantic in the Florida Keys and the Dry Tortugas in the Gulf of Mexico. Maximum known size is 25.4 mm SL. references: Longley 1927; Longley and Hildebrand 1941; Stephens 1963, 1970; Robins and Ray 1986 (as Coralliozetus diaphanus); Smith 1997; Williams 2002d.

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CHAENOPSIDAE 621

Hemiemblemaria simulus Wrasse blenny

Longley and Hildebrand, 1940

Hemiemblemaria simulus has a protruding lower jaw, no cirrus above eye, and a broad stripe on flank. Head in profile is gently curved from midsnout to dorsal fin origin and pointed in dorsal view. Snout is acute and depressed. Anterior naris has a simple, short cirrus, and posterior naris has a raised rim. Nasal ridges are smooth. Maxilla extends slightly beyond postorbital margin. Jaws have an outer row of enlarged, recurved teeth and an inner band of small teeth. Vomer possesses crescentic patch of minute teeth, and palatine has a single row of teeth. Measurements are expressed as percent of SL: head length 27.2%–33.5%, snout length 5.1%– 6.2%, orbit diameter 5.1%–10%, upper jaw length 13.1%–14.6%, pre-dorsalfin length 19.4%–21.5%, pre-anal-fin length 49.3%–56.1%, head depth 14.9%–15.3%. Pectoral fin is acutely rounded and has 14 rays. Dorsal fin originates closer to posterior margin of operculum than to posterior rim of orbit and has 21 to 23 spines and 17 rays. Spinous section of dorsal fin is about even in height and lower than rayed section of fin. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 22 or 23 rays. Dorsal and anal fins are free of

caudal fin. Caudal fin is emarginate to rounded. Adults have three color phases: yellowish or greenish yellow with a broad dark stripe, light ground color with a stripe broken into a number of blocklike segments, or stripe absent and yellow dorsal coloration grading into white on lower body. Juveniles are translucent with a black stripe. This species occurs in the western Atlantic in southern Florida and from the Bahamas to Honduras, including the eastern Gulf of Mexico, the Florida Keys, and Cuba. In the Gulf of Mexico it is known only from the Dry Tortugas. It is associated with coral reefs and lives in burrows of dead boring mollusks, with only its head protruding. This species closely resembles juveniles, females, and young males of Thalassoma bifasciatum. It swims freely with this species, and this association enables it to approach small ray-finned fishes on which it feeds. Maximum known size is 88.7 mm SL.

references: Stephens 1963, 1970; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997; Williams 2002d.

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622 PERCIFORMES

Stathmonotus hemphillii Blackbelly blenny

Bean, 1885

Stathmonotus hemphillii has a median notch in upper lip, a dorsal fin supported entirely by spines, and a greatly reduced pectoral fin. Head in profile is weakly convex to dorsal fin origin, and snout is acute. Anterior naris is a short tube, and posterior naris has a raised rim. Maxilla extends to or beyond posterior margin of orbit. Jaws have an outer row of canine teeth and an inner band of small teeth on either side of symphyses. Vomer has teeth, but palatine lacks teeth. Measurements are expressed as percent of SL: head length 15%–15.9%, snout length 2.4%–2.8%, eye diameter 2.1%– 3.3%, upper jaw length 6%–7.1%, predorsal-fin length 16.1%–19.1%, pre-anal-fin length 49.1%–53%, body depth 7.6%– 8.8%. Pectoral fin is very small and has 4 or 5 weak rays. Dorsal fin originates over pectoral fin base and has 45 to 53 spines. Pelvic fin has 1 spine and 2 rays. Anal fin has 2 widely separated, stout spines and 23 to 29 rays. Dorsal and anal fins are joined to caudal fin by membranes. Caudal fin is slightly emarginate. Two infraorbital pores are located between eyes.

Vertebrae number 50 to 58: 20 to 25 precaudal and 30 to 34 caudal. Color is sexually dimorphic and variable in males. Some males are colorless or variously pigmented on head and abdomen; other males are uniformly darkly pigmented except for margin of dorsal and caudal fins. Females lack pigment except for network of four to six dark lines and spots radiating from eyes. This species occurs in the western Atlantic from southern Florida and the Bahamas to Nicaragua, including the Florida Keys, the Dry Tortugas in the Gulf of Mexico, and the Greater and Lesser Antilles. In the Lesser Antilles it is known only from Antigua. It is associated with rock and rubble near coral reefs and sea-grass beds from the shoreline to 6 m. Maximum known size is 45 mm SL. references: Longley and Hildebrand 1941; Springer 1955a; Böhlke and Chaplin 1968; Robins and Ray 1986; Hastings and Springer 1994; Smith 1997; Williams 2002d.

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CHAENOPSIDAE 623

Stathmonotus stahli Eelgrass blenny

(Evermann and Marsh, 1899)

Stathmonotus stahli has a median notch in upper lip, a preopercular cirrus, and a dorsal fin supported entirely by spines. Head in profile is weakly convex to dorsal fin origin, and snout is acute. Anterior naris is a short tube, and posterior naris has a raised rim. Maxilla extends to or beyond posterior margin of orbit. Jaws have an outer row of canine teeth and an inner band of small teeth on either side of symphyses. Vomer has teeth, but palatine lacks teeth. Supraorbital cirrus is rounded and flaplike, and nasal and nuchal cirri are present. Measurements are expressed as percent of SL: head length 21.2%–22.5%, snout length 3.5%– 4.6%, eye diameter 3.7%–5.1%, upper jaw length 10%–11.6%, pre-dorsal-fin length 24.1%–25%, pre-anal-fin length 49.9%–51%, body depth 12.3%–14%. Pectoral fin is small and has 8 or 9 rays. Dorsal fin originates over opercular margin and has 39 to 45 spines. Pelvic fin inserts anterior to pectoral fin base and has 1 spine and 2 rays. Anal fin has 2 widely separated, stout spines and 21 to 26 rays. Dorsal and anal fins are joined to caudal fin by membranes. Caudal fin has 10 to 13 segmented rays and is slightly convex. Scales are present

on body. Vertebrae number 44 to 49: 16 to 19 precaudal and 27 to 31 caudal. Most specimens lack pigment, but some males have an irregular pattern of melanophores posterior to eye or melanophores on supraorbital cirri and pelvic fin. Melanophores may also occur along margins of dorsal, anal, and caudal fins. This species occurs in the western Atlantic from southern Florida and the Bahamas to Venezuela, including the Florida Keys, the Gulf of Mexico, and the Greater and Lesser Antilles. In the Gulf of Mexico it occurs in the Dry Tortugas. This species is associated with rock and rubble near coral reefs and grass beds from the shoreline to 6 m. Specimens from Florida, the Greater Antilles, Mexico, and Central America are considered a separate subspecies, S. stahli tekla, from the specimens from the Lesser Antilles and islands off South America, S. stahli stahli. Maximum known size is 40 mm TL. references: Longley and Hildebrand 1941 (as Auchenistius stahli); Springer 1955a; Böhlke and Chaplin 1968; Robins and Ray 1986; Hastings and Springer 1994; Smith 1997; Williams 2002d.

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624

BLENNIIDAE

Combtooth blennies

Combtooth blennies are generally elongate, slender to moderately deep bodied, and moderately compressed, often with cirri or fleshy flaps or tentacles on eye and occasionally with cirri or fleshy flaps on anterior naris and nape. Lateral profile of head is usually moderately to strongly blunt. Eye is of moderate size and located near dorsal margin of head. Nostril is paired, with anterior naris often tubelike and posterior naris porelike. Mouth is low on head, terminal to subterminal, and small to moderate in size. Upper jaw is not protrusible. Jaw teeth are incisor-like and in a single row, and are often closely packed and loosely attached, with some species having a pair of canine teeth at posterior end of each jaw. Teeth are present or absent in vomer but absent in palatine. First gill arch has gill rakers. Gill membranes are either connected under posterior surface of head or separated. Pectoral fin is well developed, with a broad vertical base and 10 to 18 rays. Dorsal fin is long and composed of 3 to 17 flexible spines and 9 to 119 rays, with spinous and rayed sections continuous or deeply notched. Dorsal fin spines are fewer than rays in all but one genus. Pelvic fin is jugular in location and consists of 1 obscure spine and 1 to 4 rays. Anal fin is long and consists of 2 spines, often obscure (especially in females), and 10 to 119 rays. All fin rays, except caudal fin rays in some genera, are unbranched. Caudal fin is truncate to convex. Lateral line tubes or canals extend the length of body or are restricted to anterior section. Scales are entirely lacking. Maximum known size is 540 mm TL, but specimens are usually less than 100 mm TL. Combtooth blennies occur worldwide in tropical to warm temperate seas, generally in less than 20 m. All species are benthic as adults, eggs are benthic, and larvae are pelagic. Most species live in shallow marine waters, but some occupy fresh or brackish waters. There are about 301 species in 53 genera, 21 species (1 introduced from the Indo-West Pacific) in 9 genera in the western Central Atlantic, and 14 species in 8 genera in the Gulf of Mexico. references: Springer 1986, 1993, 2001; Fritzsche and Fuiman 1982; Nelson 1994; Smith 1997; Smith-Vaniz et al. 1999; Williams 2002e.

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BLENNIIDAE 625

Key to the Species of the Gulf of Mexico (Adapted from Williams 2002e) 1a. Segmented caudal fin rays 10 or 11; pectoral fin rays usually 12; dorsal fin spines usually 11 . . . . . . . . . . . . . . . . . 2 1b. Segmented caudal fin rays usually 13; pectoral fin rays usually 13 to 15; dorsal fin spines usually 12 or 13 . . . . . . . . 3 2a. Lower jaw with prominent lip flaps . . . . . . . . . . . . . . . . . . . . . . . . Chasmodes saburrae p. 628 2b. Lower jaw without prominent lip flaps . . . . . . . . . . . . . . . . . . . . . Chasmodes longimaxilla p. 627 3a. Pectoral fin rays usually 15; lateral line discontinuous, with posterior end of anterior portion overlapping anterior end of posterior portion; total dorsal fin elements 31 or more . . . . . . . . . . . . . . . . . . . . Ophioblennius macclurei p. 638 3b. Pectoral fin rays usually 13 or 14; lateral line continuous or discontinuous, but if discontinuous, portions not overlapping; total dorsal fin elements 25 to 30 . . . . . . . . . . . . 4 4a. Ventral edge of upper lip smooth medially and crenulate laterally; small cirrus on nape between level of eye and origin of dorsal fin; dorsal fin completely or nearly completely divided into two sections; dorsal fin spines usually 13, with last very small and difficult to see; vomer with teeth . . . . . . . . . . . . . . . . . . . . . Entomacrodus nigricans p. 629 4b. Ventral edge of upper lip smooth over length; cirri on nape, if present, and numerous both anterior and posterior to eye; dorsal fin continuous, and if notched, notch not extending to base; dorsal fin spines usually 12, with last of moderate size; vomer without teeth . . . . . . . . . . . . . . . . . 5 5a. Numerous cirri on nape in longitudinal row and on eye . . . . . . . . . . . . . . . . . . . . . . . . . . . Scartella cristata p. 640 5b. Nuchal cirri absent, cirri on eye present or absent . . . . . . 6 6a. Gill openings continuous across isthmus . . . . . . . . . . . . . 7 6b. Gill openings restricted to side of head. . . . . . . . . . . . . . . 9 7a. Pectoral fin rays usually 13; total dorsal fin elements 25 to 27; single simple cirrus on eye (may be frayed at tip) . . . . 8 7b. Pectoral fin rays usually 14; total dorsal fin ray elements 28 to 30; several cirri on eye . . . . . . . . . . . . . . . . . . . . . Parablennius marmoreus p. 639 8a. Dorsal fin rays 13 or 14 (usually 14); anterior dorsal fin rays about same length as posterior rays; side of body with seven or eight prominent dark irregular bars; mature males without prolongation of dorsal fin; supraorbital cirri relatively large . . . . . . . . . . . . . Lupinoblennius vinctus p. 637 8b. Dorsal fin rays 14 or 15 (usually 15); anterior dorsal fin rays longer than posterior rays; side of body without prominent dark irregular bars; supraorbital cirri relatively small . . . . . . . . . . . . . . . . . . . . . Lupinoblennius nicholsi p. 636

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626 PERCIFORMES

9a. An enlarged canine tooth present in one or both jaws . . . 10 9b. No enlarged canine teeth in jaws . . . . . . . . . . . . . . . . . . 12 10a. Dorsal fin only slightly indented between spiny and rayed sections, with last dorsal fin spine not reduced and more than two-thirds the length of first dorsal fin ray; pelvic fin with 1 spine and 3 rays; body plain colored or mottled but without broad saddles . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10b. Dorsal fin with deep indentation between spiny and rayed sections, with last dorsal fin spine less than two-thirds the length of first dorsal fin ray; pelvic fin with 1 spine and 4 rays; body with six broad saddles above midline . . . . . . . . . . . . . . . . . . Hypleurochilus bermudensis p. 630 11a. Caudal fin with three or four dark bands on translucent background; supraorbital cirrus with 8 or fewer branches . . . . . . . . . . . . . . . . . Hypleurochilus caudovittatus p. 631 11b. Caudal fin uniformly pigmented; supraorbital cirrus with 21 or fewer branches . . . . Hypleurochilus multifilis p. 632 12a. Gill openings extending below level of pectoral fin base; dorsal fin rays 11 or 12; pelvic fin rays 4; head and usually base of pectoral fin with numerous small, pale spots (orange in life); last dorsal fin spine 3.7% to 7.5% of SL . . . . . . . . . . . . . . . . . . . . . . Hypsoblennius invemar p. 634 12b. Gill openings extending to fourth to eleventh pectoral fin ray; dorsal fin rays 13 to 16; pelvic fin rays 3; head and pectoral fin base without pale spots; last dorsal fin spine 8.5% to 15.5% of SL . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13a. Upper lip with free dorsal margin extending around snout; fleshy lobes of lower lip semicircular in lateral view and separated anteroventrally by broad shieldlike lobe; dark spot usually absent on spinous section of dorsal fin, but if present, centered on third spine; mandibular pores three . . . . . . . . . . . . . . . . . . . . . . Hypsoblennius ionthas p. 635 13b. Upper lip without free dorsal margin extending around snout; fleshy lobes of lower lip almost linear in lateral view and separated anteroventrally by broad, smooth area; dark spot usually present on spinous section of dorsal fin and centered on second spine; mandibular pores four . . . . . . . . . . . . . . . . . . . . . . . . Hypsoblennius hentz p. 633

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BLENNIIDAE 627

Chasmodes longimaxilla

Williams, 1983

Chasmodes longimaxilla has a restricted gill slit and a lower jaw lacking prominent lip flaps, and either lacks or has obsolete supraorbital cirri in adults. Dorsal profile of head is moderately sloping and slightly convex, and snout is acute. Upper lip has narrow free dorsal margin that is discontinuous over center of jaw. Anterior naris is tubular, with a filamentous flap extending from posterior margin. Cirrus on dorsal surface of eye is simple and decreases in size with growth. Jaw teeth are closely packed and acutely pointed. Vomer and palatine lack teeth. Gill membranes are broadly joined to isthmus, restricting gill opening to distance from upper margin of operculum to about upper one-fourth of pectoral fin base. Gill rakers on first arch number 9 to 12 (modally 11). Measurements are expressed as percent of SL: head length 29.2%–33%, snout length 7.2%– 8.8%, eye diameter 5.1%– 6.1%, interorbital width 2.9%–3.8%, maxillary length 14%– 22.5%, gill opening length 5.1%– 8%, body depth at pelvic insertion 24%–26.1%, caudal peduncle depth 7.4%– 8.3%. Pectoral fin has

11 to 13 (usually 12) rays. Dorsal fin has 11 spines and 16 to 19 (usually 18 or 19) rays. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 16 to 20 (usually 18 or 19) rays. Caudal fin has 10 or 11 segmented rays. Color in life is olivaceous, with light stripes or mottling, an iridescent blue spot on membrane between first and second dorsal fin spines, and an orange streak from spot to about tenth spine. This species occurs in the western Atlantic in the Gulf of Mexico from Pensacola to Veracruz. It is associated with oyster beds. It is very similar to C. bosquianus, which occurs from New York to northeastern Florida and was described as a subspecies of this taxon (Williams 1983). Maximum known size is 86 mm SL. references: Springer 1959a; Williams 1983 (as C. bosquianus longimaxilla), 2002e; Robins and Ray 1986; Boschung 1992; Smith 1997; CastroAguirre et al. 1999 (all above as C. bosquianus except Williams).

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628 PERCIFORMES

Chasmodes saburrae Florida blenny

Jordan and Gilbert, 1882

Chasmodes saburrae has a restricted gill slit and a lower jaw possessing prominent lip flaps, and either lacks or has obsolete supraorbital cirri in adults. Dorsal profile of head is moderately steep and moderately convex, and snout is acute. Upper lip has a narrow free dorsal margin that is discontinuous over center of jaw. Anterior naris is tubular, with a filamentous flap extending from posterior margin. Cirrus on dorsal surface of eye is simple and decreases in size with growth. Jaw teeth are closely packed and broadly rounded. Vomer and palatine lack teeth. Gill membranes are broadly joined to isthmus, restricting gill opening to distance between upper margin of operculum and about upper fourth of pectoral fin base. Gill rakers on first arch number 12 to 14. Measurements are expressed as percent of SL: head length 26.6%–31.6%, snout length 6.4%– 8.3%, eye diameter 5.2%–7.6%, interorbital width 3.1%–3.9%, maxillary length 9.5%– 12.3%, gill opening length 5%–7.1%, body depth at pelvic insertion 24.9%–28.4%, caudal peduncle depth 7.7%–9.5%. Pectoral fin has 11 to 13 (usually 12) rays. Dorsal fin has

10 or 11 spines and 16 to 20 (usually 18 or 19) rays. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 17 to 20 (usually 18) rays. Caudal fin has 10 to 13 (usually 11) segmented rays. Color in life is olivaceous or brown, with light stripes or mottling, an iridescent blue spot on membrane between first and second dorsal fin spines, and a streak from spot to about tenth spine. This species occurs in the western Atlantic from Edgewater on the east coast of Florida to the Chandeleur Islands in the northern Gulf of Mexico. It overlaps with C. longimaxilla between Pensacola, FL, and the Chandeleur Islands. Chasmodes longimaxilla is restricted to oyster reefs, and C. saburrae is restricted to sea-grass beds. Food consists of annelids, amphipods, isopods, shrimps, and ray-finned fish larvae. Maximum known size is 80 mm SL. references: Springer 1959a (as C. bosquianus saburrae); Springer and Woodburn 1960; Williams 1983, 2002e; Robins and Ray 1986; Boschung 1992; Smith 1997.

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BLENNIIDAE 629

Entomacrodus nigricans Pearl blenny

Gill, 1859

Entomacrodus nigricans has simple cirri on eyes and on nape, separate dorsal fins, and an incomplete lateral line. Anterior profile of head is nearly vertical to eye. Dorsal margin of upper lip is free laterally but discontinuous over center of jaw, and ventral edge is smooth medially and crenulate laterally. Lower lip has a short, free lower margin. Anterior naris has a raised margin and a palmate cirrus extending from posterior margin. Posterior naris is porelike. Cirri on dorsal surface of eye number 1 to 11 (increase with growth) and are shorter than eye diameter. Bony interorbital space is slightly concave. A short, slender simple cirrus is present on each side of nape. Comblike jaw teeth are very slender, bluntly pointed, and embedded in gums but not in jaw bones. A moderately large, recurved canine tooth is located medial to incisor-like teeth in corner of lower jaw. Teeth are present on vomer. Gill rakers on first arch number 14 to 20 (usually 15 to 18). Gill membranes are broadly connected but are free of isthmus. Measurements are expressed as percent of SL: head length 22.1%–23.2%, snout length 5.5%– 6.2%, eye diameter 3.5%– 4.6%, interorbital width 2.5%–3.1%, gill opening length 7.4%–9.9%, body depth at pelvic insertion 18.5%–22.7%, caudal peduncle depth 8.7%–10.5%. Pectoral fin has 13 or 14 rays. Dorsal fin has 13 spines (last very small) and 13 to 16 (usually 14 or 15) rays. Pelvic fin

has 1 spine and 4 rays. Anal fin has 2 spines and 14 to 17 rays (usually 16), with spines and first 1 to 4 rays bearing fleshy rugosities in mature males. Caudal fin has 13 segmented rays. Lateral line is continuous but terminates under spiny dorsal fin. Vertebrae number 33 to 35. Color is tan to light, slightly darker dorsally than ventrally, with six or six and one-half dark brown bars extending to ventral midline. Narrow light bands often subdivide bars, and interspaces between bands are often marked with white spots. Head is brown to dark brown, with dark spot posterior to eye and 7 to 10 dark stripes originating on upper lip and converging along ventral midline. Vertical fins are brown to dark brown, with dark spots and tan streaks on fin rays. This species occurs in the western Atlantic from southern Florida and Bermuda to Venezuela, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is known from the Dry Tortugas and Campeche Bank, at East Triangles Reef, in the Gulf of Mexico. This species is associated with rocky substrates from near the surface to one meter in depth. Maximum known size is 63 mm SL. references: Longley and Hildebrand 1941 (as Salarichthys textiles); Springer 1967; Randall 1996; Smith 1997; Smith-Vaniz et al. 1999; Williams 2002e.

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630 PERCIFORMES

Hypleurochilus bermudensis Barred blenny

Beebe and Tee-Van, 1933

Hypleurochilus bermudensis has a branched cirrus on eye, a restricted gill slit, and a pair of canine teeth in both jaws. Anterior profile of head is steep and rounded. Upper lip has a free dorsal margin that is discontinuous over center of jaw. Lower lip has a fleshy lobe extending from corner of mouth to about half of jaw length and running about parallel to inner margin of lip. Anterior naris has a long flap bearing one to three cirri extending from posterior margin. Cirrus on dorsal surface of eye is longer than eye, is round in cross section, and may have one to six simple branches. Jaw teeth are elongate, closely packed, and bluntly rounded. Corner of each jaw bears a canine tooth, with that of upper jaw external to and smaller than that of lower jaw. Vomer lacks teeth. Gill membranes are broadly joined to isthmus, restricting gill opening to distance from upper margin of operculum to ventral end of pectoral fin base. Measurements are expressed as percent of SL: head length 26.7%– 31.7%, snout length 9%–11.4%, eye diameter 6.1%–7.3%, interorbital width 2.9%– 4%, length of gill opening 11.5%–14.8%, body depth at pelvic insertion 25.5%–26.5%, caudal peduncle depth 7.8%–9.9%. Pectoral fin has 13 or 14 rays. Dorsal fin has 12 spines and

13 (rarely 12 or 14) rays, with last spine less than two-thirds the length of first ray. Pelvic fin has 1 spine and 4 rays. Anal fin has 2 spines and 15 rays, with spines and first ray bearing fleshy rugosities in mature males. Lateral line is disjunct, with anterior section originating at upper margin of operculum and extending to near end of spinous section of dorsal fin and consisting of biramous pores, and posterior section extending along midflank and consisting of short canal segments. Color is grayish tan dorsally and laterally, with six distinct bars along upper trunk, and grayish tan ventrally. Caudal fin is light tan, with a small, elongate brown spot on base and three to five dark bands distally. This species occurs in the western Atlantic from the Florida Keys, Bermuda, the Bahamas, and the Gulf of Mexico. In the Gulf of Mexico it occurs at the Dry Tortugas and Flower Garden Banks and at Isla de Lobos off Veracruz. It occurs to about 27 m on coral reefs and rocky substrates. Maximum known size is 40 mm SL.

references: Longley and Hildebrand 1941; Randall 1966b; Boschung 1992; Bath 1994; Smith 1997; Smith-Vaniz et al. 1999; Williams 2002e.

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BLENNIIDAE 631

Hypleurochilus caudovittatus

Bath, 1994

Hypleurochilus caudovittatus has a branched cirrus on eye, a restricted gill slit, and a pair of canine teeth in both jaws. Anterior profile of head is steep and rounded. Upper lip has a free dorsal margin that is discontinuous over center of jaw. Lower lip has a fleshy lobe extending from corner of mouth to about half of jaw length and running parallel to inner margin of lip. Anterior naris has a long flap bearing one to three cirri extending from posterior margin. Cirrus on dorsal surface of eye is longer than eye, is round in cross section, and may have three to seven simple branches. Jaw teeth are elongate, setiform, and bluntly rounded. Corner of each jaw bears a large canine, with that of upper jaw external to and smaller than that of lower jaw. Vomer lacks teeth. Gill membranes are broadly joined to isthmus, restricting gill openings to distance from upper margin of operculum to ventral end of pectoral fin base. Measurements are expressed as percent of SL: head length 25.4%–30.8%, snout length 8.3%–9.8%, eye diameter 6.2%– 6.6%, length of gill opening 14.5%–15.2%, body depth at pelvic insertion 24.7%–29.5%, caudal pe-

duncle depth 8.1%–9.1%. Pectoral fin has 13 or 14 rays. Dorsal fin has 12 or 13 spines and 14 or 15 rays, with last spine more than twothirds the length of first ray. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 16 rays, with spines and first ray bearing fleshy rugosities in mature males. Lateral line is disjunct, with anterior section originating at upper margin of operculum and extending to near end of spinous section of dorsal fin and consisting of biramous pores, and posterior section extending along midflank and consisting of short canal segments. Color is grayish brown. A dark spot often occurs on membrane between first and second dorsal fin spines. Caudal fin is transparent except for a dark spot on base and three to five dark bands distally. This species occurs in the western Atlantic from the eastern Gulf of Mexico between Sarasota and St. Andrew Bay, FL. It is generally associated with sandy substrates. Maximum known size is 47.8 mm SL. references: Bath 1994; Sarah Hopfensperger, pers. comm., February 26, 2002; Williams 2002e.

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632 PERCIFORMES

Hypleurochilus multifilis

(Girard, 1858)

Hypleurochilus multifilis has a branched cirrus on eye, a restricted gill slit, and canine teeth in both jaws. Dorsal profile of head is steep and rounded. Mature males possess a large predorsal-fin neck fold. Upper lip has a free dorsal margin that is discontinuous over center of jaw. Lower lip has a fleshy lobe extending from corner of mouth to about half of jaw length and running about parallel to inner margin of lip. Anterior naris has a long flap bearing 2 to 15 cirri extending from posterior margin. Cirrus on dorsal surface of eye is very elaborate, longer than eye, and round in cross section, and it bears a series of fine secondary cirri, about half the length of major cirrus, on each side of base. Jaw teeth are elongate, incisorlike, and bluntly rounded. Corner of each jaw bears a large canine, with that of upper jaw external to that of lower jaw. Vomer lacks teeth. Gill opening extends from upper margin of operculum to ventral end of pectoral fin base. Measurements are expressed as percent of SL: head length 26.7%–30.6%, snout length 8.8%–9.6%, eye diameter 4.2%–5.5%, interorbital width 2%–2.8%, length of gill opening 15.8%–20%, body depth at pelvic insertion 23.8%–26.6%, caudal peduncle depth 7.8%– 10.1%. Pectoral fin has 13 or 14 rays. Dorsal fin has 12 (rarely 11 or 13) spines and 14 or

15 (rarely 13 or 16) rays, with last spine more than two-thirds the length of first ray. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 16 or 17 (rarely 15) rays, with spines and first ray bearing fleshy rugosities in mature males. Lateral line is disjunct, with anterior section originating at upper margin of operculum, extending to near end of spinous section of dorsal fin, and consisting of biramous pores, and posterior section extending along midflank and consisting of short canal segments. Color is uniformly grayish brown in males and mottled grayish brown in females. A dark spot often occurs on membrane between first and second dorsal fin spines, and caudal fin may have a dark spot on base. Females often have faint bars on transparent membrane. This species occurs in the western Atlantic in the Gulf of Mexico. In the Gulf it occurs from St. Andrew Bay, FL, to Port Aransas, TX. It is associated with oyster reefs and pier pilings. Maximum known size is 100 mm TL. references: Springer and Woodburn 1960; Randall 1966b; Bath and Wirtz 1981; Robins and Ray 1986; Boschung 1992 (all as H. geminatus); Bath 1994; Sarah Hopfensperger, pers. comm., February 26, 2002; Williams 2002e.

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BLENNIIDAE 633

Hypsoblennius hentz Feather blenny

(Lesueur, 1825)

Hypsoblennius hentz has a cirrus on eye, a restricted gill slit, and an incomplete lateral line. Dorsal profile of head is steep and indented posterior to eye. Upper lip has a free dorsal margin that is narrowly discontinuous over center of jaw. Lower lip has a fleshy lobe extending from corner of mouth to about half of jaw length and running about parallel to inner margin of lip. Anterior naris has a long filamentous flap extending from posterior margin. Cirrus on dorsal surface of eye is shorter to slightly longer than eye diameter and bears a number of unbranched or branched secondary cirri. Bony interorbital space is concave. Jaw teeth are setiform and bluntly pointed. Vomer lacks teeth. Gill membranes are broadly joined to isthmus, restricting gill opening to distance from upper margin of operculum to about midlength of pectoral fin base. Measurements are expressed as percent of SL: head length 27.9%– 29.5%, snout length 6%– 8.2%, eye diameter 4.7%–5.2%, interorbital width 1.9%–2.9%, length of gill opening 10%–12.3%, body depth at pelvic insertion 25.5%–29.1%, caudal peduncle depth 7.6%–9.1%. Pectoral fin has 13 to 15 rays. Dorsal fin is continuous and has 11 to 13 robust and stiff spines and 13 to 16 rays. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 14 to 17 rays, with spines and first ray bearing fleshy rugosities in mature males.

Caudal fin has 13 segmented rays. Color is yellowish brown dorsally and whitish ventrally, with many variably shaped dark brown spots that are better defined on head and often form irregular bands on body. Brown spots form two or three dark V-shaped bands on ventral aspect of head. A large, dark oval-shaped mark is located posterior to eye and separated from eye by a narrow pale area. A dark spot is usually present on anterior section of dorsal fin and is centered on second spine. This species occurs in the western Atlantic from Nova Scotia to Campeche Bay in shallow water but is discontinuous around Florida. On the east coast it has not been collected south of St. Lucie Inlet, FL, and on the west coast it is not known from south of Sanibel Island, FL. It is common in sea-grass flats. Spawning takes place during the summer on natural oyster reefs. Food consists of plant material, hydroids, annelids, and amphipods. Eggs are deposited in empty, intact oyster shells. Maximum known size is 103 mm SL.

references: Hildebrand and Schroeder 1928; Hildebrand and Cable 1930; Springer and Woodburn 1960; Hoese and Moore 1977, 1998; SmithVaniz 1980; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Williams 2002e.

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634 PERCIFORMES

Hypsoblennius invemar Tessellated blenny

Smith-Vaniz and Acero P., 1980

Hypsoblennius invemar has a cirrus on eye, a restricted gill slit, and an incomplete lateral line. Dorsal profile of head is steep and indented posterior to eye. Upper lip has a free dorsal margin that is broadly discontinuous over center of jaw. Lower lip has a fleshy lobe extending from corner of mouth to about half of jaw length and running about parallel to inner margin of lip. Anterior naris has a short, simple to moderately palmate cirrus extending from posterior margin. Cirrus on dorsal surface of eye is simple in juveniles and subdivided into four branches in adults, with that of males considerably longer than that of females. Bony interorbital space is concave. Jaw teeth are bluntly pointed. Vomer lacks teeth. Gill membranes are broadly fused to isthmus, restricting gill opening to distance from upper margin of operculum to below pectoral fin base. Measurements are expressed as percent of SL: head length 26.5%–32.5%, snout length 7.5%– 8.8%, eye diameter 6.3%– 6.9%, interorbital width 1.6%–1.9%, length of gill opening 16%–18.1%, body depth at pelvic inser-

tion 27%–30%, caudal peduncle depth 9.4%– 10.4%. Pectoral fin has 13 to 15 (usually 14) rays. Dorsal fin has 11 or 12 spines and 11 or 12 rays, with last spine 3.7%–7.5% of SL. Pelvic fin has 1 spine and 4 rays. Anal fin has 2 spines and 13 or 14 rays, with spines and first ray bearing fleshy rugosities in mature males. Caudal fin has 13 segmented rays. Color in life is greenish brown. Head and, usually, pectoral fin base are blue, with bright orange to brick red spots surrounded by dark ring. In preservative, head and body are brown dorsally and dark tan ventrally, with numerous pale spots on head and, usually, pectoral fin base. This species occurs in the western Atlantic from the northern Gulf of Mexico off Louisiana, the Lesser Antilles, Venezuela, and Colombia. It is associated with empty barnacle (Balanus) tests. Maximum known size is 47 mm SL.

references: Smith-Vaniz 1980; Smith 1997; Hoese and Moore 1998; Williams 2002e.

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BLENNIIDAE 635

Hypsoblennius ionthas Freckled blenny

(Jordan and Gilbert, 1882)

Hypsoblennius ionthas has a cirrus on eye, a restricted gill slit, and an incomplete lateral line. Dorsal profile of head is steep and indented posterior to eye. Upper lip has a free dorsal margin that is continuous over center of jaw. Lower lip has a fleshy semicircular lobe, separated from opposite lobe by a broad shield-shaped medial lobe. Anterior naris has a long filamentous flap extending from posterior margin. Cirrus on dorsal surface of eye is longer than eye diameter and bears a number of unbranched or branched secondary cirri. Bony interorbital space is concave. Jaw teeth are bluntly pointed. Vomer lacks teeth. Gill membranes are joined to isthmus, restricting gill opening to distance from upper margin of operculum to base of fourth to eleventh pectoral fin ray. Measurements are expressed as percent of SL: head length 23.8%–25.5%, snout length 8.1%– 8.9%, eye diameter 4%– 5.2%, interorbital width 0.8%–2.1%, length of gill opening 7.6%–11.5%, body depth at pelvic insertion 24.2%–26.7%, caudal peduncle depth 9.3%–10.9%. Pectoral fin has

13 to 15 rays. Dorsal fin has 11 to 13 robust and stiff spines and 13 to 16 rays, with last spine 8.5%–15.5% of SL. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 14 to 17 rays, with spines and first ray bearing fleshy rugosities in mature males. Color is yellowish brown dorsally and whitish ventrally, with dark bands on body. Head has two dark lines from eye to mouth and a dark bar on cheek with a blackish anterior edge. Dark spot is usually absent on anterior section of dorsal fin, but if present, is centered on third spine. Females have dark freckles on lower part of head that are occasionally continued onto body. This species occurs in the western Atlantic from North Carolina to southeastern Florida (Matanzas Inlet), and from Cedar Key, FL, to Aransas Bay, TX, in the Gulf of Mexico. It is associated with shallow bays and estuaries. Maximum known size is 70 mm SL. references: Hoese and Moore 1977, 1998; SmithVaniz 1980; Robins and Ray 1986; Boschung 1992; Smith 1997; Williams 2002e.

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636 PERCIFORMES

Lupinoblennius nicholsi Highfin blenny

(Tavoga, 1954)

Lupinoblennius nicholsi has a cirrus on eye, gill slits continuous across isthmus, and a discontinuous and incomplete lateral line. Snout is moderately acute, and dorsal profile of head is strongly oblique. Lips are not conspicuous and lack free dorsal margin across symphyses. Anterior naris consists of a short, slender tube and has a small, triangular terminal flap. Posterior naris is a moderately large pore with a raised margin. Cirrus on dorsal surface of eye is flat, slender, and lanceolate. Cirri are about twice orbit diameter in males and rudimentary in females and juveniles. Jaw teeth are bluntly pointed, with a posterior canine in both jaws in males and in lower jaw in females. Posterior canines of males are larger than that of females. Vomer lacks teeth. Gill membranes are free of isthmus. Measurements are expressed as percent of SL: head length 27%–31.3%, body depth at pelvic fin insertion 25.6%–27%, caudal peduncle depth 8.5%– 8.9%; or as percent of head length: snout length 14.3%–20%, eye diameter about 25%. Pectoral fin has 13 rays. Dorsal fin has 12 or 13 spines and 13 or 15 (usually 15) rays, with anterior 4 spines prolonged, especially in mature males. Pelvic fin

has 1 spine and 4 rays. Anal fin has 2 spines and 15 to 17 (usually 16 or 17) rays, with spines and anterior rays bearing fleshy rugosities in males. Caudal fin is broadly rounded and has 13 segmented rays. Anterior portion of lateral line extends from upper opercular angle to sixth to eleventh dorsal fin spine, and midlateral portion originates over anal fin and consists of separate pores. A single pore is located on head between dorsal fin and lateral line. Color is olivaceous green in life, with two dark stripes running from eye to mouth and from eye to gular region. Males are dark and have 8 to 12 light oblique lines on side. Males and juveniles are light, with fine whitish reticulations on dorsal and lateral surfaces. This species occurs in shallow water in the western Atlantic from southeastern Florida and the southern and southwestern Gulf of Mexico. Maximum known size is 44 mm SL. references: Fowler 1954 (as Semablennius gallowayi); Dawson 1970 (as Blennius nicholsi); Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Williams 2002e.

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BLENNIIDAE 637

Lupinoblennius vinctus Mangrove blenny

(Poey, 1867)

Lupinoblennius vinctus has a cirrus on eye, gill slits continuous across isthmus, and a discontinuous and incomplete lateral line. Snout is moderately acute, and dorsal profile of head is moderately oblique. Lips are not conspicuous and lack free dorsal margin across symphyses. Anterior naris ends as a short, slender tube, usually with a small terminal filament. Posterior naris is a large pore with a raised margin. Cirrus on dorsal surface of eye is flattened, with several short fringes on distal margin. Cirri of males are about twice as long as those of females and juveniles. Jaw teeth are bluntly pointed, with males possessing an enlarged posterior canine in both jaws and females possessing a smaller posterior canine in lower jaw. Gill membranes are free of isthmus. Gill opening originates at level of upper margin of operculum. Measurements are expressed as percent of SL: head length 28.6%–32.7%, snout length 8.1%– 8.9%, eye diameter 4%–5.2%, body depth at pelvic insertion 22.7%–28.7%, caudal peduncle depth 7.8%–9.7%. Pectoral fin has 12 to 14 rays. Dorsal fin has 12 spines and 13 or 14 rays. Pelvic fin has 1 spine and 4 rays. Anal fin has 2 spines and 14 to 16 (usually 15 or 16) rays, with spines and anterior rays bear-

ing fleshy knobs in mature males. Caudal fin is subtruncate to broadly rounded and has 13 segmented rays. Anterior portion of lateral line extends from upper opercular angle to ninth or tenth dorsal fin spine and consists of an unbranched, pored tubiform segment followed by four short two-pored tubes. Midlateral portion of lateral line is located above anterior section of anal fin and consists of four to eight separated short two-pored segments. Two pores are located on head between dorsal fin and lateral line. Color is tan to brown, with dusky bar between orbits. Side of body is patterned with seven or eight irregular dark bars, often with light blotches superimposed on bars, to form series of O- or U-shaped markings. A dark blotch occurs on membrane between first and second dorsal fin spines. This species occurs in the western Atlantic from the southern Gulf of Mexico to Panama and from the Antillean islands of Jamaica, Antigua, and Trinidad. It is often associated with low-salinity estuaries. Maximum known size is 36 mm SL. references: Dawson 1970; Smith 1997 (both as L. dispar); Williams 2002e.

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638 PERCIFORMES

Ophioblennius macclurei Redlip blenny

(Silvester, 1915)

Ophioblennius macclurei has cirri on the eye and nape and a discontinuous lateral line. Profile of head is slightly negatively sloped. Mouth is terminal to slightly inferior. Lips are not conspicuous, and dorsal lip lacks a free dorsal margin across symphysis and possesses a fringed ventral margin. Anterior naris terminates as a short tube that bears a branched cirrus. Posterior naris is a large pore with a slightly raised margin. Supraorbital cirrus is unbranched, very slender, and shorter than eye diameter. A pair of small, slender cirri is located on side of nape. Jaw teeth are very narrow and bluntly pointed, with an additional stout canine tooth located medial to last incisor of lower jaw. Gill membranes are broadly joined but free of isthmus. Measurements are expressed as percent of SL: head length 19.6%– 21%, snout length 6.5%–7.4%, eye diameter 3.9%– 4.4%, upper jaw length 7%–9.7%, body depth at pelvic fin insertion 24.4%– 29.4%, caudal peduncle depth 6.7%–7.9%. Pectoral fin has 14 to 15 rays. Dorsal fin has 12 spines and 19 to 21 rays. Pelvic fin has 1 embedded spine and 4 rays. Anal fin has 2 spines and 20 to 22 rays. Caudal fin is broadly

rounded to truncate and has 13 segmented rays. Lateral line is interrupted at about midlength, with anterior and posterior sections overlapping medially. Color is dark brown or reddish brown, with lower lip, lower section of pectoral fin, and margins of dorsal and anal fins bright red. A dark spot is located behind eye, and caudal fin has a dark blotch along median rays. Juvenile specimens may have pale bars separating squarish blotches on side. This species occurs in the western Atlantic from New York and Bermuda to northern South America, including the Gulf of Mexico. In the Gulf of Mexico it occurs off the west coast of Florida, in the Dry Tortugas, on reefs off the Texas coast, along the coast of Veracruz, and on Alacrán Reef off Yucatán. Food consists of filamentous algae. Maximum known size is about 125 mm TL.

references: Longley and Hildebrand 1941 (as Rupiscates atlanticus); Springer 1962; CastroAguirre and Márquez-Espinoza 1981; Randall 1996; Schaldach et al. 1997; Smith 1997; SmithVaniz et al. 1999 (all as Ophioblennius atlanticus); Williams 2002e.

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BLENNIIDAE 639

Parablennius marmoreus (Poey, 1876) Seaweed blenny

Parablennius marmoreus has several cirri on eye, and gill openings continuous across isthmus. Snout is blunt, and dorsal profile of head is very steep. Lips are not conspicuous, dorsal lip lacks a free margin across symphysis, and ventral lip has a poorly developed lip fold. Anterior naris terminates as a short tube that bears a triangular flap. Posterior naris is a small pore with a slightly raised margin. Supraorbital cirrus has several branches. Jaw teeth are bluntly pointed, with posterior canines in both jaws. Vomer lacks teeth. Gill rakers on first arch number 11 to 13. Measurements are expressed as percent of SL: head length 25.7%– 29.6%, snout length 5.3%–9.3%, eye diameter 5.2%–7.1%, upper jaw length 9.9%– 11.7%, body depth at pelvic fin insertion 21.7%–23.8%, caudal peduncle depth 8%– 8.2%. Pectoral fin has 14 rays. Dorsal fin has 12 spines and 17 to 18 rays. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 19 or 20 rays. Caudal fin is broadly rounded and has 13 segmented rays. Lateral line extends

from upper opercular angle to near end of spinous section of dorsal fin and has 11 to 13 pores. Color is tan, rusty, or orangish, with orange spots on head and brown spots on side. Spots may coalesce to form a broad dusky stripe extending from head to caudal fin, with stripe most distinct anteriorly. Some specimens are predominately yellow. This species occurs in the western Atlantic from New York and Bermuda to northern South America, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is known from the northern Gulf of Mexico. This species is associated with rock reefs. Food consists mostly of algae. Maximum known size is about 80 mm SL. references: Kanazawa 1952; Springer and Woodburn 1960; Briggs et al. 1964; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; SmithVaniz et al. 1999; Williams 2002e.

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640 PERCIFORMES

Scartella cristata Molly miller

(Linnaeus, 1758)

Scartella cristata has cirri on eye and nape, and gill openings are continuous across isthmus. Snout is blunt, and dorsal profile of head is steep and rounded. Lips are not conspicuous, dorsal lip lacks a free margin across symphysis, and ventral lip has a poorly developed lip fold. Anterior naris terminates as a short tube that bears a branched cirrus. Posterior naris is a large pore with a slightly raised margin. Supraorbital cirrus has several slender branches. Nape possesses a dense patch of slender cirri along dorsal midline. Jaw teeth are bluntly pointed, with a stout canine located medial to last incisor of lower jaw. Gill membranes are free of isthmus. Gill rakers on first arch number 17 to 21. Measurements are expressed as percent of SL: head length 22.9%– 28.8%, snout length 7.6%–11.2%, eye diameter 4.6%– 6%, upper jaw length 7.5%–9.4%, body depth at pelvic fin insertion 25%–30.3%, caudal peduncle depth 8.8%–10.1%. Pectoral fin has 14 rays. Dorsal fin has 12 spines and 14 or 15 rays. Pelvic fin has 1 spine and 3 rays. Anal fin has 2 spines and 16 or 17 rays, with spines fleshy and thickened in males. Caudal

fin is broadly rounded to truncate and has 13 segmented rays. Lateral line is continuous but is prominent only from upper margin of operculum to near end of spinous section of dorsal fin. Color is dull green to olive, with dark bars on body extending onto lower part of dorsal fin. Pale spots often occur on dorsal fin. This species occurs in the tropical Atlantic. In the western Atlantic it occurs from Florida and Bermuda to Brazil, including the northern and southern Gulf of Mexico and the Greater and Lesser Antilles. It occurs in tide pools and shallow water to 10 m along rocky shores. It also occurs in Sargassum rafts, in which case it is brilliant yellow in color (J. J. Williams, pers. comm., March 18, 2002). Maximum known size is about 100 mm TL.

references: Longley and Hildebrand 1941 (as Blennius cristatus); Randall 1968a, 1996; Hoese and Moore 1977, 1998; Castro-Aguirre 1978 (as B. cristatus); Castro-Aguirre and MárquezEspinoza 1981; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; SmithVaniz et al. 1999; Williams 2002e.

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641

GOBIESOCIDAE

Clingfishes

Clingfishes are generally depressed anteriorly and subcylindrical or compressed posteriorly, with a single dorsal fin and pelvic fins and part of pectoral fins modified into thoracic sucking disc. Snout is short to long. Nostril is paired, with anterior naris tubular and posterior naris usually tubular. Eye is on dorsolateral aspect of head and is small to moderate in size. Mouth is terminal to subterminal and small to moderate in size. Jaw teeth range from villiform to fanglike and are arranged in patches or rows. Gill membranes are usually free of isthmus but are occasionally attached, and gills are located on three to three and one-half arches (no slit behind last arch). Pectoral fin is broad and fanlike. Dorsal fin is single, consists entirely of rays, and is posteriorly located. Pelvic fins consist of four rays that are joined to form part of adhesive sucking disc located between head and trunk. Pelvic rays make up lateral edges of disc, and fourth ray is joined to lower portion of pectoral fin base by membrane. Disc bears flattened papillae along its anterior lateral margin, posterior margin, and central region. Anal fin is similar in size, shape, and position to dorsal fin and lacks spines. Scales are absent, and lateral line consists of pores that are well developed on head only. Vertebrae number 25 to 54. This description does not apply to Alabes, an eel-shaped taxon from Australia. Maximum known size is less than 90 mm TL. Clingfishes occur worldwide in tropical to warm temperate seas and in brackish and fresh waters from near shore to 200 m. Sucking disc is used to attach fish to hard substrates and plants in areas subjected to wave or tidal action. Food consists of small invertebrates and epiphytes. There are about 120 species in 36 genera worldwide, 25 species in 8 genera in the western Central Atlantic, and 2 or 3 species in 1 or 2 genera in the Gulf of Mexico. references: Briggs 1955; Fritzsche and Fuiman 1982; Nelson 1994; Smith 1997; Smith-Vaniz et al. 1999; Hutchins 2001a; McEachran 2002b.

Key to the Species of the Gulf of Mexico (Adapted, in part, from Briggs 1955) 1a. Upper lip narrow and about same width in front of snout as on side; lowest 5 to 7 pectoral fin rays markedly reduced; disc length 15% to 22% of SL . . . . . . . . . . . . . . . . . . . . . . . . . Acyrtops beryllinus p. 643 1b. Upper lip broad and much wider in front of snout than on side; lowest 5 to 7 pectoral fin rays not markedly reduced; disc length 24% to 43% of SL . . . . . . . . . . . . . . . . . . . . . 2

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642 PERCIFORMES

2a. Central margin of upper lip without shallow lobes; pectoral fin rays 19 to 22 (usually 21) . . . . . . . . . . . . . . . . . . . . . . . Gobiesox punctulatus p. 644 2b. Central margin of upper lip with well-developed papillae; pectoral fin rays 22 to 26 (usually 24) . . . . . . . . . . . . . . . . . . . . . . . . Gobiesox strumosus p. 645

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GOBIESOCIDAE 643

Acyrtops beryllinus Emerald clingfish

(Hildebrand and Ginsburg, 1926)

Acyrtops beryllinus has a narrow upper lip, short median fins, and a relatively small sucking disc. Head is slightly broader than body, and snout is continuous with head. Nares are tubelike, with anterior naris possessing a small flap on posterior margin, and posterior naris located just anterior to anterior margin of eye. Eye diameter is 66.7% to 76.9% of interorbital distance. Fleshy lobes on head are limited to lower jaw. Upper lip is narrow. Lower jaw is smooth. Teeth in front of upper jaw are pointed and compressed, with conical teeth behind these and on sides of jaw. Teeth in front of lower jaw are large and incisor-like, with conical teeth behind these and on side of jaw. Suboperculum bears a sharp spine covered by skin and extending from operculum. Gill membranes are united and are free of isthmus. Second and third gill arches each have six or seven gill rakers. Anus is closer to anal fin origin than to posterior margin of disc. Measurements are expressed as percent of SL: head length 32.9%–34.9%, head width 28.5%–32.8%, snout length 9.2%–11.4%, disc length 18.4%– 20.4%, body depth 14.9%–19.2%. Pectoral

fin has 21 to 23 rays, and lower 5 to 7 rays are reduced in size. Fleshy pad on pectoral fin base is poorly developed and lacks free posterior margin. Dorsal fin has 5 to 7 rays, and origin is closer to caudal fin base than to upper pectoral fin base. Pelvic sucking disc is single and rather small, and bears papillae on anterior, posterior, and single middle sections. Anal fin has 5 or 6 rays. Caudal fin is truncate and has 8 rays. Color is pale emerald green to brownish, with golden to brassy lines occasionally radiating from eye. This species occurs in the western Atlantic from southern Florida and the Bahamas to Belize, including the Florida Keys, the Gulf of Mexico, and the Lesser Antilles. In the Gulf of Mexico there is an unconfirmed record from Veracruz. It is closely associated with turtle grass. Maximum known size is 20 mm SL. references: Briggs 1955; Gould 1965; Böhlke and Chaplin 1968; Johnson and Greenfield 1983; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; McEachran 2002b.

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644 PERCIFORMES

Gobiesox punctulatus Stippled clingfish

(Poey, 1876)

Gobiesox punctulatus has poorly developed papillae on head, a broad upper lip, and a relatively large sucking disc. Head is broader than body, and snout is continuous with head. Upper lip is broadest near symphysis. Nares are tubelike, with anterior naris possessing a bilobed flap on posterior margin, and posterior naris located just anterior to anterior margin of eye. Upper lip is smooth, and papillae are lacking on snout above premaxillary groove. Teeth in front of upper jaw are conical, irregular in shape and size, and arranged in a patch, and those on side are conical and arranged in a single row. Teeth in lower jaw are arranged in two rows, with those in front narrow and incisor-like but pointed, and those on side conical. Suboperculum bears a sharp spine covered by skin extending from operculum. Gill membranes are united and free of isthmus. Second and third gill arches each have seven gill rakers. Anus is about mid-distant between anal fin origin and posterior margin of disc. Measurements are expressed as percent of SL: head length 41.7%– 45.5%, head width 37%– 41.7%, snout length 11.7%–14.5%, disc length 37.4%– 40%, body depth 17.5%–22.2%. Pectoral fin has 19 to 22 (usually 21) rays, and lower 5 to 7 rays are not reduced in size. Fleshy

pad on pectoral fin base is well developed and has free posterior margin extending from upper gill membrane attachment to lowest part of pectoral fin base. Dorsal fin has 11 or 12 (usually 11) rays, and origin is closer to upper pectoral fin base than to caudal fin base. Sucking disc is single and very large, and bears papillae on anterior, posterior, and paired middle sections. Anal fin has 7 to 9 (usually 8) rays. Caudal fin is broadly rounded. Color is gray to olive and stippled with small black spots. Occasionally specimens have bars on head. This species occurs in the western Atlantic from the northern Gulf of Mexico to northern South America. In the Gulf of Mexico it is recorded from the Texas coast, although Hoese and Moore (1998) consider the Texas records to be unverified. It is thought to be rare to absent off the Gulf coasts of Alabama and Florida, but there is one verified record from western Florida (TCWC 7302.07, 2517N, 8252W). This species is associated with limestone substrates in clear, shallow water. Maximum known size is 63 mm TL.

references: Briggs 1955; Robins and Ray 1986; Smith 1997; McEachran 2002b.

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GOBIESOCIDAE 645

Gobiesox strumosus Skilletfish

Cope, 1870

Gobiesox strumosus has a broad upper lip and a relatively large sucking disc. Head is broader than body, and snout is continuous with head. Upper lip is broadest near symphysis. Nares are paired and tubelike, with anterior naris possessing a bilobed flap on posterior margin, and posterior naris located just anterior to anterior margin of eye. Well-developed papillae occur on head and upper lip but not on snout above premaxillary groove. Teeth in front of upper jaw are conical, irregular in shape and size, and arranged in a patch. Teeth on side of upper jaw are conical and arranged in a single row. Teeth in front of lower jaw are arranged in two rows, with those in front incisor-like and bearing trifid tips in juveniles and rounded tips in adults. Teeth on side of lower jaw are conical. Suboperculum bears a sharp spine covered by skin extending from operculum. Gill membranes are united and are free of isthmus. Second and third gill arches each have six to nine gill rakers. Anus is closer to anal fin origin than to posterior margin of disc. Measurements are expressed as percent of SL: head length 38.5%– 43.5%, head width 37%– 41.7%, snout length 10.8%–13.1%, disc length 30.9%–34.3%, body depth 15.9%–21.3%.

Pectoral fin has 22 to 26 (usually 24) rays, and lower 5 to 7 rays are not reduced in size. Fleshy pad on pectoral fin base is well developed and has a free posterior margin extending from upper gill membrane attachment to lowest part of pectoral fin base. Dorsal fin has 10 to 13 (usually 12) rays, and origin is closer to upper pectoral fin base than to caudal fin base. Sucking disc is single and very large and bears papillae on anterior, posterior, and paired middle sections. Anal fin has 9 to 11 rays. Caudal fin is broadly rounded. Color is dark olive brown, with six faint lines radiating from eye and a mottled, netlike pattern and a dark band at base of caudal peduncle. This species occurs in the western North Atlantic from New Jersey and Bermuda to the southern Gulf of Mexico. It is found throughout the Gulf of Mexico in shallow water, including grassy areas and rocky tide pools. Maximum known size is 80 mm SL. references: Briggs 1955; Springer and Woodburn 1960; Hoese and Moore 1977, 1998; Johnson and Greenfield 1983; Robins and Ray 1996; Boschung 1992; Schaldach et al. 1997; Smith 1997; McEachran 2002b.

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646

CALLIONYMIDAE

Dragonets

Dragonets are elongate and moderately depressed, with a broad depressed head, widely separated pelvic fins, and no scales. Head is little to moderately elevated. Nostrils are undivided. Eye is moderate to large, and high on head. Mouth is terminal, and upper jaw is very protrusible. Jaw teeth are villiform, and teeth are absent in vomer and palatine. Preoperculum has a stout and variably shaped spine. Operculum and suboperculum lack spines. Gill opening is restricted to a small dorsal or sublateral pore. Gill rakers are few or absent. Pectoral fin inserts low on flank and is large and fanlike. Dorsal fin is divided into separate spinous and rayed fins, with first fin consisting of three to five weak spines and second fin consisting of 7 to 10 rays. In males, first dorsal fin is usually high and colorful. Pelvic fin is large, has one spine and 5 rays, and inserts anterior to pectoral fin base, with inner ray attached to body anterior to pectoral fin base. Anal fin consists of 4 to 10 rays. Caudal fin is generally greatly expanded and of variable shape, with central rays elongated as filaments in males. Lateral line is complete. Color ranges from drab to bright, with spots and stripes. Maximum known size is about 250 mm TL. Dragonets occur worldwide in tropical to temperate seas from the shoreline to 800 m. Eggs, larvae, and postlarvae are pelagic, and adults are benthic. Adults are often associated with sandy to muddy substrates and sea-grass beds, but some species occur on coral reefs. There are 125 species in nine genera worldwide, 5 species in three genera in the western Central Atlantic, and 4 species in three genera in the Gulf of Mexico. references: Fricke 1981, 1986, 2001; Nakabo 1982b; Nelson 1994; Smith 1997; Smith-Vaniz et al. 1999; Hartel and Nakabo 2002a.

Key to the Species of the Gulf of Mexico (Adapted from Hartel and Nakabo 2002a) 1a. Ventrolateral fold from belly to caudal fin base; anal fin rays 4 . . . . . . . . . . . . . Diplogrammus pauciradiatus p. 648 1b. No ventrolateral fold from belly to caudal fin base; anal fin rays 7 or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Preopercular spine with antrorse ventral barb and several upwardly directed barbs . . . . . . . . . . . . . . . . . . . . Paradiplogrammus bairdi p. 651 2b. Preopercular spine without antrorse ventral barb and only two upwardly directed barbs at posterior preopercular tip .............................................3 3a. First dorsal fin spine elongate and filamentous; second dorsal fin slightly emarginate; black mark on first dorsal fin

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CALLIONYMIDAE 647

small or absent; anal fin without blackish band in males . . . . . . . . . . . . . . . . . . . . . Foetorepus goodenbeani p. 650 3b. First dorsal fin spine not elongate and filamentous; second dorsal fin convex (males) or nearly straight (females); black mark on first dorsal fin large; anal fin with blackish band in males . . . . . . . . . . . . . . . . . . Foetorepus agassizii p. 649

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648 PERCIFORMES

Diplogrammus pauciradiatus Spotted dragonet

(Gill, 1865)

Diplogrammus pauciradiatus has head and anterior part of trunk laterally expanded, preopercular spine with three dorsally projecting barbs, and ventrolateral keel extending from belly to caudal fin base. Snout is short and triangular in dorsal view. Jaw teeth are villiform and arranged in bands. Eye is very large and directed dorsolaterally. Preopercular spine is broad based, lacks antrorse spine or hook on base, is upturned near tip, and bears three triangular barbs. Gill opening is oval and dorsolateral in position. Measurements are expressed as percent of SL: head length 25.8%– 30.8%, snout length 8.4%–9.6%, eye diameter 6.4%– 8.1%, upper jaw length 7.3%– 7.7%, pre-dorsal-fin length 27.5%–30.5%, body depth 12%–15.4%. Pectoral fin has 16 to 19 rays. First dorsal fin has four spines that are very long and terminate as filaments in males. Second dorsal fin has 6 rays, and all but first are branched, with last ray divided at base. Pelvic fin is connected to pectoral fin by broad membrane. Anal fin has 4 rays, and only last is branched and divided at base. Caudal fin is long. Lateral line is single and complete. Color is mottled tan, with reticulations ven-

trally. A row of dark spots occurs on fleshy keel along side. Most males have a dark bar across mouth and a dark diagonal bar across eye. Some large males have orange pigment on anterior part of head and around eye. This species occurs in shallow water in the western Atlantic from North Carolina and Bermuda to Colombia, including the eastern and southern Gulf of Mexico and the Bahamas. It is associated with sea-grass beds. Feeding takes place during the day, and food consists of harpacticoid copepods. At night, individuals bury themselves in the substratum or rest on the substratum. Spawning takes place in the water column, about 30 cm from the bottom, over open sand. Maximum known size is 43 mm SL for males and 27 mm SL for females.

references: Jordan and Evermann 1898; Springer and Woodburn 1960 (both as Callionymus calliurus); Davis 1966; Sogard 1984; Robins and Ray 1986; Boschung 1992; Harrington 1997; Smith 1997; Smith-Vaniz et al. 1999; Hartel and Nakabo 2002a.

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CALLIONYMIDAE 649

Foetorepus agassizii Spotfin dragonet

(Goode and Bean, 1888)

Foetorepus agassizii has a slightly depressed head, a preopercular spine with two barbs at tip, and a dark spot on first dorsal fin. Snout is short and triangular in dorsal view. Jaw teeth are villiform and arranged in broad bands. Eye is very large and directed dorsolaterally. Preopercular spine has a smooth base, lacks an antrorse spine or hook on base, is upturned near tip, and bears two dorsal barbs. Gill opening is small, oval, and dorsolateral in position. Measurements are expressed as percent of SL: head length 27.8%–29.8% in males and 25.8%–30.6% in females, snout length 7.7%– 8.6% in males and 7.8%– 8.7% in females, eye diameter 9%–10.6% in males and 9%–11.1% in females, upper jaw length 8.1%– 8.8% in males and 8.3%– 8.9% in females, pre-dorsalfin length 31.2%–32.9% in males and 33.1%– 33.6% in females, body depth 11.6%–14.2% in males and 12.5%–14.6% in females. Pectoral fin has 20 to 23 rays (usually 2 simple rays and 18 to 21 long, branched rays). First dorsal fin has four spines, with first just slightly longer than second and lacking long filament. Second dorsal fin is convex in males and nearly straight in females and has 8 branched rays, with last ray divided to base. Pelvic fin is con-

nected to pectoral fin by broad membrane. Anal fin has 7 rays, and only last is branched and divided to base. Caudal fin is diamond shaped, with middle 2 rays extending as filaments in males. Lateral line is single and complete. In life, upper body and head are carmine red, with dark red markings on body. Belly is white. First dorsal fin has a jet black spot between third and fourth spines, and anal fin has a black band near margin. In preservative, body and head are yellow, with a dark spot on dorsal fin and a dark band on anal fin. This species occurs in the western Atlantic from Canada to northern Brazil, including the eastern, northwestern, and southern Gulf of Mexico and the Antilles, between 91 and 650 m. It is absent along the coast of Brazil between 5N and 20S. Maximum known size is 250 mm TL.

references: Jordan and Evermann 1898 (as Callionymus himantophorus); Fricke 1981; Robins and Ray 1986; Boschung 1992; Smith 1997 (last four references as Foetorepus agassizii, in part); Nakabo and Hartel 1999; Hartel and Nakabo 2002a.

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650 PERCIFORMES

Foetorepus goodenbeani

Nakabo and Hartel, 1999

Foetorepus goodenbeani has a slightly depressed head, a preopercular spine with two barbs at tip, and a shallowly emarginate second dorsal fin. Snout is short and triangular in dorsal view. Eye is very large and directed dorsolaterally. Maxilla extends almost to anterior margin of eye. Jaw teeth are villiform and arranged in broad bands. Occipital region, behind eye, bears very low bony protuberances. Preopercular spine has a smooth base, lacks an antrorse spine or hook on base, is upturned near tip, and bears two dorsal barbs. Gill opening is small, oval, and dorsolateral in position. Measurements are expressed as percent of SL: head length 24.5%–27.7% in males and 25.4%–34.7% in females, snout length 7.5%–10.8%, eye diameter 6.9%–10%, upper jaw length 7.8%–9%, pre-dorsal-fin length 26.3%–30.7% in males and 28.8%–29.5% in females, body depth 11.1%–13.7% in males and 12.5%–14% in females. Pectoral fin usually has 1 simple ray and 19 to 22 branched rays. First dorsal fin has four spines, with first spine about 2 times longer than second

spine. Second dorsal fin has 8 rays, and all are branched, with last ray divided to base and elongated in males. Pelvic fin is connected to pectoral fin by broad membrane. Anal fin has 7 rays, and only last is branched and divided to base. Caudal fin is emarginate dorsally and round ventrally, with middle 2 rays very long in males. Lateral line runs from preorbital region to caudal fin base. In life, body and head are red dorsally, with faint marbled olive brown marks, and white ventrally, with two wavy yellow lines on side. First dorsal fin is dark yellow. In preservative, body and head are tan, with blotches dorsally, and whitish yellow ventrally. This species occurs between 47 and 396 m in the western Atlantic from Georges Bank off Massachusetts to the northern Gulf of Mexico off Texas. It is most common between 100 and 200 m. Maximum known size is 300 mm TL. references: Longley and Hildebrand 1941 (as F. agassizii); Davis 1966 (as F. agassizii, in part); Nakabo and Hartel 1999; Hartel and Nakabo 2002a.

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CALLIONYMIDAE 651

Paradiplogrammus bairdi (Jordan, 1888) Lancer dragonet

Paradiplogrammus bairdi has a moderately depressed head, a forwardly projecting barb on underside and series of barbs on upper side of preopercular spine, and a mottled body. Snout is rather long and triangular in dorsal view. Jaw teeth are minute and arranged in broad bands. Eye is very large and directed laterally. Preopercular spine has an antrorse barb or hook on base and three to nine barbs on upper side. Gill opening is slitlike and sublateral in position. Measurements are expressed as percent of SL: head length 24.1%–28.7%, snout length 8.7%–9%, eye diameter 7.1%– 8%, upper jaw length 8.1%–10%, pre-dorsal-fin length about 34%, body depth 12.1%–15%. Pectoral fin has 21 rays. First dorsal fin has four spines and is considerably higher than second dorsal fin in males, with a well-developed membrane behind fourth spine. Second dorsal fin has 9 rays,

with only last one branched and divided to base. Pelvic fin is connected to pectoral fin by a broad membrane. Anal fin has 8 rays, and only last is branched and divided to base. Caudal fin is truncate to slightly rounded. Lateral line is single and complete. Body is light gray with red mottling. Males have a yellow spinous dorsal fin and dark bars on cheek. This species occurs in the western Atlantic from Cape Hatteras and Bermuda to northern South America, including the northern and southern Gulf of Mexico and the Bahamas. It is associated with sand around reefs to 91 m. Maximum known size is 114 mm TL. references: Jordan and Evermann 1898 (as Callionymus bairdi); Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997 (as C. bairdi); SmithVaniz et al. 1999; Hartel and Nakabo 2002a.

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652

DRACONETTIDAE

Draconetts

Draconettids are elongate and slender, depressed anteriorly, and compressed posteriorly, with strong, pointed antrorse spines on operculum and suboperculum, two dorsal fins, and no scales. Head is large, and snout is acutely pointed both laterally and vertically. Nostrils are unpaired and terminate as a tube near tip of snout. Eye is large and located high on head. Mouth is terminal, and upper jaw is very protrusible. Jaw teeth are small, and vomer and palatine lack teeth. Preoperculum lacks spines. Gill opening is relatively broad, extending from near upper margin of pectoral fin base to just above pelvic fin base. Pectoral fin inserts on midflank and is broad based. First dorsal fin has three spines, and second dorsal fin has 12 to 15 rays. Pelvic fin inserts anterior to posterior margin of operculum and consists of one spine and 5 rays. Anal fin consists of 12 or 13 rays. Caudal fin is generally truncate or convex. Lateral line is confined to head. Vertebrae number 23. Draconettids occur in the western Atlantic and western and Central Pacific, off Japan and Hawaii, along the outer continental and insular shelves and upper slopes. There are 12 species in a single genus, 2 species in the western Central Atlantic, and 1 species in the Gulf of Mexico. references: Briggs and Berry 1959; Nakabo 1982a; Fricke 1992; Nelson 1994; Hartel and Nakabo 2002b.

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DRACONETTIDAE 653

Centrodraco acanthopoma

(Regan, 1904)

Centrodraco acanthopoma has a relatively broad opercular opening and spines on operculum and suboperculum. Upper jaw projects beyond lower jaw and extends to anterior margin of orbit. Jaw teeth are arranged in narrow bands. Gill opening extends along length of pectoral fin base. Measurements are expressed as percent of SL: head length 32.4%– 36.8%, snout length 7.7%–9.4%, eye diameter 11.4%–15.2%, upper jaw length 8.5%– 10%, first dorsal spine length 4.9%– 6.4%, body depth at pelvic fin origin 12.6%–16.9%; caudal peduncle depth 6.3%–7.4%. Pectoral fin is acutely pointed and has 25 or 26 rays. First dorsal fin has three spines, and second dorsal fin has 13 to 15 rays. Anal fin has 13 rays. Caudal fin is truncate and has 13 segmented rays. Color is light, with dark round

spots and irregular blotches on side of body, a large dark blotch on operculum, and a large round blotch on caudal fin base. This species occurs in the western Atlantic from Massachusetts to Venezuela, including the Florida Keys and the southern Gulf of Mexico. There is one record from the southern Gulf of Mexico (2043N, 8651W). Postlarvae and small juveniles are mesopelagic between 200 and 400 m. Records from north of Cape Hatteras are postlarvae and small juveniles (15 to 28 mm SL) captured at mesopelagic depths. Known depth range is 210 to 300 m. Maximum known size is 110 cm TL.

references: Briggs and Berry 1959; Davis 1966; Nakabo 1982a; Hartel and Nakabo 2002b.

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654

ELEOTRIDAE

Sleepers

Sleepers are moderately elongate and moderately stout, with two dorsal fins and thoracic pelvic fins that are either close together or united at bases but distally separate. Head is moderately elevated, short, broad, and usually flat on top, with eyes widely separated. Mouth is terminal to superior and rather large. Jaw teeth are small, conical, and arranged in several rows. Gill membranes are broadly joined to isthmus. Branchiostegal rays number 6. Pectoral fin is fanlike and consists of 14 to 25 rays. First dorsal fin consists of 2 to 8 weak spines, and second consists of 1 weak spine and 6 to 15 rays. Base length of second dorsal fin is less than distance between its insertion and caudal fin base. Pelvic fin consists of 1 spine and 5 rays. Anal fin originates posterior to origin of second dorsal fin and has 1 weak spine and 6 to 12 rays. Caudal fin has 15 to 17 segmented rays. Body and most of head are covered with cycloid or ctenoid scales. Lateral line canal is absent on body, but canals and pores are present on head. Maximum known size is about 60 cm TL. Sleepers occur worldwide in tropical and subtropical regions. Most species are benthic, although some freely swim through the water column, and they are generally associated with fresh and brackish waters, but some are found on coral reefs. There are approximately 150 species in about 40 genera worldwide, 10 species in 5 genera in the western Central Atlantic, and 5 species in 5 genera in the Gulf of Mexico. references: Fritzsche and Fuiman 1982; Hoese 1986b; Nelson 1994; Smith 1997; Larson and Murdy 2001a; Murdy and Hoese 2002a.

Key to the Species of the Gulf of Mexico (Adapted from Murdy and Hoese 2002a) 1a. Ventrally directed spine covered with skin on preoperculum .............................................2 1b. Spine absent on preoperculum . . . . . . . . . . . . . . . . . . . . . 3 2a. Scales are cycloid and number 90 to 110 in horizontal series on body; caudal fin with procurrent rays; body slender, elongate, and terete. . . . . . . Erotelis smaragdus p. 658 2b. Scales on most of body are ctenoid and number 40 to 58 in horizontal series on body; caudal fin without procurrent rays; body not slender. . . . . . . . Eleotris amblyopsis p. 657 3a. First dorsal fin with six spines; body with about 40 to 65 horizontal scale rows; body and head strongly compressed . . . . . . . . . . . . . . . . . . . . . . . Gobiomorus dormitor p. 659 3b. First dorsal fin with seven spines; body with fewer than 40

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ELEOTRIDAE 655

or more than 90 horizontal scale rows; body and head not compressed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4a. Horizontal scale rows 100 to 110 . . . . . . . . . . . . . . . . . . . . . . . . . . . Guavina guavina p. 660 4b. Horizontal scale rows about 25 to 35 . . . . . . . . . . . . . . . . . . . . . . . Dormitator maculatus p. 656

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656 PERCIFORMES

Dormitator maculatus Fat sleeper

(Bloch, 1792)

Dormitator maculatus has a smooth preopercular margin and a fully scaled head. Head is broad and flat dorsally. Lips are thin, upper lip is continuous across symphysis, and upper jaw is moderately protrusible. Mouth is oblique, and upper jaw fails to reach anterior margin of eye. Jaw teeth are arranged in several rows, with those of outer row slightly movable and slender. Teeth are present in vomer but absent in palatine. Tongue has a rounded margin. Preopercular margin is fleshy. Gill opening is slightly restricted dorsally but extends ventrally to below posterior margin of orbit. Measurements are expressed as percent of SL: head length 29%–33%, snout length 8%–9%, eye diameter 5%– 8%, interorbital width 11%– 14%, upper jaw length 8%–10%, pectoral fin length 24%–30%, body depth 27%–32%. Pectoral fin is broad and fanlike and has 14 rays. First dorsal fin has seven spines, and second has one spine and 8 rays. Pelvic fins are close set, and each has one spine and 5 rays. Anal fin has one spine and 9 rays. Caudal fin is rounded. Body and most of head, including entire cheek, are covered with scales; those on

head, nape, chest, and abdomen are cycloid, and the remainder are ctenoid. Head scales are arranged in a regular mosaic pattern. Scales in horizontal series number 25 to 35. Vertebrae number 27: 12 precaudal and 15 caudal. Color is dark brown to blackish, with a large bluish black blotch above base of pectoral fin and the outer margin of dorsal and anal fins dull red. Occasionally specimens are patterned with several series of horizontally arranged spots on body. This species occurs in the western Atlantic from Chesapeake Bay and the Bahamas to southeastern Brazil, including the northern and southern Gulf of Mexico and the Antilles. It is associated with fresh and brackish waters on soft substrates. Maximum known size is 380 mm TL. references: Springer and Woodburn 1960; Böhlke and Chaplin 1968; Dawson 1969b; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith 1997; Castro-Aguirre et al. 1999; Murdy and Hoese 2002a.

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ELEOTRIDAE 657

Eleotris amblyopsis Spinycheek sleeper

(Cope, 1871)

Eleotris amblyopsis has a superior mouth, a strong spine on preoperculum, and a robust caudal fin. Head is slightly laterally expanded and depressed, with profile from snout to origin of first dorsal fin more or less horizontal. Lips are thin, and upper lip is continuous across symphysis. Mouth is strongly oblique, and upper jaw extends to level of pupil. Jaw teeth are small and arranged in two bands, with those in inner row slightly enlarged. Tongue has a rounded margin. Angle of preoperculum bears a strong, ventrally directed spine that is embedded in integument. Gill opening is restricted dorsally but extends ventrally to below operculum. Measurements are expressed as percent of SL: head length 27%–36%, snout length 7%– 8%, eye diameter 4%–5%, interorbital width 8%–9%, upper jaw length 12%– 14%, pectoral fin length 19%–31%, caudal fin length 13%–33%, body depth 18%–21%. Pectoral fin is broad and fanlike, with 15 to 18 rays. First dorsal fin has six spines, and second has one spine and 7 to 9 rays. Pelvic fins are close set, and each has one spine and 5 rays. Anal fin is similar in shape and size to second dorsal fin and has one spine and 7 or 8 rays. Caudal fin is rounded. Body and most of head, including entire cheek, are covered with scales, with those on head, nape, chest, and abdo-

men cycloid and the remainder ctenoid. Scales in horizontal series number 40 to 58 (generally 50 to 56 in specimens from North America), predorsal scales number 29 to 46, and transverse scale rows number 11 to 18. Vertebrae number 25: 10 precaudal and 15 caudal. Color is tan to brown, with minute dark spots and ventral surface lighter than dorsal surface. Cheek has two dark streaks radiating posteriorly from eye, and pectoral fin base has a large spot. This species occurs in the western Atlantic from North Carolina and Bermuda to southern Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. Adults are associated with fresh and brackish waters on soft substrates, and juveniles occur along coasts. Maximum known size is 250 mm SL. references: Böhlke and Chaplin 1968; Dawson 1969b; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997 (as E. abacurus in part); Smith 1997; Castro-Aguirre et al. 1999 (as E. abacurus in part); Smith-Vaniz et al. 1999 (all previous references, except Boschung, Schaldach et al., and CastroAguirre et al., as E. pisonis or in part as E. pisonis); Murdy and Hoese 2002a; Pezold and Cage 2002.

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658 PERCIFORMES

Erotelis smaragdus Emerald sleeper

(Valenciennes, 1837)

Erotelis smaragdus has a superior mouth, skin displaying a myomere pattern, and a robust caudal fin. Head is slightly laterally expanded and depressed, with lateral profile from snout to origin of first dorsal fin nearly horizontal. Lips are thin, and upper lip is continuous across symphysis. Mouth is strongly oblique, and upper jaw extends to about level of pupil. Jaw teeth are small and arranged in bands, with outer row slightly enlarged. Tongue has a rounded margin. Preoperculum has ventrally directed spine. Gill opening is restricted dorsally but extends ventrally to below operculum. Measurements are expressed as percent of SL: head length 26%–27%, snout length 4%–5%, eye diameter about 3%, interorbital width 3%– 4%, upper jaw length 7%– 8%, pectoral fin length 17%–20%, caudal fin length 25%– 28%, body depth 12%–13%. Pectoral fin is broad and fanlike, with 17 rays. First dorsal fin has six spines, and second has one spine and 11 rays. Pelvic fins are close set, and each has one spine and 5 rays. Anal fin has one spine and 9 rays. Caudal fin extends dorsally far anterior of hypural plate and is narrowly

rounded. Body, top of head, cheeks, and operculum are covered with cycloid scales, and those on head are partially embedded. Scales in horizontal series number 90 to 110. Vertebrae number 25: 10 precaudal and 15 caudal. Color is blackish brown with a greenish sheen dorsally and tan ventrally. Upper section of pectoral fin base has a dark spot, and dorsal, anal, and caudal fins are light tan. In life, lateral body surface has a series of chevronshaped markings, and dorsal, anal, and caudal fins have narrow brown bands. This species occurs in the western Atlantic from southeastern Florida and the Bahamas to Brazil, including the Gulf of Mexico and the Antilles. It is rare in the northern Gulf of Mexico. This species is associated with coastal waters on sandy marl to silty bottoms and apparently burrows into the substratum. Maximum known size is 200 mm SL. references: Böhlke and Chaplin 1968; Dawson 1969b; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Smith 1997; CastroAguirre et al. 1999; Murdy and Hoese 2002a.

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ELEOTRIDAE 659

Gobiomorus dormitor Bigmouth sleeper

Lacepède, 1800

Gobiomorus dormitor has a large terminal mouth, a smooth preopercular margin, and a broad gill opening. Head is rather narrow and flattened dorsally. Lips are thin, and upper lip is continuous across symphysis. Upper jaw is moderately protrusible. Mouth is moderately oblique, with lower jaw projecting beyond upper jaw and upper jaw extending to anterior margin of eye. Jaw teeth are arranged in bands, and teeth are slender and sharp pointed. Teeth are present in vomer but absent in palatine. Tongue is emarginate. Preopercular margin is fleshy. Gill opening is slightly restricted dorsally but extends ventrally to below level of eye. Measurements are expressed as percent of SL: head length 29%–30%, snout length 8%–9%, eye diameter 4%– 6%, interorbital width 4%– 6%, upper jaw length 9%–12%, pectoral fin length 19%–21%, body depth 13%–16%. Pectoral fin is broad and fanlike, with 16 or 17 rays. First dorsal fin has six spines, and second has one spine and 9 rays. Pelvic fins are close set, and each has one spine and 5 rays.

Anal fin has one spine and 9 rays. Caudal fin is rounded. Body and most of head, including entire cheek, are covered with ctenoid scales. Scales in horizontal series number 45 to 65. Vertebrae number 26: 12 or 13 precaudal and 13 or 14 caudal. Color is dark brown to olive brown dorsally and tan to yellowish tan on sides and ventrally, with dark lines on cheek and operculum and a dark brown to olive brown stripe between pectoral fin base and caudal peduncle. This species occurs in the western Atlantic from southern Florida and the Bahamas to eastern Brazil, including the Gulf of Mexico. In the Gulf of Mexico it occurs along the coasts of southern Texas, southern Florida, Veracruz, Tabasco, and Campeche. It is associated with fresh and brackish waters on soft substrates. Maximum known size is 600 mm SL. references: Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith 1997; Castro-Aguirre et al. 1999; Murdy and Hoese 2002a.

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660 PERCIFORMES

Guavina guavina

(Valenciennes, 1837)

Guavina gauvina has a superior mouth, a robust caudal fin, and small scales. Head is depressed and slightly elevated. Lips are slightly fleshy, and upper lip is continuous across symphysis. Mouth is moderately oblique, and upper jaw extends to pupil of eye. Jaw teeth are moderately small and arranged in broad bands, with outer teeth in lower jaw slightly enlarged. Tongue has a rounded margin. Preoperculum lacks spine. Gill opening is restricted dorsally but extends ventrally to below preoperculum. Measurements are expressed as percent of SL: head length 29%–30%, snout length 7%– 8%, eye diameter about 4%, interorbital width 9%–10%, upper jaw length 12%–13%, pectoral fin length 18%–19%, caudal fin length 21%–22%, body depth 20%–21%. Pectoral fin is broad and fanlike, with 16 rays. First dorsal fin has seven (rarely six) spines, and second has one spine and 9 rays. Pelvic fins are close set, and each has one spine and 5 rays. Anal fin is similar in shape and size to second dorsal fin and has one spine and 8 or 9 rays. Caudal fin is

broadly rounded. Body and most of head— including dorsal aspect, entire cheek, and operculum—are covered with scales; those on head and upper and lower surfaces of body are cycloid, and the remainder are ctenoid. Head scales are embedded. Scales in horizontal series number 100 to 110. Vertebrae number 27: 11 precaudal and 16 caudal. Color is dark brown to grayish brown and slightly lighter ventrally than dorsally. This species occurs in the western Atlantic from the southern Gulf of Mexico and the Antilles to Brazil in freshwater with access to the sea and in brackish to hypersaline water. In the Gulf of Mexico it has been recorded from Veracruz at Lago de Sontecomapan. Maximum known size is about 300 mm SL. references: Jordan and Evermann 1898; Meek and Hildebrand 1923 –1928; Taylor 1996; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Murdy and Hoese 2002a.

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661

GOBIIDAE

Gobies

Gobies are elongate and subcylindrical anteriorly and compressed posteriorly, with eyes located on dorsolateral aspect of head, prominent cheeks, and pelvic fins usually united to form a disc. Head is short and broad, and snout is rounded. Nostril is paired; anterior naris consists of a tube, and posterior naris is generally porelike and surrounded by a raised rim. Mouth is terminal to subterminal. Jaw teeth are small, conical, and arranged in one or more rows, often with some teeth enlarged, recurved, and caninelike. Vomerine teeth usually are absent but are present in some species. Gill opening is generally restricted both dorsally and ventrally and is broadly joined to isthmus. Branchiostegal rays number 5. Pectoral fin is well developed and consists of 12 to 60 rays (14 to 22 rays in western Atlantic species). Dorsal fin is generally divided, with first (when present) consisting of 2 to 8 flexible spines and second consisting of 0 or 1 flexible spine and 5 to 37 rays. Western Atlantic species have 4 to 8 weak spines in first dorsal fin and 1 weak spine and 9 to 18 rays in second dorsal fin. Second dorsal fin base is subequal to or longer than distance from insertion to caudal fin base. Pelvic fins are completely separated, partially connected, or connected and forming disc. Spine of pelvic fin is displaced forward and ventrally and is not in line with rays. Often membrane connects adjacent pelvic fin spines to form pelvic frenum, and frenum can vary from delicate to thick and fleshy. Anal fin is generally similar in shape to second dorsal fin and consists of 0 to 1 flexible spine and a variable number of rays. Western Atlantic species have 1 weak spine and 9 to 18 rays in anal fin. Terminal rays of second dorsal fin and anal fin are divided to base but counted as a single ray. Caudal fin varies from rounded to lanceolate and has 16 or 17 segmented rays. Body is naked, or partially or fully covered with cycloid or ctenoid scales. Lateral line canals are absent on body but present on head, generally in supraorbital, postocular, and preopercular regions. Many species have cutaneous papillae on head. The majority of the species are less than 100 mm SL, but one reaches 800 mm SL. Gobies occur worldwide in tropical, subtropical, and warm temperate regions. Most are benthic in habitat, but some are freeswimming and schooling species. Most species are marine between the shoreline and 500 m, but a number inhabit estuaries and freshwater. They are especially diverse on coral reefs. A number of species are symbiotic with other marine organisms. Freshwater species inhabit mountain streams to altitudes of 2,000 m on oceanic islands. There are about 1,875 species in 220 genera, 124 species in 33 genera in the western Central Atlantic, and 58 to 62 species in 26 genera in the Gulf of Mexico.

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662 PERCIFORMES

references: Fritzsche and Fuiman 1982; Hoese 1986a; Miller 1986; Birdsong et al. 1988; Nelson 1994; Larson and Murdy 2001b; Murdy and Hoese 2002b.

Key to the Species of the Gulf of Mexico (Adapted from Murdy and Hoese 2002b) 1a. Dorsal and anal fins connected to caudal fin; a single dorsal fin; body very elongate . . . . Gobioides broussonetii p. 704 1b. Dorsal and anal fins separated from caudal fin; two dorsal fins; body robust or elongate . . . . . . . . . . . . . . . . . . . . . . 2 2a. Lower jaw teeth incisor-like, typically in a single row; pelvic fins mostly fused to belly by membrane . . . . . . . . . . . . . . . . . . . . . . . Sicydium gymnogaster p. 728 2b. Lower jaw teeth conical or compressed, typically in more than one row; pelvic fins largely free of belly . . . . . . . . . . 3 3a. Pair of pores (anterior interorbital pores) above anterodorsal margin of eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3b. Usually a single pore (anterior interorbital pore) above anterodorsal margin of eyes, or head pores absent . . . . . 16 4a. Low membranous crest on nape from above preoperculum to first dorsal fin origin . . . . . . . . . . . . . . . . . . Oxyurichthys stigmalophius p. 723 4b. No membranous crest on nape. . . . . . . . . . . . . . . . . . . . . 5 5a. Shoulder girdle, under operculum, with distinct fleshy lobes. . . . . . . . . . . . . . . . . . . . . . . . Awaous banana p. 668 5b. Shoulder girdle, under operculum, without fleshy lobes .............................................6 6a. Teeth compressed, with bilobed tips; mouth slightly inferior . . . . . . . . . . . . . . . . . . . . . . . . . Evorthodus lyricus p. 702 6b. Teeth conical, pointed; mouth terminal or inferior . . . . . . 7 7a. Tongue distinctly bilobed; sides of head scaled to below eye; mouth inferior . . . . . . . . . . Gnatholepis thompsoni p. 703 7b. Tongue pointed or rounded, not bilobed; sides of head without scales, but scales may occur on operculum; mouth terminal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8a. Lateral cephalic canal with four pores; first gill arch with numerous gill rakers on upper and lower limbs . . . . . . . . . . . . . . . . . . . . . . . Gobionellus oceanicus p. 705 8b. Lateral cephalic canal with two pores; first gill arch with no gill rakers on upper limb and four or five on lower limb .............................................9 9a. Second dorsal fin and anal fin with 11 and 12 elements, respectively. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9b. Second dorsal fin and anal fin with 12 and 13 rays, respectively. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10a. Side of head patterned with black circles; many green spots on side of body. . . . . . . . . . Ctenogobius smaragdus p. 692 10b. Side of head not patterned with black circles; about five

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GOBIIDAE 663

round to elongate dark blotches along midflank of body, some with diagonal marks extending dorsally to form V shapes . . . . . . . . . . . . . . . . Ctenogobius boleosoma p. 687 11a. Margin of preoperculum darkly pigmented . . . . . . . . . . 12 11b. Margin of preoperculum no darker than remainder of head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 12a. Nape with 10 to 12 predorsal scales; spines of first dorsal fin not produced . . . . . . . . Ctenogobius stigmaturus p. 694 12b. Nape with few or no predorsal scales; third spine of first dorsal fin often greatly produced in males . . . . . . . . . . . . . . . . . . . . . . . Ctenogobius fasciatus p. 689 13a. Cheek with three broad, dark vertical bars; middle of lower jaw with laterally or horizontally projecting tusklike canine tooth . . . . . . . . . . . Ctenogobius stigmaticus p. 693 13b. Cheek without three broad, dark vertical bars; lower jaw with or without canine tooth, but if present, tooth not projecting laterally or horizontally from mouth . . . . . . . . . . 14 14a. Cheek with dominant suborbital bar extending from lower rim of orbit to corner of jaw (may not reach corner of jaw) . . . . . . . . . . . . . . . . . . . . Ctenogobius saepepallens p. 690 14b. Cheek with dominant horizontal bar across midcheek from upper preopercular canal to corner of jaw or with dominant streak on snout from eye to midlateral portion of upper jaw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 15a. Pelvic fin in adult males dusky; pelvic fin in females with bilateral streaks paralleling innermost ray running posteriorly from fin base; adult males typically with third dorsal fin spine elongate. . . . . . . . . . Ctenogobius claytonii p. 688 15b. Pelvic fin in adult males with bilateral streaks paralleling innermost ray running posteriorly from fin base; pelvic fin in females without streaks; adult males lack elongate third spine in first dorsal fin . . . . . . Ctenogobius shufeldti p. 691 16a. First dorsal fin with two to six spines . . . . . . . . . . . . . . . 17 16b. First dorsal fin with seven or eight spines . . . . . . . . . . . . 35 17a. Second dorsal fin with more than 20 elements; pelvic fins separate, with one spine and 4 rays . . . . . . . . . . . . . . . . . . . . . . . . . Ptereleotris calliura p. 726 17b. Second dorsal fin with fewer than 20 elements; pelvic fins separate or connected by membrane, with one spine and 5 rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 18a. No head pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 18b. Head pores present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 19a. Body nearly naked to naked, few scales just anterior to caudal fin base or none; body depth 11%–14.3% of SL; second dorsal fin with one spine and 10 to 15 rays . . . . . . . . . . . . . . . . . . Evermannichthys spongicola p. 701 19b. Body fully scaled, scales reaching anteriorly to at least first dorsal fin; body depth 14.3%–25% of SL; second dorsal fin with one spine and 8 to 11 rays. . . . . . . . . . . . . . . . . 20

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664 PERCIFORMES

20a. Top of head with scales to posterior margin of eye; gill opening extending to below posterior preopercular margin; spines of pelvic fins not connected by membrane . . . . . . . . . . . . . . . . . . . . . . . . . . . Priolepis hipoliti p. 725 20b. Top of head naked; gill opening not extending to below posterior preopercular margin; spines of pelvic fins connected by membrane and forming cup-shaped disc. . . . . 21 21a. Body usually uniformly pigmented, lacking bands, bars, or stripes . . . . . . . . . . . . . . . . . . . . Lythrypnus elasson p. 714 21b. Body with bands, bars, and/or stripes . . . . . . . . . . . . . . 22 22a. Pale bands on body with dark center lines . . . . . . . . . . . 23 22b. Pale bands on body, if present, without dark center lines, but blue bands on body with darker center line . . . . . . . . . . . . . . . . . . . . . . . . . . Lythrypnus spilus p. 717 23a. Dark bands on body divided by pale central areas; pectoral fin base with two spots, one ventral to other (occasionally barely separated) . . . . . . . . . . Lythrypnus phorellus p. 716 23b. Dark bands on body not divided by pale central areas; pectoral fin base with one or two spots . . . . . . . . . . . . . . . . . . . . . . . . Lythrypnus nesiotes p. 715 24a. Upper three or four pectoral fin rays filamentous and free of membrane; scales extending onto top of head . . . . . . 25 24b. No free pectoral fin rays; scales not extending onto top of head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 25a. Lateral scale series, from upper opercular margin to last scale overlapping central caudal fin rays, 36 or fewer. . . 26 25b. Lateral scale series, from upper opercular margin to last scale overlapping central caudal fin rays, 37 to 44 . . . . . . . . . . . . . . . . . . . . . . Bathygobius soporator p. 672 26a. Pectoral fin rays 19 or 20; scales in horizontal series 33 to 36 (usually 35) . . . . . . . . . . Bathygobius mystacium p. 671 26b. Pectoral fin rays 16 or 17; scales in horizontal series 31 to 34 (usually 33) . . . . . . . . . . . . Bathygobius curacao p. 670 27a. Prominent crest extending from between eyes to origin of dorsal fin . . . . . . . . . . . . . Lophogobius cyprinoides p. 713 27b. No crest but often low, fleshy ridge extending from between eyes to origin of dorsal fin . . . . . . . . . . . . . . . . 28 28a. Pelvic fins forming circular disc . . . . . . . . . . . . . . . . . . . 29 28b. Pelvic fins forming emarginate disc. . . . . . . . . . . . . . . . . 30 29a. Prominent dark spot on lower half of pectoral fin base . . . . . . . . . . . . . Coryphopterus punctipectophorus p. 685 29b. No spot on lower half of pectoral fin base . . . . . . . . . . . . . . . . Coryphopterus glaucofraenum p. 681 30a. Pelvic frenum (membrane connecting two pelvic fin spines) present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 30b. Pelvic frenum absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 31a. Upper part of pectoral fin base with dark spot . . . . . . . . . . . . . . . . . . . . . . . . Coryphopterus thrix p. 686

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GOBIIDAE 665

31b. Upper part of pectoral fin base without dark spot . . . . . . . . . . . . . . . . . . . . . . Coryphopterus eidolon p. 680 32a. Pelvic fins separate or nearly separate. . . . . . . . . . . . . . . 33 32b. Pelvic fins united . . . . . . . . . . Coryphopterus dicrus p. 679 33a. Three interorbital pores between eyes . . . . . . . . . . . . . . . . . . . . . Coryphopterus hyalinus p. 682 33b. Two interorbital pores between eyes. . . . . . . . . . . . . . . . 34 34a. Second dorsal fin and anal fin each with 11 elements; no stripes behind eye. . . . . . Coryphopterus personatus p. 684 34b. Second dorsal fin and anal fin each with 10 elements; several stripes behind eye. . . . . Coryphopterus lipernes p. 683 35a. Head with three or more pairs of barbels; body without scales . . . . . . . . . . . . . . . . . . . Barbulifer ceuthoecus p. 669 35b. Head without elongate barbels but may have one or two pairs of short bumps; body with or without scales . . . . . 36 36a. Pelvic fins completely separate . . . . . . . . . . . . . . . . . . . . 37 36b. Pelvic fins connected by membrane . . . . . . . . . . . . . . . . 39 37a. Pelvic fin rays unbranched but with expanded tips; tongue bilobed . . . . . . . . . . . . . . . . . . . . Varicus marilynae p. 729 37b. Pelvic fin rays branched and without expanded tips; tip of tongue rounded or truncate . . . . . . . . . . . . . . . . . . . . . . 38 38a. Second dorsal fin with 10 elements; anal fin with 9 elements . . . . . . . . . . . . . . . . . . . . . . . . Chriolepis benthonis p. 677 38b. Second dorsal fin with 11 elements; anal fin with 8 elements . . . . . . . . . . . . . . . . . . . . . . . . . . . Chriolepis vespa p. 678 39a. Mouth not completely closing, with protruding outwardly curved teeth. . . . . . . . . . . . . . . . . . . . . . Risor ruber p. 727 39b. Mouth closing normally, without protruding teeth. . . . . 40 40a. Vomer with large teeth . . . . Palatogobius paradoxus p. 724 40b. Vomer without teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 41a. Head without pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 41b. Head with pores. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 42a. Head very depressed, broader than it is deep; body dark ventrally, pale dorsally; pelvic fin spines not connected by membrane. . . . . . . . . . . . . . . . . . . . Gobulus myersi p. 712 42b. Head compressed, deeper than it is wide; body with diffuse spots; pelvic fin spines connected by membrane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nes longus p. 722 43a. Scales present anterior to pelvic fin base; top of head scaled; head compressed. . . . . . . . . . . . . . . . . . . . . . . . . 44 43b. Scales absent anterior to pelvic fin base; top of head without scales, or with scales but head depressed . . . . . . . . . 47 44a. Second dorsal fin with 12 elements. . . . . . . . . . . . . . . . . 45 44b. Second dorsal fin with 13 to 15 elements . . . . . . . . . . . . 46 45a. Anal fin with 12 elements; no black band on upper lip . . . . . . . . . . . . . . . . . . . . . . . . . . Bollmannia litura p. 676 45b. Anal fin with 13 elements; black band on upper lip . . . . . . . . . . . . . . . . . . . . . . Bollmannia eigenmanni p. 675

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666 PERCIFORMES

46a. Second dorsal fin with 13 elements; row of scales along lower margin of cheek . . . . . . . . . . . . . . . . . . . Bollmannia boqueronensis p. 673 46b. Second dorsal fin typically with 14 elements, occasionally 13 or 15; no row of scales along lower margin of cheek . . . . . . . . . . . . . . . . . . . . . . . Bollmannia communis p. 674 47a. Tongue bilobed; no pore above and between eyes. . . . . . 48 47b. Tongue usually rounded, rarely bilobed; medial pore above and between eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 48a. Preopercular sensory canal with three pores; second dorsal fin usually with more than 17 elements; anal fin usually with more than 18 elements; scales in lateral series greater than 65 . . . . . . . . . . . . . . . . Microgobius microlepis p. 720 48b. Preopercular sensory canal with two pores; second dorsal fin usually with 17 or fewer elements; anal fin usually with 18 or fewer elements; scales in lateral series fewer than 65 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 49a. Most scales on body ctenoid; lower jaw with four enlarged caninelike teeth in outer row; interorbital width about 4% of SL; side of body with one broad and two narrow yellow stripes . . . . . . . . . . . . . . . . . . . . . Microgobius carri p. 718 49b. Most scales on body cycloid; lower jaw with about eight enlarged caninelike teeth in outer row; interorbital width less than 3% of SL; no yellow stripes on body . . . . . . . . 50 50a. Lateral cephalic sensory canal with three pores; body with numerous dark blotches; mouth of males greatly enlarged (greater than 15% of SL in males) . . . . . . . . . . . . . . . . . . . . . . . . Microgobius gulosus p. 719 50b. Lateral cephalic sensory canal with two pores; body without dark blotches; mouth of males little enlarged (less than 15% of SL in males) . . . . . Microgobius thalassinus p. 721 51a. Body with prominent stripes or transverse bars, or spotted; head distinctly compressed, deeper than it is wide . . . . . 52 51b. Body with diffuse transverse bars or uniformly gray or brown, never with stripes; head rounded or depressed, broader than it is deep . . . . . . . . . . . . . . . . . . . . . . . . . . 57 52a. No transverse bars on body; prominent pale-colored stripe from eye to posterior margin of head and typically extending along body to caudal fin base; typically black stripe ventral to pale-colored stripe . . . . . . . . . . . . . . . . . . . . . 53 52b. Prominent transverse bars or bands on body; no stripes extending entire length of body and if stripe present, extending length of head only . . . . . . . . . . . . . . . . . . . . . . . . . . 56 53a. Rostral frenum present; mouth distinctly inferior, snout overhanging upper lip . . . . . . . Elacatinus oceanops p. 697 53b. Rostral frenum absent (upper lip separated from snout by deep groove); mouth usually terminal or subterminal. . . 54 54a. Tip of snout uniformly dusky (nares may be set in pale patch on either side in juveniles) . . . . . . . Elacatinus horsti p. 695

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GOBIIDAE 667

54b. Tip of snout with distinct pale marking, including anterior midline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 55a. Dark stripe on body extending ventrally to ventral midline; body with lateral pale stripe about as wide as eye width; predorsal area without pale median streak . . . . . . . . . . . . . . . . . . . . . . . . Elacatinus prochilos p. 698 55b. Dark stripe on body not extending ventrally to ventral midline; lateral pale stripe narrower than eye width; predorsal dark area with pale median streak . . . . . . . . . . . . . . . . . . . . . . Elacatinus xanthiprora p. 700 56a. Body straw colored, with 13 prominent dark mahogany– colored bands posterior to pectoral fin base . . . . . . . . . . . . . . . . . . . . . . . . Elacatinus macrodon p. 696 56b. Body variously spotted or banded, if banded, with fewer than 13 bands and not dark mahogany colored . . . . . . . . . . . . . . . . . . . . . . . . . . Elacatinus saucrus p. 699 57a. Short segment of lateral line present above operculum, consisting of canal with pore at each end; body naked to scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58 57b. Short segment of lateral line absent above operculum, a single sensory pore above preoperculum may be present; body completely naked . . . . . Gobiosoma robustum p. 711 58a. Body naked; anal fin ray elements usually 11 (rarely 10 or 12); second dorsal fin ray elements usually 13 (rarely 12 or 14) . . . . . . . . . . . . . . . . . . . . . . Gobiosoma bosc p. 706 58b. Body with at least two basicaudal scales, posterior section of body occasionally extensively scaled; anal fin ray elements usually 10 (except usually 11 in G. ginsburgi); second dorsal fin ray elements usually 10 to 12 (rarely 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 59a. Two small basicaudal scales, located at upper and lower end of caudal fin base; pectoral fin rays 15 to 19 . . . . . . 60 59b. Sides of caudal peduncle scaly, typically with more than eight transverse rows of scales; pectoral fin rays 18 to 22 (except 16 to 18 and usually 17 in G. grosvenori) . . . . . 61 60a. Anal fin ray elements usually 11 (rarely 10 or 12); pectoral fin rays 18 or 19 (rarely 17) . . . . . . . . . . . . . . . . . . . . . . . Gobiosoma ginsburgi p. 707 60b. Anal fin ray elements usually 10 (rarely 9); pectoral fin rays 16 (rarely 15 or 17) . . . . . . . . Gobiosoma longipala p. 710 61a. Second dorsal fin ray elements 10; anal fin ray elements 9; pectoral fin rays 17 (rarely 16 or 18); scales in horizontal series 31 to 35 . . . . . . . . . . . Gobiosoma grosvenori p. 708 61b. Second dorsal fin ray elements 12 (rarely 11 or 13); anal fin ray elements 10 (rarely 9); pectoral fin rays 20 or 21; scales in horizontal series 34 to 36 . . . . . . . . . . . . . . . . . . . . Gobiosoma hemigymnum p. 709

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668 PERCIFORMES

Awaous banana River goby

(Valenciennes, 1837)

Awaous banana has head pores, several fleshy lobes on anterior margin of shoulder girdle under operculum, and a scaled body. Anterior profile of head is broadly rounded, and snout is long and conical. Eye is high on head. Mouth is large and terminal to slightly subterminal. Lips are fleshy, dorsal lip is continuous across symphysis, and lower lip connects with narrow frenum at symphysis. Jaw teeth are slender, long, and unicuspid, with those in upper jaw arranged in two rows and those in outer row slightly enlarged. Teeth in lower jaw are arranged in a band that narrows posteriorly, with outermost teeth slightly enlarged. Measurements are expressed as percent of SL: head length 23%–34%, snout length 11.1%– 14.5%, eye diameter 3.8%– 4.3%, upper jaw length 8%–21% in males and 7%–14% in females, pectoral fin length 21.8%–23.8%, pelvic fin length 17.8%–21.5%, body depth 13.8%–16.4%. Pectoral fin has 14 to 17 rays, first dorsal fin has six spines, second dorsal fin has one spine and 9 to 12 (usually 10) rays, and

anal fin has one spine and 9 to 11 (usually 10) rays. Pelvic fins are broadly united to form disc, and rear edge of disc is rounded or complete. Caudal fin is broadly rounded. Body is covered with small ctenoid scales. Vertebrae number 26: 10 precaudal and 16 caudal. Color is pale yellowish tan, with many small dark blotches and vermiculations along side of body. Dorsal fin is yellowish green. This species occurs in the western Atlantic from Florida to Brazil, including the Gulf of Mexico and the Antilles. In the Gulf of Mexico it occurs from the Florida Panhandle and from southern Texas to Laguna de Términos, Campeche. Adults burrow into sandy and gravelly river bottoms, and juveniles occur in estuaries. Maximum known size is 300 mm TL. references: Robins and Ray 1986; Boschung 1992 (both as A. tajasica); Watson 1996; Smith 1997; Castro-Aguirre et al. 1999; Murdy and Hoese 2002b.

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GOBIIDAE 669

Barbulifer ceuthoecus Bearded goby

(Jordan and Gilbert, 1884)

Barbulifer ceuthoecus has chin, snout, and suborbital barbels and lacks scales. Anterior and posterior nares are tubular and widely separated, with anterior naris directed anterodorsolaterally and posterior naris directed vertically. Head barbels are short, with one located at tip of snout, another between eye and lip, a third on posterior aspect of lip, and 14 to 16 barbels on ventral surface of head. Upper lip has free fold, lacks rostral frenum. Jaw teeth are conical, sharp pointed, and slightly recurved. Tongue is free and bilobed at tip. Gill opening is restricted ventrally to about pectoral fin base. Measurements are expressed as percent of SL: head length 28.9%–31.7%, snout length 5.1%– 6.5%, eye diameter 6.8%– 8.5%, pectoral fin length 24.6%–29%, pelvic fin length 19.1%–20.7%, body depth 14.9%– 17.6%. Pectoral fin has 16 to 19 rays. First dorsal fin has seven spines, and second has one spine and 8 to 10 rays. Pelvic fins are broadly united to form disc, and inner rays are longest. Anal fin has one flexible spine and 7 to 9 rays.

Body is naked. Lateral line system on head consists of an intraorbital pore, an infraorbital pore, two preopercular pores, and a lateral pore above preopercular canal. Vertebrae number 27: 11 precaudal and 16 caudal. Color pattern is variable and consists of dark bars on light background, with adjacent bars often coalesced along lateral midline or totally coalesced dorsally except for light blotches on dorsal midline. Ventral surface is light. This species occurs in the western Atlantic from southern Florida and the Bahamas to Venezuela, including the Gulf of Mexico and the Florida Keys. In the Gulf of Mexico it is known from Florida Bay and Alacrán Reef. This species is associated with rocks and silty rubble in very shallow water. Maximum known size is 30 mm SL.

references: Robins 1958; Hildebrand et al. 1964; Böhlke and Chaplin 1968; Böhlke and Robins 1968; Smith 1997; Murdy and Hoese 2002b.

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670 PERCIFORMES

Bathygobius curacao Notchtongue goby

(Metzelaar, 1919)

Bathygobius curacao has free upper pectoral fin rays, a moderately long upper jaw, and a fully scaled body. Anterior naris is a short tube, and posterior naris has a raised rim. Mouth is terminal, with upper jaw extending to about center of eye and lower jaw just slightly included in upper jaw. Upper lip has free dorsal margin. Jaw teeth are arranged in a narrow band, with outermost teeth enlarged. Tongue is deeply notched. Gill opening extends ventrally to pectoral fin base. Measurements are expressed as percent of SL: head length 26.9%– 29.5%, snout length 5.8%–7.4%, eye diameter 5.1%– 6.3%, upper jaw length 10.3%– 12.7%, pre-dorsal-fin length 34%–37%, pelvic fin length 20%–22.6%, caudal fin length 24.9%–29%, body depth 18.4%–21.4%. Pectoral fin rays number 15 to 18, with upper 4 or 5 rays distally free of membranes and filamentous. First dorsal fin has six spines, and second has one spine and 8 or 9 rays. Pelvic fins are united by membrane to form disc. Anal fin has two spines and 6 or 7 rays. Caudal fin is rounded. Body is completely scaled except for

side of head. Scales in lateral series number 31 to 34 (usually 33). Single sensory pores are located between eyes, behind eyes, and medial to anterior naris. Vertebrae number 27: 10 precaudal and 17 caudal. Color is tan to light brown, with five dark saddles on upper body and small and large dark blotches on flanks. Second dorsal fin and anal fin generally have a tan margin. This species occurs in the western Atlantic from southern Florida and Bermuda to northern South America, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is known from Florida Bay, the Dry Tortugas, and Veracruz in the Gulf of Mexico. This species is associated with tide pools, mangrove habitats, and sea-grass beds. Maximum known size is 75 mm TL. references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Smith 1997; Castro-Aguirre et al. 1999; SmithVaniz et al. 1999; Murdy and Hoese 2002b.

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GOBIIDAE 671

Bathygobius mystacium Island frillfin

Ginsburg, 1947

Bathygobius mystacium has free upper pectoral fin rays, a relatively short upper jaw, and a fully scaled body. Mouth is terminal, upper jaw extends to about center of eye, and lower jaw is slightly included in upper jaw. Upper lip has free dorsal margin. Jaw teeth are arranged in a narrow band, with outermost teeth enlarged. Tongue is rounded. Gill opening extends ventrally to pectoral fin base. Measurements are expressed as percent of SL: head length 28%–30.6%, snout length 6.8%– 8.4%, eye diameter 6.6%– 8.2%, upper jaw length 7.9%–11.4%, pre-dorsal-fin length 34.8%– 37.6%, pelvic fin length 18.8%–24.2%, caudal fin length 20.3%–29%, body depth 19.2%– 22.1%. Pectoral fin rays number 19 or 20, with upper 4 or 5 rays distally free of membranes and filamentous. First dorsal fin has six spines, and second has one spine and 8 or 9 rays. Pelvic fins are united by membrane to form disc. Anal fin has one spine and 7 or 8 rays. Body

is completely scaled except for side of head. Single sensory pores are located between eyes, behind eyes, and medial to anterior naris. Scales in lateral series number 33 to 36 (usually 35). Vertebrae number 27: 10 precaudal and 17 caudal. Color is tan to light brown, with dark brown blotches forming irregular bars or a checkerboard pattern on flanks. White specks often occur on scales. This species occurs in shallow water in the western Atlantic from southern Florida and the Bahamas to Central America, including Campeche Bay and Veracruz in the southern Gulf of Mexico and the Greater and Lesser Antilles. It is associated with sandy to rocky bottoms exposed to wave action. Maximum known size is 150 mm TL. references: Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997; Murdy and Hoese 2002b.

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672 PERCIFORMES

Bathygobius soporator Frillfin goby

(Valenciennes, 1837)

Bathygobius soporator has upper pectoral fin rays free of membranes, a relatively long upper jaw, and a fully scaled body. Mouth is terminal, upper jaw extends to about center of eye, and lower jaw is slightly included in upper jaw. Lips are thick and swollen, and upper lip has a free dorsal margin. Jaw teeth are arranged in a narrow band, with outermost teeth enlarged. Tongue is slightly notched. Gill opening extends ventrally to pectoral fin base. Gill rakers on first arch number 9 to 11. Measurements are expressed as percent of SL: head length 28.9%–33.8%, snout length 7.4%–9.3%, eye diameter 5%–7.6%, upper jaw length 11.2%– 14.6%, pre-dorsal-fin length 35.3%–36.8%, pelvic fin length 20.3%–23.7%, caudal fin length 24.2%–30.2%, body depth 18.9%– 26.3%. Pectoral fin rays number 18 to 21 (usually 19 or 20), with upper 4 or 5 rays distally free of membrane and filamentous. First dorsal fin has six spines, and second has one spine and 9 or 10 rays. Pelvic fins are united by membrane to form disc. Anal fin has one spine and 7 to 9 rays. Body is completely scaled except for side of head. Single sensory pores are located between eyes, behind eyes, and medial to anterior naris. Scales on dorsal side of

head and chest are cycloid and embedded, and scales on remainder of body are weakly ctenoid. Scales in horizontal series number 37 to 44. Vertebrae number 27: 10 precaudal and 17 caudal. Color is brown, either blotched or uniform, with five black saddles; the one over first dorsal fin is broadest. A blue spot is located on upper region of gill opening. This species occurs in the tropical Atlantic. In the western Atlantic it occurs from North Carolina and Bermuda to southeastern Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with irregular substrates in very shallow water from brackish estuaries to rocky tide pools. Food consists of insect larvae and caridean shrimps. Maximum known size is 75 mm SL.

references: Longley and Hildebrand 1941; Springer and Woodburn 1960; Böhlke and Chaplin 1968; Dawson 1969b; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Randall 1996; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Murdy and Hoese 2002b.

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GOBIIDAE 673

Bollmannia boqueronensis White-eye goby

Evermann and Marsh, 1899

Bollmannia boqueronensis has scales on top of head and on side of body anterior to pectoral fin. Head is moderately elevated and convex and moderately compressed. Mouth is terminal and moderately oblique, with upper jaw extending to posterior margin of pupil. Rostral frenum is absent. Jaw teeth are arranged in narrow bands, with outermost teeth slightly enlarged. Gill opening extends ventrally the length of pectoral fin base. Measurements are expressed as percent of SL: head length 29%– 31.6%, snout length 5.1%–5.9%, eye diameter 9.9%–11.5%, pectoral fin length 25.2%– 27.7%, pelvic fin length 22.2%–29.2%, caudal fin length 32.7%– 61.1%, body depth 18.2%–21%. Pectoral fin rays number 20 or 21. First dorsal fin has seven spines, and second has one spine and 12 rays. Pelvic fins are united by membrane to form disc and possess frenum uniting spines of each fin. Anal fin has

one spine and 12 rays. Caudal fin is lanceolate. Body, including chest, is covered with large deciduous scales, and scales also occur on top of head, nape, upper section of operculum, and upper and lower cheek. Scales in horizontal series number 27 to 30. Vertebrae number 27: 11 precaudal and 16 caudal. Color is pale, with head yellowish, iris white, a dark ocellus on first dorsal fin, and yellow to orangish stripes and spots on second dorsal fin. This species occurs between 27 and 55 m in the western Atlantic from southern Florida to northern South America, including the eastern Gulf of Mexico and the Lesser Antilles. In the Gulf of Mexico it occurs at the Dry Tortugas. Maximum known size is 100 mm SL. references: Evermann and Marsh 1900; Ginsburg 1935; Longley and Hildebrand 1941; Robins and Ray 1986; Smith 1997; Murdy and Hoese 2002b.

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674 PERCIFORMES

Bollmannia communis Ragged goby

Ginsburg, 1942

Bollmannia communis has scales on top of head and on side of body anterior to pectoral fin. Head is moderately elevated and convex and moderately compressed. Mouth is terminal and moderately oblique. Rostral frenum is absent. Jaw teeth are arranged in two rows, with outer row enlarged and caninelike. Gill opening extends ventrally the length of pectoral fin base. Measurements are expressed as percent of SL: head length 29.1%–31.8%, snout length 5.8%– 6.7%, eye diameter 7.6%– 8.2%, interorbital width 1%–1.9%, pre-dorsal-fin length 32.1%–35.8%, pelvic fin length 24.3%– 27.6%, caudal fin length 32.7%– 61.1%, body depth 20.3%–23.4%. Pectoral fin rays number 21 to 23. First dorsal fin has seven spines, and second has one spine and 13 to 15 (usually 14) rays. Pelvic fins are united by membrane to form disc and possess frenum uniting spines of each fin. Anal fin has one spine and 12 to 14 (usually 13) rays. Caudal fin is lanceolate.

Body, including chest, is covered with large deciduous scales, and scales also occur on top of head, nape, upper section of operculum, and upper cheek posterior to eye. Vertebrae number 27: 11 precaudal and 16 caudal. Color is pale brown and darker dorsally than ventrally, with narrow white stripes and a black blotch on spiny dorsal fin. Iris of eye is dark. In males, second dorsal, anal, and caudal fins are greatly elongated, and first dorsal fin has a black band above blotch. This species occurs in the western Atlantic from southern Florida and the Gulf of Mexico between 10 and 70 m. It occurs throughout the Gulf of Mexico within its depth range. Maximum known size is 100 mm TL. references: Ginsburg 1942; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Murdy and Hoese 2002b.

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GOBIIDAE 675

Bollmannia eigenmanni

(Garman, 1896)

Bollmannia eigenmanni has scales on top of head and on side of body anterior to pectoral fin. Head is moderately elevated and convex and moderately compressed. Mouth is terminal and moderately oblique. Rostral frenum is absent. Jaw teeth are arranged in two rows, with outer row enlarged and caninelike. Gill opening extends ventrally the length of pectoral fin base. Measurements are expressed as percent of SL: head length 25.6%–29.8%, snout length 4%– 4.4%, eye diameter 9.4%–9.8%, interorbital width 2%–1.6%, pre-dorsal-fin length 32.4%–33.3%, pelvic fin length 23%– 26.8%, caudal fin length 36.5%– 40.5%, body depth 17.8%–17.9%. Pectoral fin rays number 23 to 25. First dorsal fin has seven spines, and second has one spine and 11 rays. Pelvic fins

are united by membrane to form disc and possess frenum uniting spines of each fin. Anal fin has one spine and 12 rays. Caudal fin is lanceolate. Body, including chest, is covered with large deciduous scales, and scales also occur on top of head, nape, upper section of operculum, and upper cheek posterior to eye. Color is pale brown and darker dorsally than ventrally, with a black band on upper lip. This species occurs in the western Atlantic from southern Florida and the eastern Gulf of Mexico off Alabama and Florida. Maximum known size is about 180 mm SL.

references: Garman 1896; Williams and Shipp 1980; Boschung 1992; Murdy and Hoese 2002b.

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676 PERCIFORMES

Bollmannia litura

Ginsburg, 1935

Bollmannia litura has scales on top of head and on side of body anterior to pectoral fin. Head is evenly convex to origin of dorsal fin and moderately compressed. Mouth is terminal and moderately oblique. Rostral frenum is absent. Jaw teeth are arranged in three irregular rows, with outer row of upper jaw and inner row of lower jaw moderately enlarged. Tongue is fleshy and broadly emarginate. Gill opening extends ventrally the length of pectoral fin base. Measurements are expressed as percent of SL: head length about 31%, snout length about 8%, eye diameter about 12%, pelvic fin length about 31%, caudal fin length about 49%, body depth about 25%. Pectoral fin rays number 20. First dorsal fin has seven spines, and second has one spine and 11 rays. Middle dorsal fin spines are elongated, with fourth the longest. Pelvic fins are united by membrane to form disc. Anal fin has one spine and 11 rays. Caudal fin is lanceolate. Body, including chest, is covered with large deciduous

scales, and scales also occur on top of head, nape, upper section of operculum, and upper cheek posterior to eye. Scales on body and caudal peduncle are weakly ctenoid, and other scales are cycloid. Scales in horizontal series number 28. Vertebrae number 27: 11 precaudal and 16 caudal. Color is pale yellow, with dorsal section of body slightly darker than ventral section and five indistinct blotches on side. Dorsal fins and upper section of caudal fin have several dark elliptical markings with lightcolored centers, and posterior section of first dorsal fin has a black blotch. This species occurs in the western Atlantic from the northwestern Gulf of Mexico, off Port O’Connor, TX, the Dominican Republic, and Puerto Rico. Maximum known size is 60 mm SL.

references: Ginsburg 1935 (as Bollmania litura); Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 677

Chriolepis benthonis

Ginsburg, 1953

Chriolepis benthonis has an enlarged head, separate pelvic fins, and scales only on posterior section of body. Mouth is terminal and moderately oblique, with upper jaw extending to pupil of eye. Teeth in jaws are in several rows, with those in outer row consisting of widely spaced, recurved canines. Tongue is rounded. Gill opening is restricted ventrally to about pectoral fin base. Pectoral fin has 16 rays. First dorsal fin has seven spines, and second dorsal fin has one spine and 9 rays. Fourth ray of pelvic fin is very long, and fifth ray is very short and closely applied to fourth ray. Anal fin has one spine and 8 rays. Scales are present from below seventh dorsal fin ray to caudal

peduncle, in nine or more rows, with anterior rows cycloid and last three rows ctenoid. Sensory pores are lacking on head, but sensory papillae are arranged in rows. Vertebrae number 27: 11 precaudal and 16 caudal. Color in life is pale yellow, with scattered chromatophores on head and body. This species occurs in the western Atlantic from the southern Gulf of Mexico off Progreso, Yucatán, and on Arrowsmith Bank off Quintana Roo at 154 to 350 m.

references: Ginsburg 1953; Hastings and Bortone 1981; Smith 1997; Murdy and Hoese 2002b.

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678 PERCIFORMES

Chriolepis vespa

Hastings and Bortone, 1981

Chriolepis vespa has an enlarged head, separate pelvic fins, and scales only on posterior section of body. Mouth is terminal and moderately oblique, with upper jaw extending to pupil of eye. Teeth in upper jaw are in five rows anteriorly and three rows posteriorly; those in outer row are widely spaced, recurved canines, and remainder are small and close set. Teeth in lower jaw are in four rows anteriorly and one row posteriorly; those in outer row are widely spaced and recurved canines, those in inner row are relatively small canines, and those in middle rows are small and close set. Tongue is truncate to very shallowly indented. Gill opening is restricted ventrally to about pectoral fin base. Measurements are expressed as percent of SL: head length 29.3%–35.4%, snout length 5.2%– 8.5%, eye diameter 9.8%– 12.5%, upper jaw length 11.1%–12.3%, pectoral fin length 28.4%–36.4%, pelvic fin length 27.1%–36.5%, body depth 13%–18.7%. Pectoral fin has 15 to 17 rays. First dorsal fin has

seven spines, and second dorsal fin has one spine and 10 rays. Fourth ray of pelvic fin is very long, and fifth ray is very short and closely applied to fourth ray. Anal fin has one spine and 7 rays. Anteriormost scales are cycloid, and the remainder are ctenoid. Sensory pores are lacking on head. Vertebrae number 27: 11 precaudal and 16 caudal. Color in life is pale yellowish, somewhat pinkish anteriorly and translucent posteriorly, with four prominent olive bars and several less prominent olive saddles on body. In preservative, body is pallid, with bars and saddles visible as concentrations of melanophores. This species occurs between 35 and 183 m in the western Atlantic from the northeastern Gulf of Mexico off Mobile Bay, Pensacola, St. Petersburg, and possibly off Guyana. Maximum known size is 40 mm SL. references: Hastings and Bortone 1981; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 679

Coryphopterus dicrus Böhlke and Robins, 1960 Colon goby

Coryphopterus dicrus has a posteriorly notched pelvic fin disc and two dark, colonshaped spots on pectoral fin base. Upper lip has free dorsal fold, lacks frenum. Poorly developed, fleshy mid-dorsal ridge extends from level of orbits to dorsal fin origin. Jaw teeth are conical, pointed, and arranged in several rows, with those of outer row enlarged. Tongue is free anteriorly and truncate. Gill opening extends ventrally to pectoral fin base, and gill membrane is broadly joined to isthmus to form V-shaped fold. Gill rakers on lower limb number seven. Measurements are expressed as percent of SL: head length 27.3%–32.2%, eye diameter 8.1%–9.8%, pectoral fin length 26.2%–31.4%, pelvic fin length 24.4%– 27.9%, body depth at dorsal fin origin 19.9%– 26.4%, caudal peduncle depth 12.6%–14.6%. Pectoral fin has 18 to 20 (usually 19 or 20) rays. First dorsal fin has six spines, and second dorsal fin has one spine and 9 rays. Pelvic fins are joined by membrane, but disc is incomplete or concave posteriorly because fifth pelvic fin ray is considerably shorter than fourth. Pelvic fin spines are not joined by frenum. Anal fin

has one spine and 9 rays. Caudal fin is truncate and has 17 segmented rays and 12 or 13 branched rays. Genital papilla of males is about half the length of anal spine. Body is covered with cycloid scales on chest and pectoral fin base and with ctenoid scales on remainder of body. Head and nape are naked. Scales in horizontal series number 26. Vertebrae number 26: 10 precaudal and 16 caudal. Color is pallid, with 10 or 11 dark reddish brown markings and small pale blue green flecks on body and two small, dark colon-shaped spots on pectoral fin base. This species occurs in the western Atlantic from southern Florida and the Bahamas to Central America, including the Florida Keys, the southern Gulf of Mexico, Puerto Rico, and the Virgin Islands. In the Gulf of Mexico it is known from Isla Pérez, Alacrán Reef. Maximum known size is 50 mm SL. references: Böhlke and Robins 1960a; Böhlke and Chaplin 1968; Robins and Ray 1986; Randall 1996; Smith 1997; Murdy and Hoese 2002b.

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680 PERCIFORMES

Coryphopterus eidolon Pallid goby

Böhlke and Robins, 1960

Coryphopterus eidolon has two pale stripes on side of head and a blackish bar at base of caudal fin. Upper lip lacks frenum, and upper jaw is moderately protrusible. Low, fleshy middorsal ridge extends from just behind orbits to dorsal fin origin. Jaw teeth are conical, pointed, and arranged in five or six rows, with those of outer and inner rows enlarged. Tongue is free anteriorly and truncate to slightly trilobate. Gill opening extends ventrally to pectoral fin base, and gill membrane is broadly joined to isthmus to form V-shaped fold. Gill rakers on first arch number six or seven on lower limb. Measurements are expressed as percent of SL: head length 31%–33.5%, eye diameter 8.4%– 9.8%, pectoral fin length 23.1%–30.9%, pelvic fin length 24%–28.3%, body depth at dorsal fin origin 23.3%–26%, caudal peduncle depth 11.6%–13.2%. Pectoral fin has 18 to 20 (usually 19 or 20) rays. First dorsal fin has six spines, and second dorsal fin has one spine and 8 to 11 (usually 10) rays. Pelvic fins are joined by membrane to form complete disc, although fifth pelvic fin ray is shorter than fourth. Opposing pelvic fin spines are con-

nected by frenum. Anal fin has one spine and 8 or 9 rays. Caudal fin is truncate and has 17 segmented and 12 or 13 branched rays. Genital papilla of males is elongate and pointed. Body is covered with cycloid scales on chest and pectoral fin base and with ctenoid scales on remainder of body. Head and nape are naked. Scales in horizontal series number 25 or 26. Vertebrae number 26: 10 precaudal and 16 caudal. Color is pallid, with small yellow spots or short bands on snout, cheek, and anterior aspect of body; a narrow yellow stripe outlined by narrow dark lines extending from eye to first dorsal fin; and a narrow black bar at base of caudal fin. Pectoral fin base lacks dark spot. This species occurs in the western Atlantic from southern Florida and the Bahamas to Haiti and the Virgin Islands, including the southern Gulf of Mexico. It is associated with coral reefs and fine coralline sands. Maximum known size is 60 mm SL. references: Böhlke and Robins 1960a; Böhlke and Chaplin 1968; Robins and Ray 1986; Randall 1996; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 681

Coryphopterus glaucofraenum Bridled goby

Gill, 1863

Coryphopterus glaucofraenum has a small spot or an elongate dark bar above operculum and three rows of dark spots on body. Upper lip lacks frenum, has free dorsal fold. Welldeveloped, fleshy mid-dorsal ridge extends from just behind orbits to dorsal fin origin. Jaw teeth are conical, pointed, and arranged in five or six rows, with those of outer and inner rows enlarged. Tongue is free anteriorly and truncate to broadly rounded. Gill opening extends ventrally to pectoral fin base, and gill membranes are broadly joined to isthmus to form V-shaped fold. Gill rakers on first arch number 11 on lower limb. Measurements are expressed as percent of SL: head length 27.4%–30.4%, eye diameter 6.4%– 8.1%, pectoral fin length 22.8%–30.1%, pelvic fin length 23%–28.6%, body depth at dorsal fin origin 21.3%–28.6%, caudal peduncle depth 11.4%–14.1%. Pectoral fin has 17 to 20 (usually 18 or 19) rays. First dorsal fin has six spines, and second has one spine and 9 rays. Pelvic fins are joined by membrane to form complete disc, fifth pelvic fin ray is nearly as long as fourth ray, and pelvic spines are joined by frenum. Anal fin has one spine and 9 rays. Caudal fin is truncate and has 17 segmented rays and 12 or 13 branched rays. Genital papilla of males is elongate and pointed. Body is covered with cycloid scales

on chest and pectoral fin base and with ctenoid scales on remainder of body. Head and nape are naked. Scales in horizontal series number 26 to 28. Vertebrae number 26: 10 precaudal and 16 caudal. Color is pallid, with a series of dark spots and X-shaped marks on flanks, a small triangular dark spot or short dark bar above operculum, and a dark bar or two vertically oriented dark spots on base of caudal fin. Specimens occupying light sandy bottoms in clear water are more lightly pigmented than specimens occupying dark substrates or turbid water. This species occurs between 5 and 32 m in the western Atlantic from North Carolina and Bermuda to northern South America, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from the Dry Tortugas, the Florida Middle Grounds, Alabama, and Campeche Bay. Maximum known size is 75 mm SL.

references: Longley and Hildebrand 1941; Robins 1958; Böhlke and Robins 1960a; Böhlke and Chaplin 1968; Robins and Ray 1986; Garzón and Acero 1990 (in part as Coryphopterus tortugae); Boschung 1992; Smith 1997; Smith-Vaniz et al. 1999; Murdy and Hoese 2002b.

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682 PERCIFORMES

Coryphopterus hyalinus Glass goby

Böhlke and Robins, 1962

Coryphopterus hyalinus has a triangular patch of pigment behind eye and a prominent black ring around anus. Upper lip has free dorsal fold, lacks frenum. Low, fleshy mid-dorsal ridge extends from level of orbits to dorsal fin origin. Jaw teeth are conical, pointed, and arranged in several rows, with those of outer row enlarged. Tongue is free anteriorly and truncate, bilobate, or trilobate. Gill opening extends ventrally to pectoral fin base, and gill membranes are broadly joined to isthmus to form V-shaped fold. Gill rakers on first arch number one on upper limb and eight or nine on lower limb. Measurements are expressed as percent of SL: head length 28.7%–31.5%, eye diameter 9.7%–11.6%, body depth at dorsal fin origin 22.8%–26%, caudal peduncle depth 11%–12.8%. Pectoral fin has 14 to 16 (usually 15) rays. First dorsal fin has five or six spines, and second has one spine and 9 rays. Pelvic fins are joined by a very narrow membrane at base. Anal fin has one spine and 9 (rarely 8 or 10) rays. Caudal fin is truncate to slightly emarginate. Body is covered with cycloid scales on chest and pectoral fin base and with ctenoid

scales on remainder of body. Head and nape are naked. A pair of sensory pores is located between anterior margins of eyes, and a single pore is located on midline posterior to eyes. Scales in horizontal series number 24 to 26. Vertebrae number 26: 10 precaudal and 16 caudal. Color is translucent, with gold or orange tint. Dark masklike pigment is present on snout and continues behind eye as a triangular patch. A distinct dark ring surrounds anus and extends onto belly anteriorly and to genital papilla posteriorly. Scales are outlined with dark pigment. This species occurs between 2.4 and 52 m in the western Atlantic from the Florida Keys and Bermuda to Colombia, including the Bahamas, Haiti, and the Lesser Antilles. It has not been recorded from the Gulf of Mexico, but it may occur there. This species is probably associated with coral reefs. Maximum known size is 29.6 mm SL. references: Böhlke and Robins 1962; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Murdy and Hoese 2002b.

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GOBIIDAE 683

Coryphopterus lipernes Bluenose goby

Böhlke and Robins, 1962

Coryphopterus lipernes has two dusky stripes behind eye and a prominent black ring around anus. Upper lip has free dorsal fold, lacks frenum. Low, fleshy mid-dorsal ridge extends from level of orbits to dorsal fin origin. Jaw teeth are conical, pointed, and arranged in several rows, with those of outer row enlarged. Tongue is free anteriorly and truncate, bilobate, or trilobate. Gill opening extends ventrally to pectoral fin base, and gill membranes are broadly joined to isthmus to form V-shaped fold. Gill rakers on first arch number one to four on upper limb and eight on lower limb. Measurements are expressed as percent of SL: head length 30.2%–31.5%, eye diameter 10.3%–11.9%, body depth at dorsal fin origin 20.2%–23.9%, caudal peduncle depth 11.6%–11.9%. Pectoral fin has 16 to 18 rays. First dorsal fin has six spines, with second spine greatly elongated. Second dorsal fin has one spine and 9 rays. Pelvic fins are joined by a very narrow membrane at base. Anal fin has one spine and 9 rays. Caudal fin is truncate to

slightly emarginate. Body is covered with cycloid scales on chest and pectoral fin base and with ctenoid scales on remainder of body. Head and nape are naked. Two sensory pores are located along midline between eyes. Scales in horizontal series number 26. Vertebrae number 26: 10 precaudal and 16 caudal. Color is translucent yellow, with reddish blotches. Dark blue pigment is present on snout, between orbits, and on lips, and a dark ring surrounds anus. Scales above midline are outlined with dark pigment. This species occurs between 15 and 24 m in the western Atlantic from the Florida Keys and the Bahamas to Central America and the Antilles. It has not been recorded from the Gulf of Mexico, but it may occur there. This species is associated with coral reefs. Maximum known size is 32 mm SL. references: Böhlke and Robins 1962; Robins and Ray 1986; Smith 1997; Murdy and Hoese 2002b.

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684 PERCIFORMES

Coryphopterus personatus Masked goby

(Jordan and Thompson, 1905)

Coryphopterus personatus has a black mask on snout and a prominent black ring around anus. Upper lip has free dorsal fold, lacks frenum. Low, fleshy mid-dorsal ridge extends from level of orbits to dorsal fin origin. Jaw teeth are conical, pointed, and arranged in several rows, with those of outer row enlarged. Tongue is free anteriorly and truncate. Gill opening extends ventrally to pectoral fin base, and gill membranes are broadly joined to isthmus to form V-shaped fold. Gill rakers on first arch number 1 or 2 on upper limb and 8 to 10 on lower limb. Measurements are expressed as percent of SL: head length 26.6%– 31.2%, eye diameter 8%–10.6%, pectoral fin length 28.2%–28.6%, pelvic fin length 21.6%– 25.1%, body depth at dorsal fin origin 21.3%– 27%, caudal peduncle depth 11%–13%. Pectoral fin has 14 to 16 (usually 15 or 16) rays. First dorsal fin has six spines, with second spine elongated in adults. Second dorsal fin has one spine and 9 or 10 rays. Pelvic fins are joined by a very narrow membrane at base. Anal fin has one spine and 9 or 10 rays. Caudal fin is emar-

ginate. Body is covered with cycloid scales on chest and pectoral fin base and with ctenoid scales on remainder of body. Head and nape are naked. Scales in horizontal series number 24 to 28. Two sensory pores are located along midline between eyes. Vertebrae number 26: 10 precaudal and 16 caudal. Color is translucent, with a gold or orange tint. Dark mask on snout extends through eye, and distinct dark ring surrounds anus and extends onto belly anteriorly and to genital papilla posteriorly. This species occurs between 2 and 30.5 m in the western Atlantic from southern Florida and Bermuda to the east coast of Yucatán, including the Gulf of Mexico, the Bahamas, Haiti, and the Virgin Islands. In the Gulf of Mexico it is known from the Dry Tortugas and Alacrán Reef. Maximum known size is 35 mm SL. references: Longley and Hildebrand 1941 (in part); Böhlke and Robins 1962; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Murdy and Hoese 2002b.

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GOBIIDAE 685

Coryphopterus punctipectophorus Spotted goby

Springer, 1960

Coryphopterus punctipectophorus has a small spot on ventral half of pectoral fin base. Upper lip lacks frenum, has free dorsal fold. Low mid-dorsal ridge extends from just behind orbits to dorsal fin origin. Jaw teeth are conical, pointed, and arranged in five or six rows, with those of outer and inner rows enlarged. Tongue is free anteriorly and truncate to broadly rounded. Gill opening extends ventrally to pectoral fin base, and gill membranes are broadly joined to isthmus to form V-shaped fold. Measurements are expressed as percent of SL: head length 26.1%–29%, eye diameter 8.3%–9%, pectoral fin length 26.2%–30.2%, body depth at dorsal fin origin 18.8%–23.7%, caudal peduncle depth 12.3%–14%. Pectoral fin has 18 to 20 (usually 18 or 19) rays. First dorsal fin has six spines, and second has one spine and 10 rays. Pelvic fins are joined by a membrane to form complete disc, with fifth pelvic fin ray nearly as long as fourth ray, and pelvic fin spines are joined by frenum. Anal fin has one spine and 9 rays. Caudal fin is truncate

and has 12 or 13 branched rays. Body is covered with cycloid scales on chest and pectoral fin base and with weak ctenoid scales on remainder of body. Head and nape are naked. Scales in lateral series number 25 to 28 (usually 26 or 27). Vertebrae number 26: 10 precaudal and 16 caudal. Color is white, with a row of dusky light buff spots on dorsal contour and two rows of coral pink spots on flank. Head and nape have several orange lines. This species occurs in the western Atlantic from the Gulf of Mexico. In the Gulf it occurs between St. Andrew Bay and St. Petersburg, FL, and off Veracruz from 18 to 36.6 m. It lives in burrows near rocky areas. Maximum known size is 75 mm SL. references: Böhlke and Robins 1960a; Springer 1960; Springer and Woodburn 1960 (as Coryphopterus sp.); Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Murdy and Hoese 2002b.

10A-T3324 9/13/05 11:18 AM Page 686

686 PERCIFORMES

Coryphopterus thrix Böhlke and Robins, 1960 Bartail goby

Coryphopterus thrix has second dorsal fin spine extended as a filament and a large black spot on pectoral fin base. Upper lip lacks frenum and is moderately protrusible. Low, fleshy mid-dorsal ridge extends from just behind orbits to dorsal fin origin. Jaw teeth are conical, pointed, and arranged in six irregular rows, with those of outer and inner rows enlarged. Tongue is free anteriorly and truncate. Gill opening extends ventrally to pectoral fin base, and gill membranes are broadly joined to isthmus to form V-shaped fold. Measurements are expressed as percent of SL: head length 31.7%– 32.9%, eye diameter 8.8%–9.7%, pectoral fin length 27%–28.5%, pelvic fin length 22.2%– 27.8%, body depth at dorsal fin origin 19.4%– 20.4%, caudal peduncle depth 9.4%–12.4%. Pectoral fin has 17 to 19 rays. First dorsal fin has six spines, and second has one spine and 8 or 9 rays. Pelvic fins are joined by a membrane to form complete disc, with fifth fin ray nearly as long as fourth ray, and pelvic fin spines are joined by frenum. Anal fin has one spine and

9 rays. Caudal fin is truncate and has 17 segmented rays and 12 or 13 branched rays. Genital papilla of males is long, pointed, and tubular. Body is covered with cycloid scales on chest and pectoral fin base and with ctenoid scales on remainder of body. Head and nape are naked. Scales in horizontal series number 25 or 26. Vertebrae number 26: 10 precaudal and 16 caudal. Color is pallid, with a large black spot on dorsal part of pectoral fin base, gill membranes and pelvic fins dusky, and a black bar on base of caudal fin. This species occurs in the western Atlantic from southern Florida, the Bahamas, and the Gulf of Mexico. In the Gulf of Mexico it has been captured off Texas. It is associated with clear water around coral heads. Maximum known size is 50 mm SL.

references: Böhlke and Robins 1960a; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 687

Ctenogobius boleosoma Darter goby

(Jordan and Gilbert, 1882)

Ctenogobius boleosoma has four or five dark blotches along midflank, and most of body is covered with scales. Mouth is terminal and slightly oblique, and upper jaw extends to middle of eye. Lips are thin, and upper lip is free at symphysis. Jaw teeth are arranged in several rows, with outer teeth slightly larger in females and immature males. Mature males have seven or eight large, recurved, widely spaced caninelike teeth in outer row of upper jaw. Gill opening extends ventrally below pectoral fin base. Measurements are expressed as percent of SL: head length 23.9%–27.2%, snout length 6.8%– 8.3%, eye diameter 4.2%– 6.3%, upper jaw length 7.7%–9.7%, pectoral fin length 20.1%–26.1%, pelvic fin length 21.3%–25.2%, caudal fin length 30.8%– 40.2%, body depth 16.6%–19.7%. Pectoral fin has 16 rays. First dorsal fin has six spines, and second has one spine and 9 to 11 (typically 10) rays. Pelvic fins are united by membrane to form broad disc, and pelvic fin spines are joined by frenum. Anal fin has one spine and 10 to 12 (typically 11) rays. Pectoral and pelvic fin rays are longer in adult males than in ju-

veniles and females. Body is covered with cycloid scales anteriorly and ctenoid scales posteriorly. Predorsal region is naked or has 1 or 2 scales. Scales in horizontal series number 29 to 33. A pair of sensory pores is located between eyes. Vertebrae number 26: 10 precaudal and 16 caudal. Color is tan to dusky, with four or five narrow longitudinal brown spots or bars along midline. Spots under second dorsal fin form V-shaped patterns. Dorsal and caudal fins are streaked with small brown spots. This species occurs in the western Atlantic from Delaware Bay and Bermuda to Brazil, including the northern and southern Gulf of Mexico and the Bahamas. It is associated with bays and with brackish and fresh waters on muddy bottoms among submerged vegetation. Maximum known size is 75 mm SL.

references: Ginsburg 1932; Böhlke and Chaplin 1968; Dawson 1969b; Gilbert and Randall 1979; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997 (all previous as Gobionellus boleosoma); Castro-Aguirre et al. 1999; SmithVaniz et al. 1999; Murdy and Hoese 2002b.

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688 PERCIFORMES

Ctenogobius claytonii

(Meek, 1902)

Ctenogobius claytonii has distinct blotches on side, and most of body is covered with scales. Mouth is terminal and slightly oblique, and upper jaw extends to middle of eye. Lips are thin, and upper lip is free at symphysis. Jaw teeth are arranged in two rows, with five or six in outer row of upper jaw enlarged and slightly recurved. Outer teeth in lower jaw are slightly enlarged. Gill opening extends ventrally below pectoral fin base. Measurements are expressed as percent of SL: head length 25.1%– 26.9%, snout length 6.2%–7.3%, eye diameter 5.9%– 6.1%, upper jaw length 8.7%–9.9%, pre-dorsal-fin length 32.3%–36.4%, pelvic fin length 19.7%–22.9%, caudal fin length 25.7%–30.9%, body depth 14.6%–17.3%. Pectoral fin has 15 to 17 rays. First dorsal fin has six spines, and second has one spine and 11 rays. Third spine of first dorsal fin of males is considerably elongate. Pelvic fins are united by membrane to form broad disc, and pelvic fin spines are joined by frenum. Anal fin has one spine and 12 rays. Body is covered with

ctenoid scales except for area between nape and origin of dorsal fin, base of pectoral fin, breast, and midline of belly. Scales in horizontal series number 32 to 37. A pair of sensory pores is located between eyes. Vertebrae number 26: 10 precaudal and 16 caudal. Color is tan, with small flecks of brownish markings on head, two to four dark narrow bars on nape, and five narrow oblong blotches on upper body under dorsal fins. Pelvic fin of adult males is dusky, and pelvic fin of females bears bilateral streaks running parallel to innermost ray. This species occurs in the western Gulf of Mexico from Texas to Veracruz. It is associated with brackish and fresh waters. Maximum known size is 60 mm SL. references: Ginsburg 1932; Gilbert and Randall 1979 (as Gobionellus atripinnis); Smith 1997; Hoese and Moore 1998 (as G. claytoni); CastroAguirre et al. 1999 (as G. claytoni); Murdy and Hoese 2002b.

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GOBIIDAE 689

Ctenogobius fasciatus

Gill, 1858

Ctenogobius fasciatus has a broad, low keel in front of dorsal fin, and most of body is covered with scales. Mouth is terminal, but lower jaw is somewhat included in upper jaw, and upper jaw extends to anterior margin of orbit. Lips are thin, and upper lip is broadly joined to snout by frenum. Jaw teeth are arranged in narrow bands, with eight in outer row of upper jaw enlarged and slightly recurved. Outer teeth in lower jaw are enlarged in males but only slightly so in females. Tongue is slightly emarginate but not bilobed. Gill opening extends ventrally below pectoral fin base. Measurements are expressed as percent of SL: head length 24.5%–26.6%, snout length 6.4%–7.9%, eye diameter 4.4%–5.3%, upper jaw length 6.7%– 8.6%, pre-dorsalfin length 32.3%–35.6%, pelvic fin length 24.6%–32.5%, caudal fin length 29.1%– 40.4%, body depth 14.2%–16.1%. Pectoral fin has 17 or 18 rays. First dorsal fin has six spines, and second has one spine and 11 (or rarely 12) rays. Pelvic fins are united by membrane to form broad disc, and pelvic fin spines are joined by frenum. Anal fin has one spine and 12 rays. Third dorsal fin spine and caudal

fin of males are considerably longer than those of females. Body is covered with ctenoid scales except for area between nape and origin of dorsal fin, base of pectoral fin, chest, and belly. Scales in horizontal series number 35 to 43. A pair of sensory pores is located between eyes, and a single pore is medial to posterior naris. Vertebrae number 26: 10 precaudal and 16 caudal. Color is tan, with five diffuse dark blotches and reticulations and marbling on body. Margin of preoperculum is darkly pigmented, and a dark line runs from posterior aspect of maxilla to behind eye. Dorsal fins and caudal fin are streaked with rows of small spots. This species occurs in the western Atlantic from the southern Gulf of Mexico off Veracruz to Venezuela, including Dominica, Trinidad, Barbados, and Cuba. It is associated with brackish and fresh waters. Maximum known size is 72 mm SL. references: Ginsburg 1932 (as Gobionellus claytonii); Robins and Lachner 1966; Smith 1997; Murdy and Hoese 2002b.

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690 PERCIFORMES

Ctenogobius saepepallens Dash goby

(Gilbert and Randall 1968)

Ctenogobius saepepallens has a broad, low keel in front of dorsal fin and a triangular dark blotch on operculum. Mouth is terminal, and upper jaw extends to middle of eye. Lips are thin, and upper lip lacks frenum. Jaw teeth are arranged in two rows, with teeth in outer row enlarged; three midlateral teeth in upper jaw and one midlateral tooth in lower jaw are recurved and caninelike. Tongue is slightly emarginate but not bilobed. Gill opening extends ventrally to just below pectoral fin base. Measurements are expressed as percent of SL: head length 25.5%–30.7%, snout length 4.5%– 8%, orbit diameter 5.3%–9%, upper jaw length 9.8%–11.8%, pectoral fin length 22%– 29.4%, pelvic fin length 23.8%–31.3%, caudal fin length 35%–53%, body depth 15.6%– 21%. Pectoral fin has 15 to 17 rays. First dorsal fin has six spines, with third spine elongated as a filament. Second dorsal fin has one spine and 11 rays. Pelvic fins are united by membrane to form broad disc, and pelvic fin spines are joined by frenum. Anal fin has one spine and 12 rays. Body is covered with ctenoid scales except for nape to origin of dorsal fin, base of

pectoral fin, chest, and part of belly. Scales in horizontal series number 29 to 34. A pair of sensory pores is located between eyes, and a single pore is medial to posterior naris. Vertebrae number 26: 10 precaudal and 16 caudal. Color is pale bluish gray, with five narrow, elongate dark blotches on midflank and a triangular blotch on middle of operculum. In life, a dark vertical line passes through eye, and blotches on side are edged with pale blue and horizontally divided by a black line. This species occurs from the shoreline to 44 m in the western Atlantic from southern Florida and the Bahamas to Venezuela, including the Florida Keys and the Greater and Lesser Antilles. It has not been reported from the Gulf of Mexico, but it may occur there. This species apparently lives in burrows with snapping shrimps. Maximum known size is 50 mm TL. references: Gilbert and Randall 1968, 1979; Robins and Ray 1986; Randall 1996; Smith 1997 (all as Gobionellus saepepallens); Murdy and Hoese 2002b.

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GOBIIDAE 691

Ctenogobius shufeldti Freshwater goby

(Jordan and Eigenmann, 1887)

Ctenogobius shufeldti has a broad, low keel in front of dorsal fin and five blotches on midside. Mouth is slightly oblique and terminal, but lower jaw is slightly included in upper jaw. Upper jaw extends to posterior margin of eye in males and to about middle of eye in females. Lips are thin, and upper lip is complete across symphysis. Jaw teeth are arranged in narrow bands, with those of outer row of upper jaw slightly enlarged and recurved in males. Tongue is truncate. Gill opening extends ventrally below pectoral fin base. Measurements are expressed as percent of SL: head length 25.4%– 29.6%, snout length 4.9%– 6.9%, eye diameter 3.8%–5.3%, upper jaw length 8.1%– 10.7%, pre-dorsal-fin length 30.7%–33.7%, pelvic fin length 21.1%–24.8%, caudal fin length 29%–33.8%, body depth 15.3%– 19.4%. Pectoral fin has 17 or 18 rays. First dorsal fin has six spines, and second has one spine and 11 (rarely 10 or 12) rays. Pelvic fins are united by membrane to form broad disc, and pelvic fin spines are joined by frenum. Anal fin has one spine and 12 (rarely 11 or 13) rays. Spinous dorsal fin of males is slightly longer than in females, and posterior rays of second

dorsal fin and anal fin are slightly longer in males than in females. Body is covered with ctenoid scales except for area from nape to origin of dorsal fin, base of pectoral fin, chest, and belly. In addition, seven partially embedded cycloid scales occur along midline anterior to dorsal fin origin and on lower half of pectoral fin base. Scales in horizontal series number 35 to 40. A pair of sensory pores is located between eyes, and a single pore is medial to posterior naris. Vertebrae number 26: 10 precaudal and 16 caudal. Color is tan to light brown, with five dark rectangular blotches on side and a dark stripe on cheek. Pelvic fin of mature males has bilateral streaks running parallel to innermost ray. This species occurs in the western Atlantic from North Carolina to the southern Gulf of Mexico. It is found throughout the Gulf of Mexico in low-salinity bays and estuaries. Maximum known size is 84 mm SL. references: Ginsburg 1932; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997 (all as Gobionellus shufeldti); Murdy and Hoese 2002b.

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692 PERCIFORMES

Ctenogobius smaragdus Emerald goby

(Valenciennes, 1837)

Ctenogobius smaragdus has black circles on side of head and a dark spot above pectoral fin base. Mouth is subterminal and slightly oblique, and upper jaw extends to about middle of eye in females and to about posterior margin of eye in males. Lips are thin, and upper lip is free at symphysis. Jaw teeth are arranged in several rows. Outer teeth of upper jaw are enlarged and increase in size toward corner of mouth, and inner teeth on either side of symphysis are enlarged in lower jaw. Jaw teeth in males are relatively larger than those of females. Tongue is emarginate but not bilobed. Gill opening extends ventrally below pectoral fin base. Measurements are expressed as percent of SL: head length 22.3%–28.6%, snout length 7.1%–11.8%, eye diameter 5.4%– 8.8%, upper jaw length 8.1%–13.6%, pectoral fin length 19.4%–25.3%, pelvic fin length 17.8%–24.7%. Pectoral fin has 15 to 18 rays. First dorsal fin has six spines, and second has one spine and 9 to 11 rays. Pelvic fins are united by membrane to form broad disc,

and pelvic fin spines are joined by frenum. Anal fin has one spine and 10 or 11 rays. Males have longer caudal fins than females. Predorsal region has about 11 scale rows. Scales in horizontal series number about 40. A pair of sensory pores is located between eyes. Vertebrae number 26: 10 precaudal and 16 caudal. Color in life is tan to dusky, with green spots outlined with bronze rings on sides of head and body. A green spot is located in mouth, and a dark spot is present on shoulder. In preservative, green spots on head and body appear as light spots surrounded by darker pigment. This species occurs in the western Atlantic from North Carolina to Cuba and Brazil, including the eastern Gulf of Mexico from Charlotte Harbor to Florida Bay. It is associated with mangroves and stagnant backwater habitats. Maximum known size is 100 mm TL. references: Pezold 1984; Robins and Ray 1986; Smith 1997 (all as Gobionellus smaragdus); Murdy and Hoese 2002b.

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GOBIIDAE 693

Ctenogobius stigmaticus Marked goby

(Poey, 1860)

Ctenogobius stigmaticus has a broad, low keel in front of dorsal fin and three dark bars on cheek. Mouth is terminal, but lower jaw is somewhat included in upper jaw, and upper jaw extends to posterior margin of eye. Lips are thin, and upper lip is broadly joined to snout by frenum. Jaw teeth are arranged in narrow bands, with 8 to 10 in outer row of upper jaw greatly enlarged and slightly recurved, especially in males. Outer teeth in lower jaw are enlarged, with one recurved canine at about midlength of jaw on both sexes but larger in males. Tongue is slightly emarginate but not bilobed. Gill opening extends ventrally below pectoral fin base. Measurements are expressed as percent of SL: head length 26.2%–28.6%, snout length 5%– 6.3%, eye diameter 6.4%– 7%, upper jaw length 8.8%–10.7%, pectoral fin length 28.2%–35%, pelvic fin length 29%– 29.6%, caudal fin length 41.3%– 44.6%, body depth 18.5%–18.9%. Pectoral fin has 18 rays. First dorsal fin has six spines, with several spines elongated as filaments. Second dorsal fin has one spine and 11 rays. Pelvic fins are

united by membrane to form broad disc, and pelvic fin spines are joined by frenum. Anal fin has one spine and 12 rays. Spinous dorsal fin and caudal fin of males are considerably longer than those of females. Body is covered with ctenoid scales except for nape to origin of dorsal fin, base of pectoral fin, chest, and belly. Scales in horizontal series number 31 to 36. A pair of sensory pores is located between eyes, and a single pore is medial to posterior naris. Vertebrae number 26: 10 precaudal and 16 caudal. Color is light tan, with three dark bars on cheek and a small spot on shoulder. This species occurs in the western Atlantic from South Carolina to Brazil, including the Gulf of Mexico. In the Gulf of Mexico it is known from the west coast of Florida, from the Keys to Pensacola, and from Port Isabel, TX. Maximum known size is 80 mm SL. references: Ginsburg 1932; Dawson 1969b; Pezold and Edwards 1983; Robins and Ray 1986; Boschung 1992; Smith 1997 (all as Gobionellus stigmaticus); Murdy and Hoese 2002b.

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694 PERCIFORMES

Ctenogobius stigmaturus Spottail goby

(Goode and Bean, 1882)

Ctenogobius stigmaturus has commalike blotches on midside. Mouth is subterminal, and upper jaw extends to at least middle of eye. Lips are thin, and upper lip is complete across symphysis. Jaw teeth are arranged in two or three rows, with two or three teeth on either side of symphysis of upper jaw enlarged and slightly recurved, and two teeth on either side of lower jaw enlarged, especially in males. Tongue is emarginate but not bilobed. Gill opening extends ventrally below pectoral fin base. Measurements are expressed as percent of SL: head length 24.5%–30.4%, snout length 6.8%–10.4%, eye diameter 8.4%– 10.4%, upper jaw length 8.6%–12.8%, pectoral fin length 20.3%–26.3%, pelvic fin length 20.3%–26.9%, caudal fin length 27.9%– 39.9%. Males have relatively longer jaws, pectoral fins, and pelvic fins than females. Pectoral fin has 16 rays. First dorsal fin has six spines, and second has one spine and 10 to 13 (usually 11) rays. Pelvic fins are united by membrane to form broad disc, and pelvic fin spines are joined by frenum. Anal fin has one spine and

10 to 13 (usually 12) rays. Predorsal region generally has 10 to 12 scale rows. Scales in horizontal series number 29 to 33. A pair of sensory pores is located between eyes, and a single pore is medial to posterior naris. Vertebrae number 26: 10 precaudal and 16 caudal. Color is tan, with dark blotches resembling commas on sides. Margin of preoperculum is darkly pigmented, and a dark blotch is located on lower portion of operculum. This species occurs in the western Atlantic from southeastern Florida and Bermuda to Panama, including the eastern Gulf of Mexico and Cuba. It has been reported from Key West, FL, and there are unconfirmed records from Pensacola and Cedar Key, FL. This species is associated with inshore and brackish water habitats. Maximum known size is 65 mm TL. references: Gilbert and Randall 1979; Pezold 1984; Robins and Ray 1986; Smith 1997 (previous references as Gobionellus stigmaturus); SmithVaniz et al. 1999; Murdy and Hoese 2002b.

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GOBIIDAE 695

Elacatinus horsti Yellowline goby

(Metzelaar, 1922)

Elacatinus horsti has a naked body and a narrow light stripe running from eye to caudal fin. Head is slightly depressed and convex between orbits. Rostral frenum is absent. Mouth is U-shaped in dorsal view and subterminal, and upper jaw extends slightly beyond posterior margin of eye. Jaw teeth are villiform and arranged in two rows. Teeth in outer row of upper jaw extend to level of anterior margin of eye, and last two teeth are recurved fangs. Last two or three teeth in inner row of lower jaw are recurved fangs. Tongue is entire. Gill opening extends ventrally the length of pectoral fin base. Measurements are expressed as percent of SL: head length 25.4%–28.1%, pre-dorsalfin length 31.2%–32.7%, pelvic fin length 14.7%–18%, caudal fin length 25.3%–28.4%, body depth 20.2%–25%; and as percent of head length: snout length 15.6%–21%, eye diameter 14.8%–16%. Pectoral fin rays number 18 or 19. First dorsal fin has seven spines, and second has one spine and 10 to 12 rays. Pelvic

fins are united by membrane to form disc. Anal fin has one spine and 10 or 11 rays. Caudal fin is truncate or rounded. A single sensory pore is located between anterior margins of eyes, and preopercular canal has two pores, but pore above operculum is absent. Vertebrae number 28: 11 precaudal and 17 caudal. Color is dark brown dorsally, with yellow or white stripe extending from eye to caudal fin base, and light tan ventrally. This species occurs in the western Atlantic from southern and southwestern Florida (off Tampa Bay), the Bahamas, Haiti, the Cayman Islands, and Belize to Curaçao. It is associated with cylindrical and vaselike sponges. Maximum known size is 45.9 mm SL. references: Ginsburg 1933b; Böhlke and Chaplin 1968; Böhlke and Robins 1968; Robins and Ray 1986; Randall 1996; Smith 1997 (all as Gobiosoma horsti); Murdy and Hoese 2002b.

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696 PERCIFORMES

Elacatinus macrodon Tiger goby

(Beebe and Tee-Van, 1928)

Elacatinus macrodon has scales on caudal peduncle and caudal fin base and 13 distinct dark bands on body. Head is moderately compressed. Mouth is terminal, and upper jaw extends to or beyond posterior margin of eye. Upper lip is continuous across symphysis. Jaw teeth are arranged in narrow bands, with outer row enlarged on either side of symphysis. Tongue is truncate. Gill opening extends ventrally the length of pectoral fin base. Measurements are expressed as percent of SL: head length 26%–34%, eye diameter 5%–10%, pre-dorsal-fin length 31%–39%, pectoral fin length 25%–34%, pelvic fin length 19%–28%, body depth 20%–24%. Pectoral fin rays number 16 to 18 (usually 17). First dorsal fin has seven (rarely six) spines, with first spine elongated as a filament in males. Second dorsal fin has one spine and 10 or 11 rays. Pelvic fins are united by membrane to form disc, and pelvic fin spines are joined by frenum. Anal fin has one spine and 8 to 10 rays. Caudal fin is rounded. Body is naked except for patch of ctenoid scales on caudal peduncle and two to four modified scales on caudal fin base. A

single sensory pore is located between anterior margins of eyes, and preopercular canal has two pores. Sensory pore segment above operculum is absent. Vertebrae number 28: 11 precaudal and 17 caudal. Color is nearly transparent, with 13 narrow dark mahoganycolored bands posterior to pectoral fin base. Bands are narrower than interspaces. This species occurs in the western Atlantic from southern Florida and Bermuda to Cuba and Haiti, including the Gulf of Mexico. In the Gulf of Mexico it is known from Sanibel Island to St. Petersburg, FL, and from Isla de Lobos off Veracruz. This species is associated with hard substrates, including rock faces, coral rubble, tide pools, pilings, and large sponges. Maximum known size is 50 mm SL.

references: Ginsburg 1933; Robins 1958 (as Garmannia macrodon); Springer and Woodburn 1960 (as Garmannia macrodon); Böhlke and Robins 1968; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999 (remainder as Gobiosoma macrodon); Murdy and Hoese 2002b.

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GOBIIDAE 697

Elacatinus oceanops Neon goby

Jordan, 1904

Elacatinus oceanops has a naked body and a pale stripe on side. Head is slightly depressed and convex between orbits. Mouth is subterminal, with snout overhanging upper jaw and upper jaw extending about to center of pupil. Upper lip is divided by rostral frenum at symphysis. Jaw teeth in upper jaw are arranged in a single row, and in lower jaw are arranged in three or four rows anteriorly and a single row posteriorly. Males have two or three recurved canines in posterior section of lower jaw. Tongue is truncate. Gill opening extends ventrally the length of pectoral fin base. Gill rakers on first arch number 8 to 10. Measurements are expressed as percent of SL: head length 20.8%–24.2%, pre-dorsal-fin length 30.6%–33.9%, pelvic fin length 10%–12.7%, caudal fin length 14.1%–21.5%, body depth 18.2%–20%; and as percent of head length: snout length 18.8%–21.6%, eye diameter about 26%. Pectoral fin rays number 16 to 19 (usually 16 or 17). First dorsal fin has seven spines, and second has one spine and 10 to 13

(usually 11 or 12) rays. Pelvic fins are united by membrane to form disc. Anal fin has one spine and 9 to 12 (usually 10 to 12) rays. Caudal fin is rounded. A single sensory pore is located between anterior margins of eyes, and preopercular canal has three pores. Vertebrae number 28: 11 precaudal and 17 caudal. Color is dark brown, with blue lateral stripe running from snout to end of caudal fin. This species occurs in the western Atlantic from southeastern Florida and the Florida Keys to Colombia, including the Gulf of Mexico. In the Gulf of Mexico it occurs off Texas, on the Flower Garden Banks, and in Campeche Bay. This species establishes cleaning stations on reefs and removes parasites and dead skin from surfaces of other fishes. Maximum known size is 50 mm TL. references: Ginsburg 1933; Böhlke and Robins 1968; Bright and Cashman 1974; Robins and Ray 1986; Smith 1997; Hoese and Moore 1998 (all as Gobiosoma oceanops); Murdy and Hoese 2002b.

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698 PERCIFORMES

Elacatinus prochilos

Böhlke and Robins, 1968

Elacatinus prochilos has a naked body and a pale stripe equal to eye width that extends the length of body. Head is slightly depressed and convex between orbits. Rostral frenum is absent. Mouth is U-shaped in dorsal view and subterminal. Upper jaw extends slightly beyond posterior margin of eye. Teeth in upper jaw are villiform and arranged in two rows, with some in outer row enlarged in males. Teeth in lower jaw attenuate from a broad band anteriorly to a single series posteriorly, with several inner teeth enlarged in males. Tongue is emarginate. First gill arch has eight very small gill rakers. Measurements are expressed as percent of SL: head length 26%– 30%, snout length 5.6%–7.3%, eye diameter 7.1%– 8.5%, pelvic fin length 15%–18%, caudal fin length 20%–22%, body depth 14%– 18%. Pectoral fin rays number 17 to 19. First dorsal fin has seven spines, and second has one spine and 10 or 11 rays. Pelvic fins are united by membrane to form disc. Anal fin has one

spine and 10 rays. Caudal fin is rounded. A single sensory pore is located between anterior margins of eyes, and preopercular canal has two pores. Sensory pores are lacking above operculum. Vertebrae number 28: 11 precaudal and 17 caudal. Color consists of a mid-dorsal dark brown stripe, a white stripe extending from tip of snout to caudal fin margin, and a broad dark stripe extending below white stripe to ventral midline. White stripe is as broad as eye and often has a bluish cast. Vertical fins and pectoral fin are dusky. This species occurs between 0.6 and 18 m in the western Atlantic from the northern Gulf of Mexico to Belize, including the Greater and Lesser Antilles. It is associated with coral heads. Maximum known size is 40 mm SL.

references: Böhlke and Robins 1968; Smith 1997 (both as Gobiosoma prochilos); Murdy and Hoese 2002b.

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GOBIIDAE 699

Elacatinus saucrus

(Robins, 1960)

Elacatinus saucrus has four or five rows of scales on caudal fin base and a boldly spotted body. Mouth is inferior. Upper lip is continuous across symphysis. Jaw teeth are arranged in narrow bands, with posterior teeth on inner and outer rows enlarged. Tongue is truncate. Gill opening extends ventrally the length of pectoral fin base. Measurements are expressed as percent of SL: head length about 27%, eye diameter about 7%, pelvic fin length about 21%, body depth about 19%. Pectoral fin rays number 15 to 17. First dorsal fin has seven spines, and second has one spine and 10 rays, with first dorsal spine extended as a filament in males. Pelvic fins are united by membrane to form disc, and pelvic fin spines are joined by frenum. Anal fin has one spine and 8 or 9 rays. Caudal fin is rounded. Body is naked except for two scales on side near base of caudal peduncle and four or five rows of scales on caudal peduncle and caudal fin base. A single sen-

sory pore is located between anterior margins of eyes, and preopercular canal has two pores. Sensory pores are lacking above operculum. Vertebrae number 28: 11 precaudal and 17 caudal. Body is straw colored, with three rows of conspicuous dark reddish brown spots surrounded by white figure-8-shaped marks. Dorsal and dorsolateral rows of spots merge under posterior section of second dorsal fin. This species occurs in the western Atlantic from the Florida Keys, the Bahamas, Jamaica, the Virgin Islands, the Lesser Antilles, and Belize. It is not recorded from the Gulf of Mexico, but it may occur there. This species lives among tentacles of living coral. Maximum known size is 16 mm SL. references: Robins 1960 (as Garmannia saucra); Böhlke and Robins 1968 (as Gobiosoma saucrus); Smith 1997 (as Gobiosoma saucrum); Murdy and Hoese 2002b.

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700 PERCIFORMES

Elacatinus xanthiprora Yellowprow goby

(Böhlke and Robins, 1968)

Elacatinus xanthiprora has a naked body and a narrow pale stripe extending the length of body. Head is convex between orbits. Rostral frenum is absent. Mouth is subterminal and U-shaped in dorsal view. Teeth in upper jaw are in several rows, with those in outer row largest, and the two posteriormost teeth are fanglike in males. Teeth in lower jaw are arranged in several rows, with those in outer row needlelike and larger than those in inner rows, and the lateralmost teeth in inner row are fanglike in males. Tongue is truncate. Gill opening extends ventrally to near upper end of pectoral fin base. Measurements are expressed as percent of SL: head length 29%–31%, snout length 6.3%–7.4%, eye diameter 6.9%– 8.3%, pectoral fin length 23%–24%, pelvic fin length 20%–27%, caudal fin length 21%–25%, body depth 20%–23%. Pectoral fin rays number 18 to 20. First dorsal fin has seven spines, and second has one spine and 10 to 12 rays. Pelvic fins are united by membrane to form disc, and pelvic fin spines are connected by frenum. Anal fin has one spine and 10 rays. Cau-

dal fin is rounded. Body is completely naked. Preopercular canal has two sensory pores, and sensory pores above operculum are lacking. Vertebrae number 28: 11 precaudal and 17 caudal. Color is dark gray dorsally and pale gray ventrally, with yellow bar on snout and two lateral stripes on side of body. Upper stripe is narrow and yellow to white, and runs from posterior margin of eye to caudal fin base. Lower stripe is broad and dark, and runs from posterior margin of eye to margin of caudal fin. This species occurs in the western Atlantic from the Florida Keys to Jamaica, including the Gulf of Mexico. In the Gulf of Mexico it is known from the Dry Tortugas and the Florida Middle Grounds. This species is associated with coral reefs and may live in sponges. Maximum known size is 40 mm SL. references: Longley and Hildebrand 1941 (as E. horsti in part); Böhlke and Robins 1968; Williams and Shipp 1980; Robins and Ray 1986; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 701

Evermannichthys spongicola Sponge goby

(Radcliffe, 1917)

Evermannichthys spongicola has a deep caudal peduncle and body patterned with numerous dark bars and saddles. Snout is acute and wedge shaped in profile and triangle shaped in dorsal view. Lips are fleshy, and fold of upper lip is interrupted at symphysis by broad frenum. Jaw teeth are small, pointed, recurved, and arranged in bands that narrow posteriorly. Some teeth along inner row of lower jaw are enlarged. Tongue is free and bilobed anteriorly, with small lobes on margin. Gill opening is straight and slightly longer than pectoral fin base. Measurements are expressed as percent of SL: head length 19%–24%, snout length 3.7%– 4.8%, eye diameter 2.6%–3.5%, upper jaw length 7.2%–9%, pectoral fin length 15%–23%, pelvic fin length 14%–20%, body depth at dorsal fin origin 11%–14%, caudal peduncle depth 8.8%–10%. Pectoral fin is fanlike and has 11 to 13 rays. First dorsal fin has five to seven (usually six or seven) spines, and second has one spine and 10 to 15 rays. Pel-

vic fins are broadly united to form oval-shaped disc, and pelvic fin spines are connected by frenum. Anal fin has one spine and 8 to 10 rays. Body is naked except for three rows of ctenoid scales extending from anal fin insertion to caudal peduncle. Vertebrae number 28 to 35: 12 to 16 precaudal and 16 to 20 caudal. Color is tan to dusky, with 17 dark bars or saddles that are best developed anteriorly. Dorsal, anal, and caudal fins have dark spots. This species occurs in the western Atlantic from North Carolina to Fort Pierce, FL, and from Panama City to the Dry Tortugas and in Campeche Bay in the Gulf of Mexico. It is associated with large sponges such as Speciospongia. Maximum known size is 30 mm SL. references: Longley and Hildebrand 1941 (as E. metzelaari); Böhlke and Chaplin 1968 (as E. metzelaari); Böhlke and Robins 1969; Robins and Ray 1986; Birdsong et al. 1988; Smith 1997; Murdy and Hoese 2002b.

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702 PERCIFORMES

Evorthodus lyricus Lyre goby

(Girard, 1858)

Evorthodus lyricus has dorsal fins close together or joined at base and a lyre-shaped mark on base of caudal fin. Mouth is slightly inferior, upper jaw extends to posterior margin of pupil, and lower jaw is slightly included in upper jaw. Lips are rather thin, and upper lip has a free dorsal margin. Teeth in upper jaw are uniserial, close set, compressed, and truncate or bilobed in females and juvenile males, and widely spaced, conical, or caninelike in adult males. Teeth in lower jaw are partially biserial, with those in outer row close set, compressed, and truncate or bilobed in females and juvenile males, and widely spaced, conical, or caninelike in adult males. Mature females have a series of two or three small, recurved, conical teeth in outer row of lower jaw. Tongue is entire or rounded. Gill opening extends ventrally below pectoral fin base. Measurements are expressed as percent of SL: head length 24%– 28.3%, snout length 5.7%–7.8%, eye diameter 4.6%– 6.2%, upper jaw length 8.7%–10.2%, pre-dorsal-fin length 31.4%–36.2%, pelvic fin length 21.9%–26.1%, caudal fin length 30.6%–50.1%, body depth 21.1%–23.7%. Pectoral fin extends beyond anal fin origin and has 15 to 17 rays. Dorsal fin is divided; first dorsal fin has six spines, and second dorsal fin has one spine and 10 rays. Pelvic fins are united by membrane to form broad disc. Anal fin has one spine and 11 rays. Spinous dorsal and cau-

dal fins of males are considerably longer than those of females. Body is completely covered with ctenoid scales, and head has cycloid scales between posterior margins of orbits and on upper section of operculum. Head has two sensory pores between eyes and one pore medial to posterior naris. Scales in lateral series number 30 to 35. Median scales preceding dorsal fin origin number 15 to 19, and scale rows on operculum number two or three. Vertebrae number 26: 10 precaudal and 16 caudal. Color is tan, with five or six irregular, narrow dark bars and median dark blotches on sides. Lyreshaped mark on base of caudal fin consists of two square dark blotches separated by a pale tan central area. Males have a pink stripe on upper and lower caudal fin lobes. This species occurs in the western Atlantic from Chesapeake Bay to northern South America, including the northern and southern Gulf of Mexico and the Antilles, but it is absent in the Bahamas. This species is associated with muddy substrates in bays and estuaries. Maximum known size is 150 mm SL. references: Ginsburg 1931; Dawson 1967b, 1969b; Hoese and Moore 1977, 1998; Wyanski and Targett 1985; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Castro-Aguirre et al. 1999; Murdy and Hoese 2002b.

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GOBIIDAE 703

Gnatholepis thompsoni Goldspot goby

Jordan, 1904

Gnatholepis thompsoni has a narrow black bar below eye and a gold spot above base of pectoral fin. Head is blunt, and mouth is subterminal. Lower jaw is included in upper jaw, and upper jaw extends to or slightly posterior to anterior margin of eye. Lips are moderately fleshy, and upper lip is continuous across symphysis. Jaw teeth are arranged in narrow bands, with those on outer row enlarged, slender, and recurved. Upper jaw has one canine tooth on side of symphysis, and lower jaw has one to several tusklike canines projecting from side of mouth. Tongue is distinctly bilobed. Gill opening extends ventrally slightly below pectoral fin base. Gill rakers on first arch number four or five. Measurements are expressed as percent of SL: head length 26.9%–29.5%, snout length 7.9%–10.5%, eye diameter 4.8%– 6%, upper jaw length 7.4%–9.9%, pre-dorsal-fin length 30.2%–33.9%, pelvic fin length 24.7%– 29.4%, caudal fin length 30.1%–35.7%, body depth 20%–22.7%. Pectoral fin has 17 rays. First dorsal fin has six spines, and second has one spine and 11 rays. Pelvic fins are united by membrane to form broad disc, and pelvic fin spines are joined by frenum. Anal fin has one

spine and 11 rays. Body and top and side of head are covered with ctenoid scales. Scales in lateral series number 30 to 32. A pair of sensory pores is located between eyes, and a single pore is medial to posterior naris. Vertebrae number 26: 10 precaudal and 16 caudal. Color is tan, with a narrow bar extending from eye to cheek, bright gold spots outlined with black pigment above pectoral fin, and spots and blotches on upper and lower side, respectively. This species occurs in the tropical Atlantic. In the western Atlantic it occurs from Florida and Bermuda to northern South America, including the Gulf of Mexico, the Florida Keys, the Bahamas, and the Antilles. In the Gulf of Mexico it occurs off southwestern Florida, the Flower Garden Banks, Isla de Lobos (off Veracruz), and Campeche Bank. It is associated with sand, rock, and rubble substrates. Maximum known size is 75 mm TL. references: Böhlke and Chaplin 1968; Bright and Cashman 1974; Castro-Aguirre and MárquezEspinoza 1981; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Murdy and Hoese 2002b.

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704 PERCIFORMES

Gobioides broussonetii Violet goby

Lacepède, 1800

Gobioides broussonetii has a continuous dorsal fin, and dorsal and anal fins are broadly connected to caudal fin. Snout is blunt, and mouth is terminal and very oblique, with lower jaw projecting beyond upper jaw and upper jaw extending to posterior margin of eye. Upper lip is complete across symphysis, expanded posteriorly, and joined with expanded lower lip at rictus. Jaw teeth are caniniform and in two or three rows, with those in outer row of lower jaw enlarged. Tongue is distinctly bilobed. Gill rakers on first arch are short and blunt and number 8 to 11 on lower limb. Measurements are expressed as percent of SL: head length 14.6%–15.6%, snout length 3.3%–3.6%, eye diameter about 1%, upper jaw length 5.3%– 6.3%, pectoral fin length 9%–10%, pelvic fin length 8.8%–11.2%, caudal fin length 25.2%– 28.1%, body depth 7.5%– 8.5%. Pectoral fin is rounded and tapered and has 17 to 20 rays. Dorsal fin has seven spines and 15 rays. Pelvic fins are united by membrane to form long disc,

and pelvic fin spines are joined by frenum. Anal fin has one spine and 15 rays. Caudal fin is lanceolate, with 17 segmented rays and 6 dorsal and 5 or 6 ventral procurrent rays. Body is covered with cycloid scales, and scales are slightly embedded. Scales in horizontal series number about 160. Vertebrae number 27: 10 precaudal and 17 caudal. Color is purplish brown dorsally and pale ventrally. Upper body is variously marked with creamy white patterns. This species occurs in the western Atlantic from Charleston, SC, to Brazil, including the Gulf of Mexico. It is widely distributed in the western and southern Gulf of Mexico. Habitats include both inshore waters and estuaries. Maximum known size is 500 mm SL. references: Dawson 1969b; Hoese and Moore 1977, 1998; Castro-Aguirre 1978; Robins and Ray 1986; Smith 1997; Schaldach et al. 1997; Murdy 1998; Murdy and Hoese 2002b.

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GOBIIDAE 705

Gobionellus oceanicus (Pallas, 1770) Highfin goby

Gobionellus oceanicus has a low keel in front of dorsal fin and a very elongate caudal fin. Mouth is terminal and rather oblique, with lower jaw even with upper jaw or slightly projecting and upper jaw extending to posterior margin of eye in males and to pupil in females. Lips are thin, and dorsal fold of upper lip is complete across symphysis. Jaw teeth are arranged in narrow bands, with those in outer row of upper jaw very slightly enlarged. Teeth in inner row of lower jaw near symphysis are enlarged. Jaw teeth are more enlarged in males than in females. Tongue is slightly emarginate. Gill opening extends to ventral margin of pectoral fin base. Gill rakers on first arch are numerous and occur on both upper and lower limbs. Measurements are expressed as percent of SL: head length 20.1%–22.5%, snout length 5.4%– 6.7%, eye diameter 3%– 4.8%, upper jaw length 8.4%–10.9%, pre-dorsalfin length 25.5%–30.4%, pelvic fin length 16.8%–22.6%, caudal fin length 31%–59.5%, body depth 10.2%–13.3%. Pectoral fin has 19 rays. First dorsal fin has six spines, with second, third, and fourth spines greatly elongated. Second dorsal fin has one spine and 13 rays. Pelvic fins are united by membrane to form broad disc, and pelvic fin spines are joined by frenum. Anal fin has one spine and 13 or 14 rays. Body and part of head are covered with scales. Scales on upper operculum and dorsal surface of head are cycloid, and scales on

body are ctenoid. Cheek generally has a variable number of scattered cycloid scales. Scales in horizontal series number 56 to 76. A pair of sensory pores is located between eyes, and a single pore is medial to posterior naris. Lateral cephalic pores number four. Vertebrae number 26: 10 precaudal and 16 caudal. Color is light brown, slightly darker on dorsal section of head and bases of dorsal fins, and lighter ventrally. A large elongate blotch is located on flank below spiny dorsal fin, and two to four small spots occur on anterior margin of spiny dorsal fin. This species occurs in shallow waters in the western Atlantic from southern Florida to Brazil, including the northern and southern Gulf of Mexico. It is apparently rare in the northern Gulf of Mexico. This species is associated with estuaries and submerged vegetation. Food consists of filamentous algae, ostracods, copepods, and insect larvae. Maximum known size is about 300 mm SL. references: Ginsburg 1932, 1953 (as G. hastatus and G. gracillimus, in part); Springer and Woodburn 1960 (as G. hastatus); Dawson 1969b (as G. hastatus, in part); Hoese and Moore 1977 (as G. hastatus, in part), 1998; Robins and Ray 1986 (as G. hastatus, in part); Pezold and Grady 1989; Boschung 1992; Schaldach et al. 1997 (as G. hastatus, in part); Smith 1997; Castro-Aguirre et al. 1999 (as G. hastatus, in part); Murdy and Hoese 2002b; Pezold 2004b.

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706 PERCIFORMES

Gobiosoma bosc Naked Goby

(Lacepède, 1800)

Gobiosoma bosc has a naked body and 9 to 11 dark bars on side. Head is moderately depressed and appears swollen in dorsal view. Rostral frenum is present. Mouth is terminal and slightly oblique, and upper jaw extends to level of pupil. Jaw teeth are villiform and arranged in bands, with outer row in upper jaw slightly enlarged and inner row of lower jaw ending as two or three recurved fangs. Tongue is truncate. Measurements are expressed as percent of SL: head length 29.2%–31.8%, predorsal-fin length 35.4%–39.4%, pelvic fin length 15%–18.3%, caudal fin length 26.1%– 27.7%, body depth 18.8%–21.2%; and as percent of head length: snout length 23.1%– 25.2%, eye diameter 16.8%–18.7%. Pectoral fin rays number 16 to 19 (usually 17 or 18) rays. First dorsal fin has six to eight spines, and second has one spine and 11 to 13 rays. Pelvic fins are united by membrane to form disc. Anal fin has one spine and 9 to 11 rays. Caudal fin is rounded. A single sensory pore is located be-

tween eyes. Preopercular canal has three pores, and two pores are located above operculum. Vertebrae number 27: 11 precaudal and 16 caudal. Color is light tan to black, with dark bars separated by narrow light bars. This species occurs in coastal areas in the western Atlantic from Massachusetts to Laguna de Términos, Campeche, in the Gulf of Mexico. It is absent off southeastern Florida and the Florida Keys. This species is found in estuaries and protected coastal regions with vegetation, rubble, and oyster beds. Maximum known size is 58 mm SL.

references: Ginsburg 1933b; Springer and Woodburn 1960; Dawson 1966b, 1969b; Böhlke and Robins 1968; Hoese and Moore 1977, 1998; Castro-Aguirre 1978; Robins and Ray 1986; Boschung 1992; Conn and Belcher 1996; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Murdy and Hoese 2002b.

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GOBIIDAE 707

Gobiosoma ginsburgi Seaboard goby

Hildebrand and Schroeder, 1928

Gobiosoma ginsburgi has two ctenoid scales on caudal fin base and about eight vertical bars on body. Head is moderately depressed and appears swollen in dorsal view. Mouth is terminal and slightly oblique, with lower jaw slightly included in upper jaw and upper jaw extending to level of pupil. Jaw teeth are pointed and in narrow bands, with some in outer row enlarged. Tongue is truncate. Gill opening extends ventrally to just above upper end of pectoral fin base. Measurements are expressed as percent of SL: head length 27.2%–28.6%, snout length 5%– 6.2%, eye diameter 5.1%– 7.2%, pectoral fin length 23.3%–25.9%, pelvic fin length 21%–26%, caudal fin length 27.7%–30.1, body depth 17%–22%. Pectoral fin rays number 17 to 20 (usually 18 or 19). First dorsal fin has seven spines, and second has one spine and 11 or 12 rays. Pelvic fins are

united by membrane to form disc. Anal fin has one spine and 9 to 11 (usually 10) rays. Caudal fin is rounded. A single sensory pore is located between eyes. Preopercular sensory canal has three pores, and two pores occur above operculum. Vertebrae number 27: 11 precaudal and 16 caudal. Color is light, with eight irregular dark bars separated by relatively broad light bars. This species occurs from near shore to 50 m in the western Atlantic from Massachusetts to southern Florida, including the eastern Gulf of Mexico at Sanibel Island off southwestern Florida. Maximum known size is 60 mm SL.

references: Hildebrand and Schroeder 1928; Ginsburg 1933b; Dawson 1963b; Robins and Ray 1986; Smith 1997; Murdy and Hoese 2002b.

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708 PERCIFORMES

Gobiosoma grosvenori Rockcut goby

(Robins, 1964)

Gobiosoma grosvenori has posterior section of body scaled and a series of midlateral dashes bisecting dusky bands. Head is moderately depressed. Mouth is terminal and slightly oblique, and upper jaw extends to level of pupil. Jaw teeth are pointed and in narrow bands, with some in outer row enlarged. Tongue is truncate. Measurements are expressed as percent of SL: head length 26%–30%, eye diameter 8%–9%, pectoral fin length 24%–28%, pelvic fin length 20%–22%, body depth 17%– 20%. Pectoral fin rays number 16 to 18. First dorsal fin has seven spines, and second has one spine and 9 rays. Pelvic fins are united by membrane to form disc. Anal fin has one spine and 8 or 9 rays. Caudal fin is rounded. Body is scaled from pectoral fin base to caudal fin base, except for area above line connecting upper end of pectoral fin base and origin of second

dorsal fin, and for area below line connecting lower end of pectoral fin base and anal fin origin. Scales in lateral series number 31 to 35. A single sensory pore is located between eyes. Preopercular sensory canal has three pores, and two pores occur above operculum. Vertebrae number 27: 11 precaudal and 16 caudal. Color is light tan, with nine short dashes along midflank from pectoral fin base to caudal peduncle. Each dash bisects a vertical band on body. This species occurs in the western Atlantic from southeast Florida and Jamaica to Venezuela, including the Gulf of Mexico off Isle de Chien. Maximum known size is 30 mm TL. references: Robins 1964 (as Garmannia grosvenori); Böhlke and Robins 1968; Robins and Ray 1986; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 709

Gobiosoma hemigymnum

(Eigenmann and Eigenmann, 1888)

Gobiosoma hemigymnum has a short barbel below posterior naris, posterior section of body scaled, and eight bars on side. Head is moderately depressed. Rostral frenum is present. Mouth is terminal and slightly oblique, and upper jaw extends to level of pupil. Jaw teeth are pointed and arranged in narrow bands, with most of those in outer row enlarged in upper jaw and most of those in outer row and all teeth near symphysis enlarged in lower jaw. Tongue is truncate. Gill opening extends ventrally from slightly above upper end of pectoral fin base to lower end of base. Pectoral fin rays number 18 to 21. First dorsal fin has seven spines, and second dorsal fin has one spine and 11 or 12 rays. Pelvic fins are united by membrane to form disc, and pelvic fin spines are joined by frenum. Anal fin has one spine and 9 rays. Caudal fin is rounded. Body is scaled from near pectoral fin base to caudal fin base, except for area above line joining upper end of

pectoral fin base and origin of second dorsal fin, and for area below line joining lower end of pectoral fin base and anal fin origin. Scales in horizontal series number about 36. A single sensory pore is located between eyes. Preopercular sensory canal has three pores, and two pores occur above operculum. Vertebrae number 27: 11 precaudal and 16 caudal. Color is light olivaceous, with eight bars running from dorsal midline to midflank and eight black dots along midflank. This species occurs in the western Atlantic from the southern Gulf of Mexico to Rio de Janeiro, Brazil, including the Antilles. In the Gulf of Mexico it is known from the Río de la Máquina, Montepio, Veracruz. Maximum known size is about 48 mm SL. references: Ginsburg 1935; Robins 1960, 1964 (as G. mediocricula); Böhlke and Robins 1968; Castro-Aguirre 1978; Murdy and Hoese 2002b.

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710 PERCIFORMES

Gobiosoma longipala Twoscale goby

Ginsburg, 1933

Gobiosoma longipala has two ctenoid scales on caudal fin base and narrow light bars on body. Head is depressed, flat on top, and concave behind eyes. Mouth is terminal, upper jaw extends beyond posterior margin of eye, and lower jaw is slightly included in upper jaw. Jaw teeth are arranged in two rows, with those in outer row enlarged. Tongue is truncate. Gill opening extends the length of pectoral fin base. Measurements are expressed as percent of SL: head length 26%–35%, pre-dorsal-fin length 33%– 42%, pelvic fin length 18%–28%, caudal fin length 18%–30%, body depth 12%– 18%; and as percent of head length: snout length 15%–24%, eye diameter 18%–29%. Pectoral fin rays number 15 to 17. First dorsal fin has seven spines, and second has one spine and 9 to 12 rays. Pelvic fins are united by mem-

brane to form disc. Anal fin has one spine and 9 rays. Caudal fin is rounded. Body is naked except for scales on upper and lower bases of caudal fin. A single sensory pore is located between eyes. Preopercular sensory canal has three pores, and two pores occur above operculum. Vertebrae number 27: 11 precaudal and 16 caudal. Color is brown, with nine narrow light bars. This species occurs in shallow water in the eastern Gulf of Mexico from Sanibel Island, FL, to Mississippi Sound, MS. Maximum known size is 50 mm TL. references: Ginsburg 1933a,b; Dawson 1963b, 1966a,b; Hoese and Moore 1977, 1998; Böhlke and Robins 1968; Robins and Ray 1986; Boschung 1992; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 711

Gobiosoma robustum Code goby

Ginsburg, 1933

Gobiosoma robustum is naked and has narrow bars on flank. Head is depressed. Upper lip is continuous across symphysis, and upper jaw is moderately protrusible. Mouth is terminal, upper jaw extends to middle of eye in juveniles and to posterior margin of eye in adults, and lower jaw is just slightly included in upper jaw. Jaw teeth are arranged in narrow bands, with outermost row enlarged in upper jaw and innermost row enlarged in lower jaw. Tongue is truncate. Gill opening extends the length of pectoral fin base. Measurements are expressed as percent of SL: head length 29.6%– 31.8%, snout length 5.2%– 6.5%, eye diameter 4.5%–5.1%, pre-dorsal-fin length 33.4%– 37.5%, pelvic fin length 20.5%–22.2%, caudal fin length 27.5%–28.9%, body depth 19.2%– 22.3%. Pectoral fin rays number 16 or 17. Dorsal fin is divided; first dorsal fin has seven spines, and second dorsal fin has one spine and 10 to 12 rays. Pelvic fins are united by mem-

brane to form disc. Anal fin has one spine and 8 to 10 rays. Caudal fin is rounded. A single sensory pore is located between eyes. Preopercular sensory canal has three pores, but pores are absent above operculum. Vertebrae number 27: 11 precaudal and 16 caudal. Color is pale to dusky tan, usually with 10 to 12 dark, irregular, and interrupted bars. This species occurs in shallow water of moderate to high salinity in the western Atlantic from eastern Florida and throughout the Gulf of Mexico. Food consists of copepods, isopods, gammarid amphipods, and shrimps. Maximum known size is 50 mm TL. references: Ginsburg 1933b; Reid 1954; Springer and Woodburn 1960; Dawson 1966b; Böhlke and Chaplin 1968; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith 1997; Castro-Aguirre et al. 1999; Murdy and Hoese 2002b.

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712 PERCIFORMES

Gobulus myersi Paleback goby

Ginsburg, 1939

Gobulus myersi is pale dorsally and dark ventrally, with a very depressed head and no head pores. Mouth is terminal and moderately oblique, with lower jaw projecting beyond upper jaw and upper jaw extending to midpupil. Free margin of upper lip is continuous across symphysis. Upper jaw teeth are arranged in four rows, with about six recurved canines in outer row and low, pointed teeth in inner rows. Lower jaw has four rows of low, pointed teeth. Tongue has a rounded margin. Gill opening has a straight margin and is slightly longer than pectoral fin base. Measurements are expressed as percent of SL: head length 28.4%–31.5%, snout length 5.2%– 6.5%, eye diameter 5.7%– 8.2%, upper jaw length 9.7%–11.9%, pectoral fin length 18.9%–25.3%, body depth at dorsal fin origin 13.2%–20.3%, caudal peduncle depth 9%–10.8%. Pectoral fin has 14 to 16 rays. First dorsal fin has seven spines, and second has one spine and 9 or 10 rays.

Pelvic fins are broadly united by membrane to form complete disc, but pelvic fin spines are not joined by frenum. Anal fin has one spine and 7 to 9 rays. Body is entirely naked. Vertebrae number 27: 11 precaudal and 16 caudal. Color is pale brown dorsally and dark brown ventrally, with two poorly defined dark saddles on back. This species occurs between 5 and 48 m in the western Atlantic from southern Florida and the Bahamas to Venezuela, including the northeastern Gulf of Mexico and the Antilles. In the Gulf of Mexico it has been mainly captured between 28 and 48 m, slightly deeper than it occurs elsewhere. It is associated with coral reefs and shell rubble. Maximum known size is 150 mm SL. references: Ginsburg 1939; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 713

Lophogobius cyprinoides Crested goby

(Pallas, 1770)

Lophogobius cyprinoides has a prominent crest extending from between eyes to near origin of dorsal fin. Head is moderately compressed, and eye is laterally located. Mouth is terminal, with snout overhanging upper jaw and upper jaw extending about to center of pupil. Teeth in upper jaw are arranged in a relatively wide band, with those in outer row enlarged and conical. Tongue is truncate to broadly rounded. Gill opening extends ventrally to isthmus. Measurements are expressed as percent of SL: head length 30.5%–34.4%, pre-dorsal-fin length 31.9%–36.1%, pelvic fin length 24%–27.4%, caudal fin length 26.8%– 35.2%, body depth 26.7%–31%; and as percent of head length: snout length 18.7%– 27.1%, eye diameter 17.8%–30.7%. Body depth increases with growth. Pectoral fin rays number 16 to 20 (usually 17 to 19). First dorsal fin has six spines, and second has one spine and 9 rays. Pelvic fins are united by membrane to form disc. Anal fin has one spine and 9 rays. Caudal fin is rounded. Body is completely

scaled, but head is naked. Preopercular sensory canal has three pores. Single pores are present medial to posterior naris and between eyes. Vertebrae number 26: 10 precaudal and 16 caudal. Color is dark gray to black. Juveniles and young adults have distinctive brown stripes and blotches. Males are uniform in color, with first dorsal fin black and possessing large orange blotches posteriorly. Breeding males are nearly uniformly blackish violet. This species occurs in the western Atlantic from southeastern Florida and Bermuda to northern South America, including the western and southern Gulf of Mexico, the Bahamas, the Greater Antilles, and the southern Lesser Antilles. It is found in shallow protected waters but also occurs in open bays with some wave action. Maximum known size is 100 mm TL. references: Böhlke and Chaplin 1968; Dawson 1972c; Robins and Ray 1986; Smith 1997; CastroAguirre et al. 1999; Smith-Vaniz et al. 1999; Murdy and Hoese 2002b.

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714 PERCIFORMES

Lythrypnus elasson

Böhlke and Robins, 1960

Lythrypnus elasson has scales on posterior section of body, and first two dorsal fin spines are elongated. Mouth is terminal and moderately oblique, with lower jaw projecting slightly beyond upper jaw and upper jaw extending beyond anterior margin of eye. Upper lip is continuous across symphysis. Jaw teeth are arranged in bands, with inner teeth small and conical and outer teeth enlarged, recurved canines. Tongue is free anteriorly and is bluntly V-shaped. Gill opening extends below pectoral fin base. Measurements are expressed as percent of SL: head length 27.4%–33.3%, eye diameter 9.4%–11.1%, pectoral fin length 24.2%–30.8%, pelvic fin length 24.2%– 32.2%, body depth at dorsal fin origin 19.2%– 25%. Pectoral fin has 14 or 15 rays, with 7 or 8 unbranched. First dorsal fin has six spines, with first two spines elongated. Second dorsal fin has one spine and 8 or 9 rays. Pelvic fins are united by membrane to form disc, and pelvic

fin spines are joined by frenum. Inner branch of fourth pelvic fin ray is prolonged. Anal fin has one spine and 8 rays. Scales occur over much of body but are absent on head, nape, chest, pectoral fin base, and area anterior to line running from upper end of pectoral fin base to origin of second dorsal fin. Scales in horizontal series number 25 or 26. Sensory pores are lacking on head. Vertebrae number 26: 10 precaudal and 16 caudal. Color is uniformly reddish gray, with ill-defined bands on dorsal half of body. This species occurs in the western Atlantic from the Bahamas, Cuba, the Cayman Islands, and the northeastern Gulf of Mexico. It is associated with coral reefs between 11 and 26 m. Maximum known size is 20 mm SL.

references: Böhlke and Chaplin 1968; Böhlke and Robins 1960a; Williams and Shipp 1980; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 715

Lythrypnus nesiotes Island goby

Böhlke and Robins, 1960

Lythrypnus nesiotes has prominent bands and scales on posterior part of body. Anterior naris is a long tube located near upper lip, and posterior naris has a raised rim and is near anterior margin of eye. Mouth is terminal and moderately oblique, with lower jaw projecting slightly beyond upper jaw and upper jaw extending beyond anterior margin of eye. Upper lip is continuous across symphysis. Jaw teeth are arranged in bands, with inner teeth villiform and outer teeth enlarged, widely spaced, and recurved. Tongue is free anteriorly, and tip is pointed. Gill opening extends ventrally to below pectoral fin base. Measurements are expressed as percent of SL: head length 28.6%– 32.1%, eye diameter 9.4%–10.9%, pectoral fin length 23.8%–28.6%, pelvic fin length 22.6%–37%, body depth at dorsal fin origin 21.4%–23.7%. Pectoral fin has 15 to 17 rays, with 7 or 8 unbranched. First dorsal fin has six spines, and second dorsal fin has one spine and 9 or 10 rays. Pelvic fins are united by membrane to form disc, and pelvic fin spines are joined by frenum. Inner branch of fourth pelvic fin ray is prolonged. Anal fin has one spine and 8 rays, and several rays may be branched. Scales occur over much of body but are absent

from head, nape, chest, pectoral fin base, and area anterior to line from upper end of pectoral fin base to midlength of first dorsal fin base. Scales in horizontal series number 24 to 26. Sensory pores are lacking on head. Vertebrae number 26: 10 precaudal and 16 caudal. Body is patterned with alternating dark brown and tan bands, with dark line running through middle of tan bands. Side of head is patterned with distinct dark spots, and one or two dark spots are located on pectoral fin base. This species occurs to a depth of 27.4 m in the western Atlantic off southern Florida, the Bahamas, the Greater Antilles, and the Gulf of Mexico. In the Gulf of Mexico it is known from northwestern Florida, Alabama, the Flower Garden Banks, and Isla de Lobos off Veracruz. It is associated with coral reefs. Maximum known size is 20 mm TL.

references: Böhlke and Robins 1960a; Böhlke and Chaplin 1968; Bright and Cashman 1974; Williams and Shipp 1980; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Greenfield 1988; Boschung 1992; Smith 1997; Murdy and Hoese 2002b.

10B-T3324 9/13/05 11:19 AM Page 716

716 PERCIFORMES

Lythrypnus phorellus Convict goby

Böhlke and Robins, 1960

Lythrypnus phorellus has prominent bands and scales on posterior section of body. Mouth is terminal and moderately oblique, with lower jaw projecting slightly beyond upper jaw and upper jaw extending beyond anterior margin of eye. Upper lip is continuous across symphysis. Jaw teeth are arranged in bands, with inner teeth villiform and outer teeth enlarged, widely spaced, and recurved. Tongue is free anteriorly, and tip is bluntly pointed. Gill opening extends below pectoral fin base. Measurements are expressed as percent of SL: head length 26.6%–33.3%, eye diameter 8.3%– 11.3%, pectoral fin length 23%–38.3%, pelvic fin length 27.3%–37.9%, depth at dorsal fin origin 23.7%–25%. Pectoral fin has 14 to 16 rays, with 7 or 8 unbranched. First dorsal fin has six spines, and second has one spine and 8 or 9 rays. Pelvic fins are united by membrane to form disc, and pelvic fin spines are joined by frenum. Inner branch of fourth pelvic ray is prolonged. Anal fin has one spine and 8 or 9 rays. Scales occur over much of body

but are absent from head, nape, chest, pectoral fin base, and area anterior to line from upper end of pectoral fin base to origin of second dorsal fin. Scales in horizontal series number 23 to 26. Sensory pores are lacking on head. Vertebrae number 26: 10 precaudal and 16 caudal. Body is patterned with alternating dark brown and tan bands, with a dark line running through middle of tan bands. Blocklike spots or broken bars radiate from eye onto snout, cheek, and occiput, and two rather faint spots are on pectoral fin base. This species occurs in the western Atlantic from North Carolina to the Florida Keys, in the northern Gulf of Mexico off Texas (Flower Garden Banks), and off the coast of Central America. It is associated with coral reefs to depths from 2 to 34 m. Maximum known size is 20 mm TL. references: Böhlke and Robins 1960a; Robins and Ray 1986; Greenfield 1988; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 717

Lythrypnus spilus Bluegold goby

Böhlke and Robins, 1960

Lythrypnus spilus has first two dorsal fin spines elongated, prominent bands, and scales on posterior section of body. Mouth is terminal and moderately oblique, with lower jaw projecting slightly beyond upper jaw and upper jaw extending beyond anterior margin of eye. Upper lip is continuous across symphysis. Jaw teeth are arranged in bands, with inner teeth villiform and outer teeth enlarged, widely spaced, and recurved. Tongue is short, rounded, and free anteriorly. Gill opening extends below pectoral fin base. Measurements are expressed as percent of SL: head length 27.6%–32.8%, eye diameter 8.8%–11.1%, pectoral fin length 29.3%– 40.4%, pelvic fin length 26.1%–37.9%, body depth at dorsal fin origin 21.1%–25%. Pectoral fin has 15 to 17 rays, with 7 or 8 unbranched. First dorsal fin has six spines, and second has one spine and 8 or 9 rays. Pelvic fins are united by membrane to form disc, and pelvic fin spines are joined by frenum. Inner branch of fourth ray of pelvic fin is prolonged. Anal fin has one spine and 8 or 9 rays. Scales occur over much of body but

are absent from head, nape, chest, pectoral fin base, and area anterior to line running from upper end of pectoral fin base to just behind origin of first dorsal fin. Scales in lateral series number 25 to 27 (usually 26). Sensory pores are lacking on head. Vertebrae number 26: 10 precaudal and 16 caudal. Body is patterned with alternating blue and orange or gold bands, with each blue band subdivided by a dark line. A dark blotch covers entire pectoral fin base. This species occurs in the western Atlantic from southern Florida and Bermuda to Haiti, including the Florida Keys, the northwestern Gulf of Mexico (Flower Garden Banks), and the Bahamas. It is associated with coral reefs, usually at depths from 3 to 26 m. Maximum known size is 25 mm TL. references: Böhlke and Robins 1960a; Böhlke and Chaplin 1968; Bright and Cashman 1974; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Murdy and Hoese 2002b.

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718 PERCIFORMES

Microgobius carri Fowler, 1945 Seminole goby Microgobius carri has a yellow stripe on body. Head is slightly compressed and concave between orbits. Mouth is terminal and moderately oblique, with lower jaw projecting beyond upper jaw, and upper jaw extending slightly beyond eye in females and far beyond eye in mature males. Jaw teeth are arranged in two rows, with those in outer row enlarged and caniniform. Side of lower jaw has four or five teeth in outer row. Tongue is bilobed. Gill opening extends the length of pectoral fin base. Gill rakers on first arch number 5 on upper limb and 15 on lower limb. Measurements are expressed as percent of SL: head length 26%– 31%, snout length 7.4%– 8.9%, eye diameter 6.7%– 8.3%, interorbital width 3%–5%, pectoral fin length 26%–34%, pelvic fin length 15%–21%, caudal fin length 24%–36%, body depth 13.2%–16.7%. Pectoral fin rays number 20 to 24. First dorsal fin has seven spines, and second dorsal fin has one spine and 14 to 17 rays. Second, fourth, and fifth dorsal fin spines of mature males are produced into filaments, with fifth longest. Pelvic fins are united by membrane to form disc. Anal fin has one spine

and 15 to 17 rays. Caudal fin is lanceolate. Body posterior to pectoral fin axil is covered with scales; those below first dorsal fin and on belly are cycloid, and the remainder are ctenoid. Cephalic sensory pores number three, and preopercular canal has two pores. Scales in horizontal series range from 50 to 62. Vertebrae number 27: 11 precaudal and 16 caudal. Color is tan to whitish gray, with two narrow pale yellow stripes on upper side and a broad orangish yellow stripe on lower side. Head is yellow between snout and eyes. This species occurs in the western Atlantic from North Carolina to the eastern Gulf of Mexico off Alabama and the Dry Tortugas, and off the Lesser Antilles. It burrows into coarse sand between 6 and 21 m and is often seen hovering above burrow. This species also forms schools and swims freely in the water column. Maximum known size is 75 mm TL. references: Fowler 1945; Springer and Woodburn 1960; Birdsong 1981; Robins and Ray 1986; Boschung 1992; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 719

Microgobius gulosus Clown goby

(Girard, 1858)

Microgobius gulosus has many dark spots and blotches on body. Head is slightly compressed and concave between orbits. Mouth is terminal and moderately oblique, with lower jaw projecting beyond upper jaw and upper jaw extending slightly beyond eye in females and far beyond eye in mature males. Jaw teeth are arranged in two rows, with those of outer row slightly enlarged and caniniform. Side of lower jaw has seven to nine teeth in outer row. Tongue is bilobed. Gill opening extends the length of pectoral fin base. Gill rakers on first arch number 3 on upper limb and 10 on lower limb. Measurements are expressed as percent of SL: head length 29%–33% in males and 26%– 31% in females, snout length 7.3%–7.5% in males and 5.4%– 6.8% in females, eye diameter 6.5%–7.6%, interorbital width 2.1%– 2.7%, pectoral fin length 21%–28%, pelvic fin length 20.7%–26.5%, caudal fin length 31.7%–34.2% in males and 30.8%–32.5% in females, and body depth 15.4%–19.7%. Pectoral fin rays number 19 to 24 (usually 20 to 23). First dorsal fin has seven spines, and second dorsal fin has one spine and 14 to 17 (usually 14 to 16) rays. Second through sixth dorsal fin spines of mature males are greatly produced into filaments. Pelvic fins are united by membrane to form disc. Anal fin has one spine and 15 to 17 rays. Caudal fin is lanceolate. Body

posterior to pectoral fin axil is covered with scales; those along lateral midline of body, on caudal peduncle, and in patch behind pectoral fin are weakly ctenoid, and the remainder are cycloid. Lateral cephalic sensory pores number two to four (usually three), and preopercular canal has two pores. Scales in horizontal series range from 44 to 54. Vertebrae number 27: 11 precaudal and 16 caudal. Color is tan, with eight or nine dark brown blotches on upper half of body. In males, second dorsal, anal, and caudal fins have a clear stripe and dark margins. In females, body has many irregular dark blotches, and dorsal fins and upper half of caudal fin have dark spots. This species occurs in the western Atlantic from Chesapeake Bay to the southern Gulf of Mexico. It is known throughout the Gulf of Mexico in shallow water. It burrows into muddy to calcareous substrates in protected water ranging from full seawater to freshwater. Food consists of algae, polychaetes, bivalve mollusks, copepods, and gammarid amphipods. Maximum known size is 75 mm TL. references: Springer and Woodburn 1960; Hoese and Moore 1977, 1998; Castro-Aguirre 1978; Birdsong 1981; Robins and Ray 1986; Boschung 1992; Murdy et al. 1997; Smith 1997; CastroAguirre et al. 1999; Murdy and Hoese 2002b.

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720 PERCIFORMES

Microgobius microlepis Banner goby

Longley and Hildebrand, 1940

Microgobius microlepis has a fleshy nuchal crest that is better developed in females than in males. Head is slightly compressed and concave between orbits. Mouth is terminal and relatively oblique, with lower jaw projecting beyond upper jaw and upper jaw extending slightly beyond eye in both sexes. Jaw teeth are arranged in two rows, with those in outer row enlarged and caniniform. Side of lower jaw has five or six teeth in outer row. Tongue is bilobed. Gill opening extends the length of pectoral fin base. Gill rakers on first arch number 4 on upper limb and 12 on lower limb. Measurements are expressed as percent of SL: head length 25%–31%, eye diameter 7%–10%, interorbital width 2.3%–3.5%, upper jaw length about 12%–15% in males and 11%– 14% in females, pre-dorsal-fin length 32%– 33.1%, pelvic fin length 18%–28%, caudal fin length 28%– 40%, body depth 16.1%–17.9%. Pectoral fin rays number 20 to 23 (usually 21 or 22). First dorsal fin has seven spines, and second dorsal fin has one spine and 15 to 19 (usually 17 or 18) rays. Second through sixth dorsal spines of both sexes are slightly produced into filaments. Pelvic fins are united by membrane to form disc. Anal fin has one spine and 17 to 19 rays. Caudal fin is lanceolate.

Body posterior to pectoral fin axil is covered with scales, but scales are absent below first dorsal fin; those on belly are cycloid, and the remainder are weakly ctenoid. Lateral cephalic sensory pores number three (rarely two), and preopercular canal has two pores. Scales in lateral series range from 68 to 78. Vertebrae number 27: 11 precaudal and 16 caudal. Males are pale gray, with a greenish, bluish, or lavender cast on dorsum. Cheeks have alternating pale blue and orangish stripes, dorsal fins are rosy with clear or yellowish stripes near margins, and dorsal section of caudal fin is yellow. Females are dusky, with dark-bordered white triangular pattern on anal fin. This species occurs between 1.5 and 5 m in the western Atlantic from southern Florida and the Bahamas to Belize, including the eastern Gulf of Mexico. It is associated with fine calcareous sands in which it constructs burrows. This species is commonly observed hovering head up and tail down 12 to 25 cm over the mouth of its burrow. Maximum known size is 50 mm TL. references: Böhlke and Chaplin 1968; Birdsong 1981; Robins and Ray 1986; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 721

Microgobius thalassinus Green goby

(Jordan and Gilbert, 1883)

Microgobius thalassinus is dusky green to bluish. Head is slightly compressed and concave between orbits. Mouth is terminal and relatively oblique, with lower jaw projecting beyond upper jaw and upper jaw extending slightly beyond eye in both sexes. Jaw teeth are arranged in two rows, with those in outer row enlarged and caniniform. Side of lower jaw has seven to nine teeth in outer row. Gill opening extends the length of pectoral fin base. Gill rakers on first arch number about 5 on upper limb and about 16 on lower limb. Measurements are expressed as percent of SL: head length 24%–28%, snout length 3.9%–5.7%, eye diameter 5.6%–7.8%, interorbital width 1.8%–3%, upper jaw length 12%–15% in males and 11%–14% in females, pre-dorsalfin length 30.3%–32.4%, pelvic fin length 18%–28%, caudal fin length 28%– 40%, body depth 16%–18.2%. Pectoral fin rays number 19 to 23 (usually 20 to 22). First dorsal fin has seven spines, and second dorsal fin has one spine and 14 to 16 (usually 15) rays. Dorsal spines of males are slightly produced into filaments. Pelvic fins are united by membrane to form disc. Anal fin has one spine and 14 to 16 (usually 15 or 16) rays. Caudal fin is lanceolate. Body posterior to pectoral fin axil is covered with scales; those behind pectoral fin and

occasionally on caudal peduncle are weakly ctenoid, and the remainder are cycloid. Scales are absent below anterior part of first dorsal fin. Lateral cephalic sensory pores number two, and preopercular canal has two pores. Scales in lateral series range from 43 to 50. Vertebrae number 27: 11 precaudal and 16 caudal. Males have two dark bars separated by a pale interspace on shoulder, dark spots with a clear submarginal stripe on first dorsal fin, and dusky pigment with a clear stripe on second dorsal fin. Females have an unspotted first dorsal fin with a dark margin. This species occurs in the western Atlantic from Chesapeake Bay to the northern Gulf of Mexico. However, it is apparently absent off southern Florida between Cape Canaveral and the Florida Keys. In the Gulf of Mexico it is present off Texas and the Tampa Bay region of Florida. This species is associated with shallow, muddy sand and muddy-bottomed protected areas in brackish to full-salinity waters. Maximum known size is 40 mm SL. references: Ginsburg 1934; Springer and Woodburn 1960; Dawson 1969b; Hoese and Moore 1977, 1998; Birdsong 1981; Robins and Ray 1986; Boschung 1992; Murdy et al. 1997; Smith 1997; Murdy and Hoese 2002b.

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722 PERCIFORMES

Nes longus (Nichols, 1914) Orangespotted goby

Nes longus has brown and orange spots on body and fins and lacks pores on head. Head is moderately compressed. Upper lip is continuous across symphysis. Mouth is terminal, and upper jaw extends slightly beyond middle of eye. Teeth in upper jaw are in two rows, with those in outermost row enlarged. Teeth in lower jaw are small and in a narrow band. Tongue is free, truncate or slightly indented anteriorly, and relatively long and narrow. Gill opening is straight and extends above and below pectoral fin base. Measurements are expressed as percent of SL: head length 21.3%–21.5%, eye diameter 4.9%–5.1%, upper jaw length 9.8%– 19%, pelvic fin length 12.7%–14.9%, body depth 13%–16.7%. Pectoral fin rays number 16 to 19. First dorsal fin has seven spines, and second dorsal fin has one spine and 12 or 13 rays. Pelvic fins are united by membrane to form disc, and pelvic fin spines are joined by frenum. Anal fin has one spine and 10 to 12 (usually 12) rays. Caudal fin is elongate and

rounded. Body is naked. Vertebrae number 28: 11 or 12 precaudal and 16 or 17 caudal. Color is pale gray to tan, with irregular yellowish brown spots on head and upper body and a row of five to seven dark brown blotches on lower side. Caudal fin has chevron markings. Males have a dusky pelvic fin and a banded caudal fin. This species occurs from shore to 9 m in the western Atlantic from southern Florida and Bermuda to Venezuela, including the Gulf of Mexico, the Bahamas, Puerto Rico, and the islands off Venezuela. In the Gulf of Mexico it occurs at the Dry Tortugas and Alacrán Reef off Yucatán. It lives with alphaeid shrimp in burrows. Maximum known size is 100 mm TL. references: Ginsburg 1933a; Böhlke and Chaplin 1968; Böhlke and Robins 1968; Robins and Ray 1986; Randall 1996; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 723

Oxyurichthys stigmalophius Spotfin goby

(Mead and Böhlke, 1958)

Oxyurichthys stigmalophius has most of body covered with scales and a large dark spot on posterior section of first dorsal fin. A low membranous crest runs from nape above preoperculum to origin of dorsal fin. Mouth is terminal and slightly oblique, with upper jaw extending to about middle of eye. Lips are thin, and upper lip is free at symphysis. Jaw teeth are conical, slightly recurved, and arranged in a single row in upper jaw and in two rows in lower jaw. Tongue is rounded. Gill opening extends ventrally below pectoral fin base. Measurements are expressed as percent of SL: head length 26.8%–27.7%, snout length 8.7%–12.6%, eye diameter 5.7%– 6.7%, upper jaw length 11.8%–13.7%, pectoral fin length 20.9%–22.7%, pelvic fin length 24.5%– 27.6%, caudal fin length 39.8%–54.9%, body depth 18.4%–19.6%. Pectoral fin has 21 or 22 rays. First dorsal fin has six spines, and second dorsal fin has one spine and 12 rays. Pelvic fins are united by membrane to form broad disc, and pelvic fin spines are joined by frenum.

Anal fin has one spine and 13 rays. Posterior section of body above midline is covered with ctenoid scales, and the remainder of body is covered with cycloid scales. Scales are absent on cheek, operculum, and nape. A pair of sensory pores is located between eyes. Scales in horizontal series number 89 to 102. Color in life is dusky brown, with four large blotches and marbling on side. Posterior section of first dorsal fin bears a large black blotch. This species occurs in the western Atlantic from Florida and the Bahamas to Suriname, including the southern Gulf of Mexico. It is associated with sandy and muddy bottoms between 2 and 60 m and apparently shares a burrow with an alphaeid shrimp. Maximum known size is 165 mm TL. references: Mead and Böhlke 1958; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997 (all as Gobionellus stigmalophius); Murdy and Hoese 2002b.

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724 PERCIFORMES

Palatogobius paradoxus Mauve goby

Gilbert, 1971

Palatogobius paradoxus has a pair of large canine teeth in each jaw, a relatively long second dorsal fin, and scales limited to posterior part of body. Mouth is large and moderately oblique, with lower jaw projecting beyond upper jaw and upper jaw extending to center of pupil. Upper lip is complete over symphysis. Jaw teeth are arranged in two rows, with those in outer row slightly enlarged. Upper jaw has a large canine on side of symphysis, and lower jaw has a large, outwardly directed canine at about midlength. A patch of pointed teeth often occurs in vomer. Tongue is rounded and partially free of floor of mouth. Gill opening extends from upper end of pectoral fin base to below posterior margin of orbit. Measurements are expressed as percent of SL: head length 22.6%–29%, snout length 5.8%–7.8%, eye diameter 7.4%– 8.7%, upper jaw length 8.7%–10.6%, pectoral fin length 16.8%– 19.8%, pelvic fin length 18.7%–28.5%, body depth 14.8%–16%. Pectoral fin has 18 to 22 rays. First dorsal fin has seven rays, and second dorsal fin has one spine and 17 to 20 rays. Dorsal fins are joined at base by membrane. Pelvic fins are joined by membrane to form

disc, but pelvic fin spines are not connected by frenum. Anal fin has one spine and 20 rays. Caudal fin is lanceolate. Scales on body are limited to posterior fourth or fifth of body. Sensory pores on head consist of one pore medial to posterior naris, one pore between anterior margins of orbits, one pore between posterior margins of orbits, and two pores posterior to orbits. Vertebrae number 27: 11 precaudal and 16 caudal. Color is pale blue dorsally and mauve ventrally, with a black stripe separating colors. Snout, cheek, and operculum are mauve; dorsal fin is yellow; and caudal fin is yellow with a blue center. This species occurs in the western Atlantic from the northeastern Gulf of Mexico, the Lesser Antilles, and the coasts of Panama and Venezuela. It is associated with isolated coral heads and coral rubble on sandy bottoms to a depth of 67 m. Maximum known size is 35 mm SL.

references: Gilbert 1971b, 1977; Robins and Ray 1986; Birdsong 1988; Boschung 1992; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 725

Priolepis hipoliti (Metzelaar, 1922) Rusty goby

Priolepis hipoliti has second dorsal spine elongated and scales on head extending to interorbital region. Mouth is terminal and moderately oblique, with lower jaw projecting slightly beyond upper jaw and upper jaw extending beyond anterior margin of eye. Upper lip is continuous across symphysis. Jaw teeth are arranged in bands, with outer teeth enlarged canines and inner teeth villiform and failing to reach corner of mouth. Tongue is free anteriorly and emarginate. Gill opening extends ventrally to level of preoperculum. Gill rakers on first arch number 15 or 16. Measurements are expressed as percent of SL: head length 29.9%– 34.4%, eye diameter 9.2%–12.8%, pectoral fin length 25.1%–35.9%, pelvic fin length 23.3%–31.4%, body depth at dorsal fin origin 23.3%–25%. Pectoral fin has 16 to 19 (usually 17 or 18) rays. First dorsal fin has six spines, and second dorsal fin has one spine and 8 to 10 rays. Pelvic fins are only united basally by membrane, and spines are not connected by frenum. Anal fin has one spine and 7 to 9 rays. Ctenoid scales cover body and occipital re-

gion of head. Scales in horizontal series number 27 to 32 (usually 28 to 31). Sensory pores are lacking on head. Vertebrae number 26: 10 precaudal and 16 caudal. Color is rusty or dull orange on body and head, with poorly defined pale bars. Fins are covered with orange spots arranged in oblique series on pectoral, dorsal, and anal fins and in vertical series on caudal fin. This species occurs in the western Atlantic from southern Florida and Bermuda to northern South America, including the Florida Keys, the Gulf of Mexico, and the Bahamas. In the Gulf of Mexico it has been captured at the Flower Garden Banks and Veracruz. This species is associated with coral reefs and rocky areas between 30 and 100 m. Maximum known size is 41 mm TL.

references: Parr 1930 (as Gobius ebriosus); Böhlke and Chaplin 1968 (as Quisquilius hipoliti); Bright and Cashman 1974 (as Q. hipoliti); Robins and Ray 1986 (as Q. hipoliti); Greenfield 1989; Randall 1996; Smith 1997; Smith-Vaniz et al. 1999; Murdy and Hoese 2002b.

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726 PERCIFORMES

Ptereleotris calliura Blue goby

(Jordan and Gilbert, 1882)

Ptereleotris calliura has nearly separate pelvic fins and a long, pointed caudal fin. Profile of head is moderately convex. Eye is large. Mouth is slightly superior and strongly oblique, with lower jaw projecting beyond upper jaw and upper jaw extending to anterior margin of eye. Upper lip lacks frenum. Jaw teeth are moderately large to large, caninelike, and arranged in two rows. In upper jaw, teeth in outer row are enlarged and slightly recurved, and those in inner row are small except for large tooth at symphysis. In lower jaw, teeth in outer row are small and close set, and those in inner row are very large, recurved, and widely spaced. Vomer and palatine have blunt teeth. Tongue is narrow and largely free of floor of mouth. Gill membranes are connected and jointed to isthmus at level of preoperculum. Gill rakers on first arch are long and slender and number about 30. Measurements are expressed as percent of SL: head length 19.8%–21.3%, snout length 4%– 4.4%, eye diameter 4.5%–5.8%, upper jaw length 7.2%– 8.1%, pre-dorsal-fin length 25.4%–26.7%, pre-anal-fin length 50.5%–53.4%, body depth 13.1%–15.9%.

Pectoral fin is fan shaped and has 20 rays, and all but outermost rays are branched. First dorsal fin has six slender spines and is partially connected to second dorsal fin. Second dorsal fin has one spine and 21 or 22 rays. Pelvic fin has one spine and 4 rays. Anal fin has one spine and 21 rays. Caudal fin is lanceolate, and median rays are elongated into a filamentous extension. Body is covered with very small embedded scales. Color is bluish gray to lavender in life, with a black stripe near margin of two dorsal fins and black pigment on anal fin and lower half of caudal fin. Color in preservative is tan, with black submarginal stripes on dorsal fins and black pigment on anal and caudal fins. This species occurs in the western Atlantic from North Carolina to southern Florida and the northern and western Gulf of Mexico. It inhabits burrows in shell rubble and sand around coral reefs between 5 and 50 m. Maximum known size is 125 mm TL. references: Hoese and Moore 1977, 1998; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Murdy and Hoese 2002b.

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GOBIIDAE 727

Risor ruber (Rosén, 1911) Tusked goby

Risor ruber has elongated, recurved canine teeth in jaws and scales limited to posterior section of body. Head is abruptly elevated, and snout is very blunt. Upper lip is interrupted medially by broad frenum. Mouth is subterminal, and upper jaw extends to or beyond pupil. Upper jaw has two to four large canine teeth recurved outward at symphysis, flanked by four or five smaller recurved canines, and an inner row of small pointed teeth. Lower jaw has two to four large canines recurved backward, flanked by three smaller recurved canines, and one to several rows of small pointed teeth. Large canines at symphyses of upper and lower jaws project from mouth. Teeth are absent on vomer and palatine. Tongue is truncate. Gill opening extends the length of pectoral fin base. Measurements are expressed as percent of SL: head length 25%–30%, eye diameter 7%–9%, upper jaw length 8%–10%, pectoral fin length 23.9%–33.8%, pelvic fin length 22%–32%, body depth 18%–26%. Pectoral fin rays number 16 or 17. First dorsal fin has seven spines, and second has one spine and 10 or 11 rays. Pelvic fins are united by membrane to form

disc, and pelvic fin spines are connected by frenum. Anal fin has one spine and 8 or 9 rays. Caudal fin is rounded and has 16 or 17 rays. Posterior aspect of body, between anal fin origin and caudal fin base, is covered with scales. Caudal fin base has four or five scales bearing enlarged ctenii. Scales in horizontal series number 14 to 19. Preopercular sensory pores are absent. Vertebrae number 27 or 28: 11 or 12 precaudal and 16 caudal. Color is pale to dark brownish gray or blackish, often with obscure dark bars. This species occurs in the western Atlantic from southern Florida and the Bahamas to Suriname, including the Gulf of Mexico. In the Gulf of Mexico it occurs along the west coast of Florida, off Alabama, and on the Flower Garden Banks. It is associated with large sponges such as loggerheads. Maximum known size is 25 mm TL. references: Böhlke and Chaplin 1968; Böhlke and Robins 1968; Bright and Cashman 1974; Williams and Shipp 1980; Robins and Ray 1986; Boschung 1992; Smith 1997; Murdy and Hoese 2002b.

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728 PERCIFORMES

Sicydium gymnogaster

Ogilvie-Grant, 1884

Sicydium gymnogaster has short pelvic fins that are largely attached to belly. Head is moderately depressed, and snout is blunt. Mouth is inferior, and upper jaw extends to pupil. Upper lip fold is broad at corner of mouth and interrupted medially by broad frenum. Lower lip fold is limited to corner of lower jaw. Upper jaw teeth are elongate, very slender, bicuspid, and arranged in a single row. Teeth in lower are incisor-like, slightly recurved, and arranged in a single row. Gill slit extends the length of pectoral fin base. Measurements are expressed as percent of SL: head length 21.6%–22.6%, snout length 7.8%–11.2%, eye diameter 4.3%– 4.7%, upper jaw length 8.5%–10%, pelvic fin length 12%–13.9%, body depth 14.7%–17.5%. Pectoral fin rays number 17 to 20. First dorsal fin has six spines, and second dorsal fin has one spine and 9 or 10 rays. Pelvic fins are united to form a broad,

oval-shaped disc. Anal fin has one spine and 10 rays. Caudal fin is truncate to slightly rounded. Body posterior to axil of pectoral fin is covered with scales except for belly. Scales on upper side anterior to dorsal fin, behind pectoral fin, and on lower side are cycloid and partially embedded; those on remainder of body are ctenoid. Scales in horizontal series number 66 to 68. Vertebrae number 26: 10 precaudal and 16 caudal. Color in preservative is yellowish tan, with eight bars on sides, and light areas between first two or three bars are reticulated with dark brown. This species occurs in the western Atlantic from the southern Gulf of Mexico at Veracruz and off Honduras. Maximum size is 130 mm TL. references: Jordan and Evermann 1898; CastroAguirre et al. 1999; Murdy and Hoese 2002b.

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GOBIIDAE 729

Varicus marilynae

Gilmore, 1979

Varicus marilynae has an enlarged head, relatively long pelvic fins, and a basal scale sheath on caudal fin. Upper lip is continuous across symphysis, and rostral frenum is lacking. Mouth is terminal and moderately oblique, with upper jaw extending to pupil of eye. Teeth in jaws are in three or more rows anteriorly, with five or six teeth in outer row of each jaw enlarged. Tongue is slightly bilobed. Gill opening is restricted to the length of pectoral fin base. Measurements are expressed as percent of SL: head length 30.1%–31.1%, snout length 5%–5.6%, eye diameter 8.9%–9.4%, upper jaw length 11.1%–11.7%, pectoral fin length 30.6%–33.3%, pelvic fin length 25%–39%, body depth 17.8%–19.4%. Pectoral fin has 16 to 18 rays. First dorsal fin has seven spines, and second has one spine and 8 rays. Pelvic fins are connected by low membrane, but pelvic fin

spines are not joined by frenum. First 4 pelvic fin rays are bifurcate and flattened distally, and bifurcate tips of first 3 rays are coalesced into a fleshy, flattened pad. Fifth pelvic fin ray is greatly reduced and close to fourth ray. Anal fin has one spine and 7 rays. Caudal fin is rounded. Body is covered with ctenoid scales except for nape and pectoral fin base. Color in life is reddish orange on lower part of head and belly, with nine brownish green bars on head and body. Pectoral fin is orange. This species occurs in the western Atlantic from the southern and western coasts of Florida. In the Gulf of Mexico it occurs off Cape St. George, FL, between 61 and 91 m. Maximum known size is 25 mm SL. references: Gilmore 1979; Greenfield 1981; Smith 1997; Murdy and Hoese 2002b.

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730

MICRODESMIDAE

Wormfishes

Wormfishes are elongate to very elongate and compressed to very compressed, with a single very long dorsal fin, a prominent chin, and small pelvic fins that are usually separate. Snout is blunt, and mouth is terminal to superior and moderately oblique to nearly vertical. Eye is large and laterally located. Jaw teeth are caniniform and in several rows. Teeth are present or absent in vomer and absent in palatine. Gill opening is laterally located. Branchiostegal rays number 5. Pectoral fin is moderate to well developed and consists of 10 to 16 rays. Dorsal fin consists of 10 to 28 flexible spines and 9 to 66 simple or branched rays. Pelvic fin inserts below pectoral fin base and has 1 spine and 2 to 5 rays. Anal fin is long and consists of 0 or 1 spine and 9 to 61 rays. Caudal fin is either free or joined to dorsal and anal fins. Scales are either cycloid, embedded, and usually nonoverlapping, or cycloid anteriorly and ctenoid posteriorly and embedded. Some species lack scales. Scales in horizontal series number 26 to 76. Lateral line is absent, and sensory pores on head are present or absent. Wormfishes occur worldwide in tropical to warm temperate seas. They burrow into soft muddy and sandy bottoms. There are about 30 species in five genera, 6 species in two genera occur in the western Central Atlantic, and 4 species in two genera occur in the Gulf of Mexico. references: Dawson 1974; Fritzsche and Fuiman 1982; Nelson 1994; Randall 1996; Smith 1997; Smith-Vaniz et al. 1999; Thacker 2000, 2002; Larson 2001; Amaya et al. 2002.

Key to the Species of the Gulf of Mexico (Adapted, in part, from Dawson 1977) 1a. Body slender and very elongate, depth usually between 3% and 9% of SL; gill opening moderate but not tubiform; scales well separated in adults; pectoral fin rays 10 to 14 .............................................2 1b. Body short and relatively stubby, depth usually 8% to 12% of SL; gill opening usually tubiform; scales separated or close set to narrowly imbricate; pectoral fin rays 11 to 15 . . . . . . . . . . . . . . . . . . . . . . . . . . . Cerdale floridana p. 731 2a. Dorsal fin spines 12; anal fin rays 55 or 56 . . . . . . . . . . . . . . . . . . . . Microdesmus lanceolatus p. 733 2b. Dorsal fin spines 19 to 27; anal fin rays 34 to 48 . . . . . . . 3 3a. Dorsal fin spines 19 to 22; anal fin rays 37 to 48; anal fin origin below dorsal fin elements 25 to 31 . . . . . . . . . . . . . . . . . . . . . Microdesmus longipinnis p. 734 3b. Dorsal fin spines 24 to 27; anal fin rays 34 to 40; anal fin origin below dorsal fin elements 33 to 38 . . . . . . . . . . . . . . . . . . . . . . . . . . Microdesmus carri p. 732

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MICRODESMIDAE 731

Cerdale floridana Longley, 1934 Rugjaw wormfish

Cerdale floridana is relatively short and stubby. Snout and lower jaw are rounded. Anterior naris is a short, widely flared tube, and posterior naris is porelike with a raised rim. Lower jaw projects beyond upper jaw, and mouth is oblique. Interorbital width is slightly less than eye diameter. Lower lip is entire and not notched below anterior naris. Jaw teeth are arranged in two rows, with those in outer row slightly larger than those in inner row. Tongue is truncate. Gill opening is a tubiform prolongation suspended from anteroventral margin of pectoral fin and is about 50% to 67% of eye diameter. Measurements are expressed as percent of SL: head length 13%–20.2%, pectoral fin length 6.8%–12.5%, pelvic fin length 4.9%–12.5%, body depth 7.7%–12.2%; and as percent of head length: snout length 17.9%– 28.5%, eye diameter 12.5%–19.1%, upper jaw length 17.9%–25%. Pectoral fin has 13 to 15 (usually 13 or 14) rays. Dorsal fin originates anterior to tip of appressed pectoral fin

and has 12 to 14 spines and 30 to 34 (usually 31 to 33) rays. Pelvic fin has 1 spine and 3 unbranched rays. Anal fin has 28 to 33 (usually 30 or 31) rays. Dorsal and anal fins are generally connected to caudal fin by a low membrane. Body is covered with small cycloid scales except for tip of snout and lower jaw. Adult squamation is not reached until 34 to 40 mm SL. Color is pale tan to yellowish green, with small dark spots. This species occurs in the western Atlantic from southern Florida and Bermuda to Colombia, including the eastern Gulf of Mexico at the Dry Tortugas, the Bahamas, and the Greater and Lesser Antilles. It is associated with coral reefs and coastal areas to 30.5 m. Maximum known size is 79.6 mm SL. references: Longley 1934; Longley and Hildebrand 1941 (as Microdesmus floridanus); Dawson 1974; Robins and Ray 1986; Smith 1997; SmithVaniz et al. 1999; Thacker 2002.

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732 PERCIFORMES

Microdesmus carri Gilbert, 1966

Microdesmus carri is slender and elongate, with a stripe running the length of dorsum. Snout is moderately blunt, and lower jaw is strongly rounded. Anterior naris is a short, widely flared tube located near upper lip, and posterior naris is porelike, with a raised rim, and located anterior to eye. Lower jaw projects beyond upper jaw, upper jaw extends to anterior margin of eye, and mouth is oblique. Interorbital width is about equal to half of eye diameter. Lower lip is entire and not notched below anterior naris. Jaw teeth are conical and arranged in two irregular series. Tongue is rounded. Gill opening is oblique and about 150% of pectoral-fin-base length. Measurements are expressed as percent of SL: head length 10%–16%, pectoral fin length 4.3%– 7.1%, body depth 4.9%–9%. Pectoral fin has 13 rays. Dorsal fin originates anterior to tip of appressed pectoral fin and has 25 or 26 spines and 41 or 42 rays. Pelvic fin has 1 spine and 3 unbranched rays. Anal fin has 33 to 36 rays.

Dorsal and anal fins are generally connected to caudal fin by a low membrane. Upper half of body is sparsely covered with minute, embedded, nonimbricated scales. Head has few scales. Color is pale, with three streaks around eye and a stripe on dorsal section of body, and upper two-thirds of body is densely pigmented with melanophores. Small specimens are unpigmented except for streaks around eye and flecks of pigment on belly. This species occurs in the western Atlantic in the southern Gulf of Mexico at Veracruz and off the coast of Costa Rica. It is associated with low- to moderately low-salinity lagoons on muddy and sandy bottoms. Populations from the Gulf of Mexico and Belize may represent an undescribed species. Maximum known size is 53 mm SL. references: Gilbert 1966a; Dawson 1972c; Smith 1997; Castro-Aguirre et al. 1999; Smith and Thacker 2000; Thacker 2002.

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MICRODESMIDAE 733

Microdesmus lanceolatus Lancetail wormfish

Dawson, 1962

Microdesmus lanceolatus is slender and elongate, with a lanceolate caudal fin and small spots along anal fin. Snout is moderately blunt, and lower jaw is strongly rounded. Anterior naris is a short, widely flared tube located near upper lip, and posterior naris is porelike, with a raised rim, and located near anterior margin of eye. Lower jaw projects slightly beyond upper jaw, upper jaw extends to about middle of eye, and mouth is oblique. Interorbital distance is about equal to eye diameter. Lower lip is entire and not notched below anterior naris. Jaw teeth are conical and arranged in two irregular series. Tongue is rounded. Gill opening is oblique and about 150% of pectoral-finbase length. Measurements are expressed as percent of SL: head length about 12.6%, pectoral fin length about 6.2%, pelvic fin length about 5.5%, depth about 6.6%; and as per-

cent of head length: snout length about 24.6%, eye diameter about 8.8%. Pectoral fin has 13 rays. Dorsal fin originates anterior to tip of appressed pectoral fin and has 12 spines and 56 rays. Pelvic fin has 1 spine and 3 unbranched rays. Anal fin has 55 rays. Dorsal and anal fins are generally connected to caudal fin by a low membrane. Body is sparsely covered with minute, embedded, nonimbricated scales. Head is naked. Color is pale straw, with very small black spots along base of anal fin. This species occurs in the western Atlantic from the northern Gulf of Mexico along Louisiana and Texas. Maximum known size is 45.3 mm SL. references: Dawson 1962a; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith 1997; Thacker 2002.

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734 PERCIFORMES

Microdesmus longipinnis Pink wormfish

(Weymouth, 1910)

Microdesmus longipinnis is slender and elongate, with a rounded caudal fin and pinkish tan coloration. Snout is moderately blunt, and lower jaw is strongly rounded. Anterior naris is a short, widely flared tube located near upper lip, and posterior naris is porelike, with a raised rim, and located near anterior margin of eye. Lower jaw projects slightly beyond upper jaw, upper jaw extends to about middle of eye, and mouth is oblique. Interorbital width is about equal to eye diameter. Upper and lower lips form moderately developed lateral flange at corner of mouth, and lower lip is entire and not notched below anterior naris. Jaw teeth are conical and arranged in two irregular series. Tongue is rounded. Gill opening is oblique and about 150% of pectoral-fin-base length. Measurements are expressed as percent of SL: head length 5.9%–16.2%, pre-anal-fin length 47.5%– 60.8%, body depth 4.2%– 8.1%. Relative preanal length and depth decrease with

growth. Dorsal fin originates anterior to tip of appressed pectoral fin and has 19 to 22 (usually 20 or 21) spines and about 45 to 53 rays. Pelvic fin has 1 spine and 3 unbranched rays. Anal fin has 41 to 47 rays. Dorsal and anal fins are generally connected to caudal fin by low membrane. Body is sparsely covered with minute, embedded, nonimbricated scales. Head is naked. Color is pinkish tan. This species occurs in the tropical Atlantic. In the western Atlantic it occurs from North Carolina and Bermuda to the Cayman Islands in the Lesser Antilles, including the Gulf of Mexico. In the Gulf of Mexico it occurs from Florida to Texas and Veracruz. Maximum known size is 257 mm SL. references: Dawson 1962a; Robins and Ray 1986; Boschung 1992; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Thacker 2002.

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735

EPHIPPIDAE

Spadefishes

Spadefishes are deep bodied, oblong or orbicular, and strongly compressed, with a short head; a series of blunt, comblike gill rakers on first arch; and slender, movable, and brushlike jaw teeth. Dorsal profile of head is parabolic. Nostril is paired. Mouth is terminal and small, and upper jaw is not protrusible. Jaw teeth are setiform or compressed, tricuspid, and arranged in bands. Palatine lacks teeth, and vomer either has or lacks teeth. Gill membranes are broadly joined to isthmus. Gill rakers on upper limb of first arch are comblike. Preoperculum is smooth or serrate. Branchiostegal rays number 6. Pectoral fin is of moderate size and rounded and inserts on lower half of flank. Dorsal fin is usually notched but occasionally entire, and has five to nine spines and 18 to 38 rays, with spines long in juveniles but short in adults. Pelvic fin has a large axillary process at base, inserts slightly in front of pectoral fin base, and has one spine and 5 rays. Anal fin has three spines and 15 to 27 rays. Caudal fin is truncate and consists of 17 principal rays. Scales are smooth or ctenoid and small to moderate in size and occur on body and on rayed sections of dorsal and anal fins. Scales are lacking on head anterior to eye. Lateral line is complete. Vertebrae number 24: 10 precaudal and 14 caudal. Gas bladder is present. Juveniles have several black bands around body, but these are lost with growth. Maximum known size is 50 cm TL. Spadefishes occur worldwide in tropical to warm temperate seas. They occur in schools and are associated with coral reefs, rocky areas, artificial reefs, and estuaries. Food consists of invertebrates, especially mollusks. Juveniles are pelagic, and juveniles of some species resemble floating leaves and bark, thereby avoiding predation. There are about 20 species in seven genera, and 1 species occurs in the western Central Atlantic, including the Gulf of Mexico. references: Fritzsche and Fuiman 1982; Johnson 1984; Smith 1987; Nelson 1994; Smith 1997; Heemstra 2001b; Burgess 2002c.

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736 PERCIFORMES

Chaetodipterus faber Atlantic spadefish

(Broussonet, 1782)

Chaetodipterus faber has a deeply notched dorsal fin that is nearly separate, and anterior part of rayed section of dorsal fin and anal fin is elongated and filamentous. Dorsal profile of head is moderately convex. Snout is very blunt. Posterior naris is slitlike. Jaw teeth are arranged in bands, with those in outermost band largest, slightly compressed, and sharply pointed. Preopercular margin is finely serrate. Operculum has a broad-based blunt spine on posterior margin. Gill rakers on first arch number 7 or 8 on upper limb and 9 or 10 on lower limb. Measurements are expressed as percent of SL: head length 33%–37%, snout length 13%–14%, eye diameter 9%–10%, upper jaw length 9%–11%, pectoral fin length 19%– 22%, body depth 77%– 82%. Pectoral fin has 17 to 19 rays. Dorsal fin has eight or nine spines

and 21 to 24 rays. Anal fin has three spines and 18 or 19 rays. Caudal fin is truncate in juveniles and emarginate in adults. Body and most of head are covered with ctenoid scales. Rayed section of dorsal fin and other fins is densely covered with small scales. Color is silvery to tan, with three or four broad dark gray to brown bands on body. Juveniles are dark brown to blackish, with a mottling of white spots. Adults may lack dark bands. This species occurs in coastal waters in the western Atlantic from Massachusetts to southeastern Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is associated with rock or reef habitats. Small juveniles are often observed in shallow water, swimming at an angle and resembling dead leaves and other plant debris. Food consists

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EPHIPPIDAE 737

of planktonic and benthic invertebrates. Maximum known size is 900 mm TL. references: Hildebrand and Schroeder 1928; Böhlke and Chaplin 1968; Randall 1968a, 1996;

Hoese and Moore 1977, 1998; Matsuura 1983h; Robins and Ray 1986; Boschung 1992; Cervigón 1993a,b; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Burgess 2002c.

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738

LUVARIDAE

Louvars

This family consists of a single species, and it occurs in the Gulf of Mexico.

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LUVARIDAE 739

Luvarus imperialis Louvar

Rafinesque, 1810

Luvarus imperialis is oval in profile and strongly compressed, with a slender keeled caudal peduncle and a lunate caudal fin. Head is moderately elevated, and snout is blunt. Forehead is compressed and keel-like. Nostril is single and porelike. Eye is rather large and laterally located at about midheight of head. A deep groove is located above eye. Mouth is terminal and very small. Jaw teeth are very slender and few in number. Upper jaw is not protrusible, and maxilla is sheathed by lachrymal bone. Gill opening is relatively short, extending from upper margin of pectoral fin base to beneath eye, where gill membrane is joined to isthmus. Preopercular and opercular margins are membranous and lack spines. Gill rakers on first arch number 10, and each possesses one or more conical teeth. Pectoral fin inserts just below midflank and has 18 rays. Dorsal fin originates just posterior to eye in juveniles and has two spines and 20 to 22 rays, and anteriormost elements are lost with growth. In adults, dorsal fin originates posterior to midpoint of body. Pelvic fin is located beneath pectoral fin base, is reduced to a flap, and covers

anus. Anal fin originates just behind pectoral fin base and has 22 rays. Anus is located in front of pectoral fin base and is partially covered by flap of pelvic fin. Body is covered with nonimbricating scale plates. Scale plates consist of basal plate bearing a single stalk that is expanded into distal stellate plate oriented horizontal to body surface. Scale plates along bases of dorsal and anal fins bear small forwardly directed thorns and large posteriorly directed thorns. Vertebrae number 22, with last two fused. Color is silvery, with a bluish tinge dorsally and a row of pink spots below dorsal fin. This species occurs worldwide in tropical to temperate seas. It is found in oceanic waters of the Gulf of Mexico between 200 and 600 m. Fecundity ranges up to 47.5 million eggs in large specimens. Maximum known size is 180 cm TL.

references: Topp and Girardin 1971; Heemstra 1986n; Robins and Ray 1986; Tyler et al. 1989; Nelson 1994.

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740

ACANTHURIDAE

Surgeonfishes

Surgeonfishes are deep bodied and very compressed, with small eyes, a small terminal mouth, and one or more spines or keeled bony plates on lateral aspect of caudal peduncle. Dorsal profile of head is strongly elevated, and snout is acute and projecting to blunt and straight to slightly concave. Nostril is paired, and nares are close set and slitlike. Eye is located near dorsal margin of head. Upper jaw is not protrusible. Jaw teeth are usually incisor-like, with serrated surfaces, and are arranged in a single row. Teeth are absent in vomer and palatine. Gill opening extends from behind eye to level of mouth. Gill membranes are broadly joined to each other and to isthmus. Gill rakers on first arch are arranged in inner and outer rows. Pectoral fin is wedge shaped and has a moderately oblique base and 14 to 18 rays. Dorsal fin is continuous, lacks notch, and consists of four to nine spines and 22 to 33 rays. Pelvic fin inserts below pectoral fin base and generally has one spine and 5 rays, although it has one spine and 3 rays in some Indo-Pacific species. Pelvic fin originates slightly behind pectoral fin base and has one spine and 3 to 5 rays. Anal fin has two or three spines and 18 to 31 rays. Caudal fin ranges from truncate to lunate, with 16 principal rays. Body and much of head are covered with small ctenoid scales. Caudal peduncle is very slender and possesses three ridged bony plates or one or more scalpel-like spines that are extended forward, away from body, when body musculature is flexed. Gas bladder is present. Vertebrae number 22. Maximum known size is 75 cm TL. Surgeonfishes occur worldwide in tropical to subtropical seas. They are usually associated with coral and rocky reefs to a depth of about 100 m and feed on filamentous algae, zooplankton, and occasionally detritus. Feeding takes place during daylight hours, and individuals often form large schools that may consist of more than one species. Bony plates and spines on caudal peduncle are defenses against predators. There are 72 species in six genera, and 3 species in a single genus occur in the western Central Atlantic, including the Gulf of Mexico. references: Nelson 1994; Krupp 1995; Randall 1996, 2001c; Smith 1997; Smith-Vaniz et al. 1999.

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ACANTHURIDAE 741

Key to the Species of the Gulf of Mexico (Adapted from Randall 1956; Briggs and Caldwell 1957) 1a. Anal fin rays 24 to 26; dorsal fin rays 26 to 28 (usually 27); depth about 58% of SL; caudal spine not outlined with dark pigment; caudal spine sheath pale . . . . . . . . . . . . . . . . . . . . . . . Acanthurus coeruleus p. 744 1b. Anal fin rays 21 to 24; dorsal fin rays 22 to 26 (usually 21 to 25); depth about 50% of SL; caudal spine outlined with dark pigment; caudal spine sheath dusky . . . . . . . . . . . . . 2 2a. Body with 10 to 12 dark brown bars but without narrow stripes; caudal fin without distinct white posterior margin . . . . . . . . . . . . . . . . . . . . . . . Acanthurus chirurgus p. 743 2b. Body without dark brown bars but with narrow stripes; caudal fin with distinct white posterior margin . . . . . . . . . . . . . . . . . . . . . . . . Acanthurus bahianus p. 742

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742 PERCIFORMES

Acanthurus bahianus Castelnau, 1855 Ocean surgeon

Acanthurus bahianus has a relatively long pectoral fin, a lunate caudal fin, and blade of caudal peduncle spine outlined with blue pigment. Head is elevated and straight to level of eye, and moderately convex from eye to dorsal fin origin. Anterior and posterior nares are both oval shaped, with a narrow flap on posterior margins. Teeth number 12 to 14 in upper jaw and 14 to 16 in lower jaw. First gill arch has 18 to 24 (usually 20 to 24) gill rakers. Measurements are expressed as percent of SL: head length 29%–32%, snout length 18%–21%, eye diameter 8%–10%, pectoral fin length 28%–30%, body depth 45%–53%. Pectoral fin has 15 to 17 rays. Dorsal fin has nine spines and 23 to 26 rays. Anal fin has three spines and 21 to 23 rays. Caudal fin is emarginate in small juveniles and lunate or deeply lunate in large juveniles and adults, respectively. Color in life is yellowish brown, with faint greenish gray stripes. Caudal fin is olive yellow, with posterior margin bluish white. Dorsal and anal fin margins are bright blue. In preservative, color

is tannish brown, with opercular margin, dorsal and anal fin margins, and area around caudal peduncle spine dark brown to black and posterior margin of caudal fin white. This species occurs in the tropical and warm temperate Atlantic Ocean. In the western Atlantic it occurs from Massachusetts and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It occurs throughout the Gulf of Mexico on coral reef habitats. Acanthurus randalli is a junior synonym of this species (W. SmithVaniz, pers. comm., June 2001). Maximum known size is 360 mm TL.

references: Longley and Hildebrand 1941; Randall 1956, 1996, 2002c; Briggs and Caldwell 1957 (as A. randalli, in part); Böhlke and Chaplin 1968; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Boschung 1992 (as A. randalli); Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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ACANTHURIDAE 743

Acanthurus chirurgus (Bloch, 1787) Doctorfish

Acanthurus chirurgus has a relatively long pectoral fin, an emarginate caudal fin, and 10 to 12 dark bars on body. Head is elevated and more or less evenly convex to dorsal fin origin. Anterior and posterior nares are oval shaped, with a narrow flap on posterior margins. Teeth number 10 to 18 in upper jaw and 14 to 16 in lower jaw, and number increases with growth. First gill arch has 16 to 19 gill rakers. Measurements are expressed as percent of SL: head length 29%–32%, snout length 19%–20%, eye diameter 8%–9%, pectoral fin length 29%–30%, body depth 52%–54%. Pectoral fin has 16 or 17 rays. Dorsal fin has nine spines and 24 or 25 rays. Anal fin has three spines and 22 or 23 rays. Caudal fin is nearly truncate in small juveniles and emarginate or lunate in large juveniles and adults, respectively. Color in life is brown or tan, with 10 to 12 narrow bars on flanks. Caudal peduncle is light, and

caudal peduncle spine is outlined with blue pigment. Posterior margin of caudal fin has a very narrow pale margin. In preservative, color is tannish brown, with opercular margin and area around caudal peduncle spine dark brown to black and caudal peduncle light. This species occurs in the tropical to warm temperate Atlantic. In the western Atlantic it occurs from Massachusetts and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with coral reefs. Maximum known size is 350 mm TL.

references: Longley and Hildebrand 1941 (as Acanthurus hepatus); Randall 1956, 1996, 2002c; Böhlke and Chaplin 1968; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Matsuura 1983o; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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744 PERCIFORMES

Acanthurus coeruleus Blue tang

Bloch and Schneider, 1801

Acanthurus coeruleus has a relatively long pectoral fin, an emarginate caudal fin, and narrow alternating dark and light stripes on body and fins. Head is steeply elevated, slightly concave to level of eyes, and straight to slightly convex from eye to origin of dorsal fin. Anterior and posterior nares are oval shaped, with a narrow flap on posterior margins. Teeth number 8 to 14 in upper jaw and 10 to 16 in lower jaw, and number increases with growth. First gill arch has 13 or 14 gill rakers. Measurements are expressed as percent of SL: head length 31%– 34%, snout length 22%–26%, eye diameter 8%–12%, pectoral fin length 29%–33%, body depth 60%– 65%. Pectoral fin has 16 or 17 rays. Dorsal fin has nine spines and 26 to 28 rays. Anal fin has three spines and 24 to 26 rays. Caudal fin is emarginate in small juveniles and falcate in large juveniles and adults. Color in life is bright blue, with narrow, dark horizontal lines on body. Caudal peduncle spine is outlined with narrow yellow line. Dor-

sal and anal fins are bright blue, with horizontal orangish brown lines. Juveniles are yellow except for blue margins of dorsal and anal fins and blue margin of eye. In preservative, color is grayish brown, with alternating slightly wavy light and dark horizontal lines on body and fins. This species occurs in the western and Central Atlantic Ocean. In the western Atlantic it occurs from New York and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with coral reefs. Maximum known size is 360 mm TL.

references: Longley and Hildebrand 1941 (as Acanthurus caeruleus); Randall 1956, 1996, 2002c; Böhlke and Chaplin 1968; Bright and Cashman 1974; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Smith 1997; Smith-Vaniz et al. 1999.

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745

SCOMBROLABRACIDAE

This family consists of a single species that occurs worldwide, including the Gulf of Mexico.

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746 PERCIFORMES

Scombrolabrax heterolepis

Roule, 1921

Scombrolabrax heterolepis is moderately elongate, elliptical in profile, and compressed, with upper jaw moderately protrusible and first dorsal fin longer than second dorsal fin. Dorsal profile of head is slightly convex. Nostril is paired, and nares consist of slitlike pores. Interorbital region is flat. Eye is very large, nearly equal to snout length. Mouth is terminal and slightly oblique, with lower jaw projecting slightly beyond upper jaw and upper jaw extending to pupil. Teeth in upper jaw consist of a row of small to moderate-sized, compressed canines and one or two very large, stout canines on either side of symphysis. Teeth in lower jaw consist of a row of moderate to moderately large, compressed canines that increase in size toward corner of mouth and a patch of small teeth at symphysis. Vomer has a V-shaped patch of small teeth, and palatine has a long row of small teeth. Ventral margin of preoperculum is serrated to angle. Operculum has 2 flat spines on posterior margin. Gill membranes are free of isthmus, and a small slit is present behind fourth gill arch. Pseudobranch on inner surface of operculum is well developed. First gill arch has four or five long, slender, and denticulate gill rakers on lower limb. Measurements are expressed as percent of SL: head length 34%–35%, snout length 9%–11%, eye diameter 9%–10%, upper jaw length 16%–17%, pre-dorsal-fin length 40%– 43%, pectoral fin length 30%–36%, pelvic fin length 14%–15%,

body depth 24%–25%. Pectoral fin is long and falcate and has 18 or 19 rays. First dorsal fin is nearly continuous with second and has 12 rather stout spines. Second dorsal fin consists of 1 spine and 14 or 15 rays. Pelvic fin inserts below pectoral fin base and has 1 spine and 5 rays. Anal fin is similar in size and shape to second dorsal fin and has 2 spines and 16 to 18 rays. Caudal fin is forked. Body is covered with small cycloid scales that are variable in size and shape and are deciduous. Lateral line runs very close to dorsal profile and ends slightly anterior to insertion of second dorsal fin. Lateral line scales are larger than other scales, are not deciduous, and number 46 to 50. Vertebrae number 30: 13 precaudal and 17 caudal. Gas bladder is present and in adults possesses bubblelike evaginations that fit into bullae of vertebral parapophyses. Color is dark brown to grayish brown. Lining of mouth is black. This species occurs in the Atlantic, Pacific, and Indian Oceans in tropical and subtropical seas between 100 and 900 m. It has not been captured in the eastern Pacific or the southeastern Atlantic Oceans. In the western Atlantic it occurs between 560 and 1,340 m in the northern and southern Gulf of Mexico to Suriname. Maximum known size is 300 mm TL. references: Potthoff et al. 1980; Fujii 1983d; Johnson 1986; Boschung 1992; Nelson 1994; Nakamura and Parin 2001a.

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747

SPHYRAENIDAE

Barracudas

Barracudas are elongate and subcylindrical to slightly compressed in cross section or moderately compressed, with a large mouth bearing fanglike teeth, two short and widely separated dorsal fins, and a falcate caudal fin. Head is long and pointed. Nostril is paired. Eye is large and located on dorsolateral aspect of head. Mouth is horizontal. Jaws are elongate, upper jaw is not protrusible, and lower jaw projects beyond upper jaw. Jaw teeth are large, sharp, and flattened or conical and of variable size. One or two large canines are usually located at tip of lower jaw. Conical teeth are also located on palatine. Preopercular and opercular margins are entire and lack spines. Gill openings are very broad, and gill membranes are separate and free of isthmus. Gill rakers, if present, are short spinules or platelets. Pseudobranch is well developed on inner surface of operculum. Branchiostegal rays number seven. Pectoral fin is small and inserts low on flank. First dorsal fin originates above or behind pelvic fin base and has five stout spines. Second dorsal fin originates above anal fin and has one spine and nine rays. Pelvic fin inserts on abdomen and has one spine and five rays. Anal fin has one or two spines and seven to nine rays. Caudal fin is deeply forked. Body and part of head are covered with scales. Scales are cycloid in all but three species, in which they are ctenoid. Lateral line is complete and straight and consists of pored scales. Vertebrae number 24: 12 precaudal and 12 caudal. Maximum known size is 205 cm TL. Barracudas occur worldwide in tropical to warm temperate seas. They are pelagic in neritic waters and are generally associated with the surface in the vicinity of reefs and sea-grass beds, although they occur to 100 m. Young individuals of large-sized species and all sizes of small-sized species form schools. There are 20 species in a single genus, and 3 or 4 species occur in the Gulf of Mexico. references: Nelson 1994; Smith-Vaniz et al. 1999; Senou 2001; Russell 2002.

Key to the Species of the Gulf of Mexico (Adapted from Böhlke and Chaplin 1968, in part) 1a. Pectoral fin, when appressed, extending beyond origin of pelvic fin; first dorsal fin originating posterior to origin of pelvic fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Pectoral fin, when appressed, not extending to origin of pelvic fin; first dorsal fin originating above or slightly anterior to origin of pelvic fin . . . . . . . . . . . . . . . . . . . . . . . . . 3

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748 PERCIFORMES

2a. Lateral line scales 75 to 87; inky blotches scattered on side . . . . . . . . . . . . . . . . . . . . . . . . Sphyraena barracuda p. 749 2b. Lateral line scales 108 to 114; no inky blotches on side . . . . . . . . . . . . . . . . . . . . . . Sphyraena guachancho p. 751 3a. Tip of snout slightly pointed . . . Sphyraena borealis p. 750 3b. Tip of snout very pointed . . . . . Sphyraena picudilla p. 752

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SPHYRAENIDAE 749

Sphyraena barracuda (Edwards, 1771) Great barracuda

Sphyraena barracuda has a relatively large mouth and dark bars and inky blotches on side. Snout is long and rounded and lacks fleshy knob. Interorbital region is flat to slightly concave. Mouth is horizontal, and upper jaw extends to anterior margin of eye in juveniles and beyond anterior margin of eye in adults. Upper jaw has several large, compressed canines on side of symphysis and a band of small, compressed, slightly posteriorly directed teeth along margin of jaw. Lower jaw has several large, compressed canines on side of symphysis and a band of large, compressed, vertically directed canines along margin, with teeth increasing in size posteriorly. Palatine has a row of large, compressed canine teeth. First gill arch lacks gill rakers but has platelets that lack distinct spines. Measurements are expressed as percent of SL: head length 31%–35%, snout length 14%–15%, eye diameter 5%– 6%, upper jaw length 14%–15%, pectoral fin length 9%–11%, body depth 13%–16%. Pectoral fin has 11 or 12 rays and, when appressed, extends to or beyond pelvic fin base. First dorsal fin originates slightly posterior to pelvic fin base. Second dorsal fin originates anterior to anal fin origin and has one spine and 8 to 10 rays. Anal fin has two spines and 7 or 8 rays. Caudal fin has 17 principal rays and 8 secondary (procurrent) dorsal rays and 7 procurrent ventral rays. In specimens larger than 50 cm TL, posterior

margin of caudal fin has a pair of large lobes arranged above and below horizontal axis. Small cycloid scales cover side of head and body. Lateral line scales number 75 to 87. Color is gray, with a greenish cast dorsally, and whitish ventrally, with several to many inky blotches on lower side. This species occurs in tropical to warm temperate seas in the Atlantic and Indo-West Pacific over continental and insular shelves. In the western Atlantic it occurs from Massachusetts and Bermuda to southern Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles, within its depth range. Juveniles occur is shallow waters over sandy substrates and vegetated areas. Adults occur in offshore waters and over reefs. Spawning occurs between April and October. Eggs and larvae are pelagic. Maturity is reached between two and three years of life. Food consists of cephalopods, shrimps, and a large variety of ray-finned fishes. Maximum known size is 165 cm SL.

references: de Sylva 1963b, 1984; Böhlke and Chaplin 1968; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Randall 1996; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Ditty et al. 1999; Smith-Vaniz et al. 1999; Senou 2001; Russell 2002.

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750 PERCIFORMES

Sphyraena borealis Northern sennet

DeKay, 1842

Sphyraena borealis has a fleshy tip on lower jaw, a moderately large mouth, and short pectoral fins. Snout is long and slightly pointed. Interorbital region is flat to nearly flat. Maxilla falls short of anterior margin of eye. Upper jaw has several large, moderately compressed canines on side of symphysis and a band of small, compressed, more or less vertically oriented teeth along margin of jaw. Lower jaw has several large, moderately compressed canines on side of symphysis and a band of large, compressed, vertically oriented round canines along margin, with teeth increasing in size posteriorly. Palatine has a row of large, slightly compressed canine teeth. Gill rakers are absent on first arch. Measurements are expressed as percent of SL: head length 29%–31%, snout length 13%–14%, eye diameter 4%–5%, upper jaw length 11%–12%, pectoral fin length 8%–10%, body depth 11%–12%. Pectoral fin has 11 or 12 rays and, when appressed, fails to reach pelvic fin base. First dorsal fin originates above or slightly anterior to pelvic fin base. Second dorsal fin originates anterior to anal fin

origin and has one spine and 8 or 9 rays. Anal fin has one or two spines and 8 or 9 rays. Caudal fin has 17 principal rays and 9 procurrent dorsal and 9 procurrent ventral rays. Small cycloid scales cover sides of head and body. Lateral line scales number 115 to 135. Color is olive brown dorsally and silvery ventrally. Second dorsal fin and anal, pelvic, and caudal fins are often yellowish. Juveniles have dusky blotches along upper flank and along lateral line. This species occurs in warm temperate seas in the western Atlantic over continental and insular shelves from southern Nova Scotia and Bermuda to Panama, including the entire Gulf of Mexico. Spawning occurs on the continental shelf in spring. Eggs and larvae are pelagic. Maximum known size is 460 mm TL.

references: Bigelow and Schroeder 1953; Hoese and Moore 1977, 1998; Robins and Ray 1986; Scott and Scott 1988; Ditty et al. 1999; SmithVaniz et al. 1999; Russell 2002.

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SPHYRAENIDAE 751

Sphyraena guachancho Cuvier, 1829 Guaguanche

Sphyraena guachancho has backwardly directed lower jaw teeth and lacks inky blotches on side. Snout is long and moderately pointed and lacks fleshy knob. Interorbital region is convex. Maxilla extends to margin of eye in adults. Upper jaw has several large, compressed canines on side of symphysis and a band of small, compressed, slightly posteriorly directed teeth along margin of jaw. Lower jaw has several large, compressed canines on side of symphysis and a band of large, posteriorly directed canines along margin, with teeth increasing in size posteriorly. Palatine has a row of large, compressed canine teeth. Gill rakers on first arch consist of small spines. Measurements are expressed as percent of SL: head length 29%– 34%, snout length 12%–15%, eye diameter 5%– 6%, upper jaw length 11%–16%, pectoral fin length 9%–12%, body depth 10%– 14%. Pectoral fin has 11 or 12 rays and, when appressed, extends to or beyond pelvic fin base. First dorsal fin originates above to slightly posterior to pelvic fin base. Second dorsal fin originates anterior to anal fin origin and has one spine and 9 or 10 rays. Anal fin has one or two

spines and 7 to 9 (usually 7 or 8) rays. Last rays of second dorsal and anal fins are slightly extended as filaments. Caudal fin has 17 principal rays and 10 procurrent dorsal and 9 procurrent ventral rays. Small cycloid scales cover sides of head and body. Lateral line scales number 108 to 114. This species occurs in tropical to warm temperate seas in the Atlantic over continental and insular shelves. In the western Atlantic it occurs from Massachusetts to Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is most common in turbid coastal waters over muddy bottoms, including bays and estuaries, and is absent from coral reefs. Spawning occurs in summer on the continental shelf. Food consists of shrimps and small ray-finned fishes. Maximum known size is 60 cm SL. references: Böhlke and Chaplin 1968; Matsuura 1983k; de Sylva 1984; Randall 1996; Schaldach et al. 1997; Smith 1997; Ditty et al. 1999; CastroAguirre et al. 1999; Russell 2002.

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752 PERCIFORMES

Sphyraena picudilla (Poey, 1860) Southern sennet

Sphyraena picudilla has a fleshy tip on lower jaw, a moderately large mouth, and short pectoral fins. Snout is long and very pointed. Interorbital region is slightly convex. Maxilla falls short of anterior margin of eye. Upper jaw has several large, moderately compressed canines on side of symphysis and a band of small, compressed, more or less vertically oriented teeth along margin of jaw. Lower jaw has several large, moderately compressed canines on side of symphysis and a band of large, compressed, vertically oriented conical canines along margin, with teeth increasing in size posteriorly. Palatine has a row of large, moderately compressed canine teeth. Gill rakers on first arch consist of small teeth. Measurements are expressed as percent of SL: head length 34%– 36%, body depth 10%–12%; and as percent of head length: snout length 45%– 48%, eye diameter about 16%, upper jaw length 40%– 42%. Pectoral fin has 11 or 12 rays and, when appressed, fails to reach pelvic fin base. First dorsal fin originates above or slightly anterior to pelvic fin base. Second dorsal fin originates anterior to anal fin origin and has one spine and 8 or 9 rays. Anal fin has two spines and

8 or 9 rays. Caudal fin has 17 principal rays and 18 dorsal and ventral procurrent rays. Small cycloid scales cover side of head and body. Lateral line scales number 113 to 133. Color is dark olive dorsally and silvery on side and ventrally. Side is marked with two reddish stripes in life. This species occurs in the tropical to subtropical western Atlantic from northern Florida and Bermuda to southern Brazil, including the eastern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is absent from the western Gulf of Mexico and continental Caribbean coast except for Venezuela. This species forms large schools and is most often found over mud bottoms. Food consists of squids, shrimps, and small ray-finned fishes. Morphologically and meristically, this species is very similar to S. borealis and may be a junior synonym of this species, as suggested by Smith-Vaniz et al. 1999. Maximum known size is 400 mm SL. references: Böhlke and Chaplin 1968; Matsuura 1983k; de Sylva 1984; Boschung 1992; Randall 1996; Smith 1997; Ditty et al. 1999.

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753

GEMPYLIDAE

Snake mackerels

Gempylids are moderately elongate, moderately deep bodied, and compressed to very elongate, slender, and compressed, with relatively large mouths, relatively large jaw teeth, and a welldeveloped caudal fin. Snout is long and acute to very acute. Nostril is paired, and nares are oval to slitlike. Eye is large and laterally located. Mouth is terminal, large, and horizontal to slightly oblique, with lower jaw projecting beyond upper jaw. Upper jaw is not protrusible, and maxilla is usually free of suborbital bones. Jaw teeth in front of upper jaw, and often in lower jaw, are fanglike. Vomer and palatine often bear small teeth. Gill opening is very extensive, and gill membranes are separate and free of isthmus. Preopercular and opercular margins often bear spines. Gill rakers on first arch are spinous tubercles or are lacking. Pectoral fin is moderately developed, has an oblique to nearly vertical base, and inserts on lower half of flank. Dorsal fins are separated and run most of length of body, with first dorsal fin consisting of spines and longer than second, which consists of rays. Pelvic fin is small, rudimentary, or absent. Anal fin is similar to second dorsal fin and possesses two or three spines and numerous rays. Several taxa have separate finlets behind second dorsal and anal fins. Caudal fin is forked. Body is covered with small to minute scales or modified scales or is naked. Lateral line is single, double, or finely branched. Caudal keels on caudal peduncle are present in only one genus. Vertebrae range from 35 to 60. Maximum known size is 300 cm SL. Snake mackerels occur worldwide in oceanic waters from 200 to 1,000 m. Species are pelagic or benthopelagic. Some species migrate to the surface at night. All species are fastswimming carnivores that feed mainly on ray-finned fishes. There are 23 species in 16 genera and 9 species in 9 genera in the western Central Atlantic and the Gulf of Mexico. references: Nakamura 1986a; Nakamura and Parin 1993, 2001b; Nelson 1994; Smith-Vaniz et al. 1999.

Key to the Species of the Gulf of Mexico (Adapted from Nakamura and Parin 1993) 1a. More than 60 dorsal fin elements; distance from anus to anal fin origin nearly equal to or greater than snout length . . . . . . . . . . . . . . . . . . . . Diplospinus multistriatus p. 755 1b. Fewer than 56 dorsal fin elements; distance from anus to anal fin origin about equal to eye diameter. . . . . . . . . . . . 2 2a. Caudal peduncle with lateral keel and small supplemental keels above and below keel; dorsal fin spines 8 or 9; lateral line single and sinuous . . . . . . . . . . . . . . . . . Lepidocybium flavobrunneum p. 758

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754 PERCIFORMES

2b. Caudal peduncle without keels; more than 12 dorsal fin spines; lateral line single or bifurcated but not sinuous. . . 3 3a. Body with rough, medium-sized scales, interspersed with spinous bony tubercles; midventral keel on belly; lateral line single and obscure . . . . . . . . Ruvettus pretiosus p. 763 3b. Body smooth, scales small, not interspersed with spinous bony tubercles; no midventral keel on belly; lateral line single or double and distinct. . . . . . . . . . . . . . . . . . . . . . . 4 4a. Pelvic fin rudimentary, with 1 spine and 0 to 4 rays, or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4b. Pelvic fin well developed, with 1 spine and 5 rays. . . . . . . 7 5a. Lateral line single . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5b. Lateral line double . . . . . . . . . . . Gempylus serpens p. 757 6a. Two free anal fin spines behind anus, first large and daggerlike; lateral line rather straight; dorsal fin spines 20 or 21 . . . . . . . . . . . . . . . . . . . . . . . . . . Nealotus tripes p. 759 6b. No free anal fin spines behind anus; lateral line curved abruptly downward over pectoral fin; dorsal fin spines 17 to 19 . . . . . . . . . . . . . . Promethichthys prometheus p. 762 7a. Lateral line double . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7b. Lateral line single . . . . . . . . . . . Nesiarchus nasutus p. 761 8a. Lower lateral line branching off under fifth to sixth dorsal fin spines; 2 small spines on lower angle of preoperculum . . . . . . . . . . . . . . . . . . . . . . . . Epinnula magistralis p. 756 8b. Both lateral lines originating at upper end of operculum; no spines on preoperculum . . . . . . . . . . . . . . . . . . . . . . Neoepinnula americana p. 760

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GEMPYLIDAE 755

Diplospinus multistriatus

Maul, 1948

Diplospinus multistriatus is very elongate and compressed, with anus about midway between tip of snout and tip of caudal fin and anterior section of anal fin very low. Head is gently elevated from snout to dorsal fin origin. Anterior naris is located closer to eye than to tip of snout, and posterior naris is elliptical and located just anterior to eye. Lower jaw projects beyond upper jaw and lacks dermal process. Tip of upper jaw possesses a conical dermal process. Upper jaw has three immovable and three or four movable fanglike teeth on side of symphysis, followed by a row of small canine teeth. Lower jaw has fixed fanglike tooth on side of symphysis, followed by a row of caninelike teeth. Vomer lacks teeth. Numerous short, spinelike gill rakers are located on first gill arch. Measurements are expressed as percent of SL: head length about 17%, body depth 6%–7%. Pectoral fin is less than onethird of head length and has 11 to 13 rays. First dorsal fin originates over operculum and has 30 to 36 spines. Second dorsal fin is located immediately behind first, is about half

the length of first, and has 35 to 44 rays. Pelvic fin consists of a minute spine in adults and is located under pectoral fin base. Anal fin originates under origin of second dorsal fin and has 2 spines and 28 to 35 rays, with anterior rays free of membranes. Lateral line is single and runs close to ventral profile. Vertebrae number 57 to 62 in the North and Central Atlantic populations. Color is silvery, with dotted lines along body and jet black gill membranes. This species occurs worldwide in tropical to temperate oceanic seas. In the western Atlantic it occurs from New England to southern Brazil, including the Gulf of Mexico. It is known from the eastern and northern Gulf of Mexico. This species is mesopelagic to 1,000 m but migrates to 100 or 200 m at night. Food consists of crustaceans and small ray-finned fishes. Fecundity is about 1,200 eggs. Females mature at about 160 mm SL, and maximum known size is 330 mm SL. references: Nakamura and Parin 1993, 2001b; Parin et al. 2002a.

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756 PERCIFORMES

Epinnula magistralis Sackfish

Poey, 1854

Epinnula magistralis is moderately elongate, fairly deep bodied, and compressed, with a double lateral line and 2 sharp spines on corner of preoperculum. Head is slightly elevated to anterior naris and moderately elevated from anterior naris to dorsal fin origin. Anterior naris is located about one-third the distance between tip of snout and anterior margin of eye, and posterior naris is elliptical and located midway between anterior naris and anterior margin of eye. Lower jaw projects beyond upper jaw, and both jaws lack dermal processes on tips. Upper jaw has several fixed and flexible fangs near symphysis, followed by a row of widely spaced small canine teeth. Lower jaw has a single fang on side of symphysis, followed by a row of widely spaced caninelike teeth. Vomer lacks teeth, and palatine has a row of small conical teeth. Operculum has 2 flat spines along posterior margin. Measurements are expressed as percent of SL: head length 24%– 33%, body depth 18%–24%. Pectoral fin has 15 rays. First dorsal fin originates above margin of preoperculum and has 15 or 16 strong spines. Second dorsal fin is located immediately

behind first, is less than half the length of first, and has 13 to 17 rays. Pelvic fin is longer than pectoral fin, consists of 1 spine and 5 rays, and inserts posterior to pectoral fin base. Anal fin is similar to second dorsal fin in shape and position and has 2 free spines, followed by 1 spine and 13 to 17 rays connected by membranes. Lateral line is double, with lower branch splitting off upper branch under fifth or sixth dorsal fin spine and running near ventral contour. Color is grayish blue, with head slightly darker than body. Fin membranes of dorsal and pelvic fins are black, basal section of caudal fin is dark blue, and remainder of fin is black. This species occurs in the western Atlantic, western North Pacific, and eastern Indian Oceans. In the western Atlantic it occurs in the northern Gulf of Mexico, off Bermuda, and in the Caribbean Sea. Maximum known size is 1,000 mm SL.

references: Grey 1953; Robins and Ray 1986 (as Epinnula orientalis); Nakamura and Parin 1993; Smith-Vaniz et al. 1999; Parin et al. 2002a.

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GEMPYLIDAE 757

Gempylus serpens Snake mackerel

Cuvier, 1829

Gempylus serpens is very elongate, slender, and compressed, with terminal processes on both jaws, series of finlets behind second dorsal and anal fins, and pelvic fin greatly reduced. Head is gently elevated from snout to dorsal fin origin. Anterior naris is closer to anterior margin of eye than to tip of snout, and posterior naris is elliptical and closer to anterior margin of eye than to anterior naris. Lower jaw projects beyond upper jaw. Maxilla is covered when mouth is closed. Upper jaw has three immovable and zero to three movable fanglike teeth on side of symphysis, followed by a row of small, compressed canine teeth. Lower jaw lacks fangs on side of symphysis but has a row of small, compressed, caninelike teeth. Vomer lacks teeth, and palatine has a row of small teeth. Numerous short, spinelike gill rakers are located on first gill arch. Measurements are expressed as percent of SL: head length 17%– 18%, body depth 6%–7%. Pectoral fin is less than one-third of head length and has 12 to 15 rays. First dorsal fin originates over upper corner of operculum and has 26 to 32 spines. Second dorsal fin is located immediately behind first, consists of 1 minute spine and 11 to 14 rays, and is followed by five or six finlets. Pelvic fin consists of 1 minute spine and 3 or 4 rays and is located under pectoral fin base.

Anal fin originates under origin of second dorsal fin, has 2 free spines followed by 1 spine and 10 to 12 rays connected by membranes, and is followed by six or seven finlets. Lateral line is double, with both branches originating below origin of first dorsal fin, upper branch running the length of first dorsal fin, and lower branch running along midflank to caudal peduncle. Vertebrae number 48 to 55: 24 to 29 precaudal and 23 to 26 caudal. Color is uniformly dark brown, including fins. This species occurs worldwide in tropical to temperate seas from the surface to 200 m. In the western Atlantic it occurs from New York and Bermuda to northern South America, including the Gulf of Mexico. Adults migrate to the surface at night, and larvae and juveniles remain near the surface only during the day. Food consists of squids, pelagic crustaceans, and ray-finned fishes, such as myctophids, exocoetids, scomberesocids, and scombrids. Fecundity ranges from 300,000 to 1,000,000 eggs. Males mature at 430 mm SL, females mature at 500 mm SL, and maximum known size is 100 cm SL. references: Grey 1953; Fujii 1983d; Robins and Ray 1986; Nakamura and Parin 1993, 2001b; Smith-Vaniz et al. 1999; Parin et al. 2002a.

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758 PERCIFORMES

Lepidocybium flavobrunneum

(Smith, 1843)

Lepidocybium flavobrunneum is moderately slender, dorsally and ventrally symmetrical, and slightly compressed, with a very low first dorsal fin, a prominent keel, and two supplementary keels on caudal peduncle. Head is evenly convex from tip of snout to dorsal fin origin. Anterior naris is about midway between tip of snout and anterior margin of eye, and posterior naris is midway between anterior naris and anterior margin of eye. Lower jaw projects slightly beyond upper jaw. Maxilla is exposed when mouth is closed. Upper jaw has two relatively small fangs on side of symphysis, followed by a row of small canine teeth. Lower jaw lacks fangs on side of symphysis but has a row of small caninelike teeth. Vomer and palatine have rows of small teeth. Measurements are expressed as percent of SL: head length 27%–28%, body depth 23%– 24%. Pectoral fin is about one-third of head length and has 15 to 17 rays. First dorsal fin originates anterior to pectoral fin base and has eight or nine short spines. Second dorsal fin is distinctly separated from first dorsal fin, has 16 to 18 rays, and is followed by four to six finlets. Pelvic fin is well developed, consists of one spine and 5 rays, and is located under pectoral fin base. Anal fin originates posterior

to origin of second dorsal fin, has one or two spines and 12 to 14 rays, and is followed by four to six finlets. Caudal fin is falcate. Lateral line is single and strongly sinuous. Body is covered with rather small scales, each of which is surrounded by a network of tubules bearing pores. Vertebrae number 31: 16 precaudal and 15 caudal. Color is uniformly dark brown, including fins. Specimens become increasingly dark with growth. This species occurs worldwide in tropical to warm temperate seas over continental and insular slopes below 200 m, but it is not known from the northern Indian Ocean. In the western Atlantic it occurs from the Scotian Shelf and Georges Bank to Suriname, including Bermuda, the Gulf of Mexico, and the Bahamas. It often migrates to shallower depths at night. Food consists of squids and ray-finned fishes, such as trachipterids, bramids, coryphaenids, and scombrids. Maximum known size is 150 cm SL. references: Schultz and Springer 1956; Bartlett and Backus 1962; Scott and Scott 1988; Boschung 1992; Nakamura and Parin 1993, 2001b; Smith 1997; Smith-Vaniz et al. 1999; Parin et al. 2002a.

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GEMPYLIDAE 759

Nealotus tripes Johnson, 1865 Black snake mackerel

Nealotus tripes is elongate, slender, and compressed, with a minute pelvic fin, 2 free anal fin spines, and a single straight lateral line. Head is moderately elevated from tip of snout to dorsal fin origin. Anterior naris is located about midway between tip of snout and anterior margin of eye, and posterior naris is elliptical and located midway between anterior naris and anterior margin of eye. Lower jaw projects beyond upper jar, and both jaws lack dermal processes on tips. Maxilla is exposed when mouth is closed. Upper jaw has zero to three depressible fangs and three fixed fangs near symphysis, followed by a row of widely spaced small canine teeth. Lower jaw has a single fang on side of symphysis, followed by a row of widely spaced caninelike teeth. Vomer lacks teeth, and palatine has a row of small teeth. Numerous short, spinelike gill rakers are located on first gill arch, with one in corner T-shaped and larger than others. Measurements are expressed as percent of SL: head length 25%– 27%, body depth 11%–14%. Pectoral fin has 13 or 14 rays. First dorsal fin originates above angle of gill opening and has 20 or 21 spines. Second dorsal fin is located close behind first dorsal fin, is less than half the length of first dorsal fin, has 16 to 19 rays, and is followed by

two finlets. Pelvic fin consists of 1 small spine and inserts below pectoral fin base. Anal fin is similar to second dorsal fin in shape and position, has 2 free spines and 15 to 19 rays, and is followed by two finlets. Anterior free anal spine is daggerlike, and posterior free anal spine is parallel to ventral contour. Lateral line slopes posteriorly to run along ventrolateral aspect of caudal peduncle. Body is covered with large deciduous scales. Vertebrae number 36 to 38: 21 or 22 precaudal and 15 or 16 caudal. Color is blackish brown, with dorsal and anal fins pale brown. This species occurs worldwide in tropical to warm temperate seas between 100 and 600 m. In the western Atlantic it occurs from Nova Scotia and the Grand Banks to southern Brazil, including the southern Gulf of Mexico. It migrates to the surface at night. Food consists of squids, crustaceans, and ray-finned fishes such as myctophids. Maturity is reached at about 150 mm SL, and maximum known size is 250 mm SL. references: Grey 1953; Backus et al. 1965; Fujii 1983e; Robins and Ray 1986; Scott and Scott 1988; Nakamura and Parin 1993, 2001b; Parin et al. 2002a.

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760 PERCIFORMES

Neoepinnula americana

(Grey, 1953)

Neoepinnula americana is moderately elongate and compressed, with a double lateral line and dorsal fin origin above angle of opercular opening. Head is moderately sloping from snout to dorsal fin origin. Anterior naris is located about midway between tip of snout and anterior margin of eye, and posterior naris is elliptical and located midway between anterior naris and anterior margin of eye. Lower jaw projects beyond upper jaw, and both jaws lack dermal processes on tips. Maxilla is exposed when mouth is closed. Upper jaw has three fangs near symphysis, followed by a row of widely spaced small canine teeth. Lower jaw has a single fang on side of symphysis, followed by a row of widely spaced caninelike teeth. Vomer and palatine have small, sharp-pointed teeth. Operculum has 2 flat spines along posterior margin. Numerous short, spinelike gill rakers are located on first gill arch. Measurements are expressed as percent of SL: head length 28%– 29%, snout length 10%–11%, eye diameter 5%– 6%, upper jaw length 13%–14%, predorsal-fin length 24%–27%, body depth 22%– 24%. Pectoral fin has 15 or 16 rays. First dorsal fin has 16 spines. Second dorsal fin is lo-

cated immediately behind first, is less than half the length of first dorsal fin, and has 1 spine and 17 to 20 rays. Pelvic fin is smaller than pectoral fin, inserts posterior to pectoral fin base, and consists of 1 spine and 5 rays. Anal fin is similar to second dorsal fin in shape and position and has 2 free spines, followed by 1 spine and 17 to 20 rays connected by membranes. Lateral line is double, and both branches originate above upper angle of gill opening. Color is brown dorsally and silvery on side and ventrally, with mouth cavity pale and branchial cavity pale to dusky. This species occurs between 166 and 457 m on or near the bottom in the western Atlantic from the northern and western Gulf of Mexico and Bermuda to northern Brazil, including the Greater Antilles. Maximum known size is 226 mm SL. references: Mead 1951 (as Epinnula orientalis); Grey 1953, 1959, 1960 (as E. orientalis americana); Fujii 1983e; Boschung 1992 (as E. orientalis); Nakamura and Parin 1993; Smith-Vaniz et al. 1999; Parin et al. 2002a.

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GEMPYLIDAE 761

Nesiarchus nasutus Black gemfish

Johnson, 1862

Nesiarchus nasutus is moderately elongate, slender, and compressed, with well-developed dermal processes at tips of both jaws and a single straight lateral line. Head is slightly elevated from tip of snout to dorsal fin origin. Anterior naris is located closer to anterior margin of eye than to tip of snout, and posterior naris is elliptical and located slightly closer to anterior margin of eye than to anterior naris. Lower jaw distinctly projects beyond upper. Maxilla is exposed when mouth is closed. Upper jaw has zero to three depressible fangs near symphysis, followed by a row of 8 to 12 small canine teeth. Lower jaw has a single fang on side of symphysis, followed by a row of 9 to 14 caninelike teeth. Vomer and palatine lack teeth. Operculum has 2 flat spines along posterior margin. Gill rakers are degenerate on first gill arch. Measurements are expressed as percent of SL: head length 22%–24%, body depth 8%–10%. Pectoral fin is short and has 12 to 14 rays. First dorsal fin originates above angle of gill opening and has 19 to 21 spines. Second dorsal fin is located immediately behind first, is less than half the length of first, has 2 spines

and 19 to 24 rays, and is followed by two finlets. Pelvic fin is small, inserts posterior to pectoral fin base, and consists of 1 spine and 5 rays. Anal fin is smaller than second dorsal fin, has 2 spines and 18 to 21 rays attached by membranes, and is followed by two finlets. Vertebrae number 34 to 36: 20 to 22 precaudal and 14 caudal. Color is dark brown with a violet sheen. Fin membranes and margin of anus are black. This species occurs worldwide, with exception of the eastern Pacific, in tropical to warm temperate seas. In the western Atlantic it occurs from North Carolina and Bermuda to central Brazil, including the Gulf of Mexico and the Caribbean Sea. Adults are benthopelagic between 200 and 1,200 m and migrate upward at night. Larvae and juveniles are epipelagic to mesopelagic. Food consists of squids, crustaceans, and ray-finned fishes. Maximum known size is 130 cm SL. references: Grey 1953; Tucker 1956; Fujii 1983e; Nakamura and Parin 1993, 2001b; Smith-Vaniz et al. 1999; Parin et al. 2002a.

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762 PERCIFORMES

Promethichthys prometheus

(Cuvier, 1832)

Promethichthys prometheus is elongate, moderately slender, and compressed, with rudimentary pelvic fins and finlets behind dorsal and anal fins. Head is slightly elevated from tip of snout to dorsal fin origin. Anterior naris is located slightly closer to tip of snout than to anterior margin of eye, and posterior naris is elliptical and located slightly closer to anterior naris than to anterior margin of eye. Lower jaw distinctly projects beyond upper jaw. Maxilla is exposed when mouth is closed. Upper jaw has three or four immobile and zero to three depressible fangs near symphysis, followed by a row of 12 or 13 small, compressed canine teeth. Lower jaw has a single short fang on side of symphysis, followed by a row of 7 or 8 caninelike teeth. Vomer lacks teeth, and palatine has a single row of small teeth. Operculum has 1 flat spine on posterior margin. Gill rakers on first arch are spinelike and have one to three cusps, with one at angle relatively long and possessing a triple root. Measurements are expressed as percent of SL: head length 27%– 29%, snout length about 11%, eye diameter about 6%, upper jaw length about 13%, body depth 8%–14%. Pectoral fin is about half of head length and has 13 to 15 (usually 13 or 14) rays. First dorsal fin originates above angle of gill opening and has 17 to 19 (usually 17 or 18) spines. Second dorsal fin is located immediately behind first dorsal fin, is less than half the length of first dorsal fin, and has 1 spine

and 17 to 19 rays. Two finlets are located behind second dorsal fin. Pelvic fin is minute, consists of 1 spine, and inserts below pectoral fin base in juvenile specimens but is absent in adults. Anal fin is similar to second dorsal fin in shape and size and has 2 or (rarely) 3 spines and 15 to 17 rays connected by membranes. Two finlets are located behind anal fin. Lateral line is single and bends ventrally to about midflank at about fourth to sixth dorsal fin spine. Vertebrae number 33 to 35: 18 to 20 precaudal and 14 to 16 caudal. Body is entirely scaled in individuals 20 to 25 cm SL and longer. Color is grayish to coppery brown, with fins blackish in large specimens (above 40 cm SL) and yellowish with blackish tips in small specimens. This species occurs worldwide in tropical to warm temperate seas, except for the eastern Pacific. In the western Atlantic it occurs from New England and Bermuda to central Brazil, including the northern and southern Gulf of Mexico and the Caribbean Sea. It is benthopelagic over continental and insular shelves and slopes between 100 and 750 m but migrates into midwater to feed on cephalopods, crustaceans, and ray-finned fishes at night. Maximum known size is 100 cm SL.

references: Grey 1960; Fujii 1983e; Nakamura and Parin 1993, 2001b; Smith-Vaniz et al. 1999; Parin et al. 2002a.

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GEMPYLIDAE 763

Ruvettus pretiosus Oilfish

Cocco, 1833

Ruvettus pretiosus is elongate, elliptical in profile, and moderately compressed, with finlets behind second dorsal and anal fins, a midventral keel on ventral contour, and a rough body surface. Head is slightly convex from snout tip to dorsal fin origin. Anterior naris is located slightly closer to anterior margin of eye than to tip of snout, and posterior naris is elliptical and located slightly closer to anterior margin of eye than to anterior naris. Lower jaw distinctly projects beyond upper jaw. Maxilla is exposed when mouth is closed. Jaws of juveniles have fixed and depressible fanglike teeth anteriorly and a row of small, sharp-pointed teeth posteriorly, but adults possess only a row of small teeth along jaw margins. Vomer and palatine have a single series of small teeth. Operculum has 2 flat spines on posterior margin. Gill rakers on first arch are spinelike and have several cusps, with one in corner larger than others and T-shaped. Measurements are expressed as percent of SL: head length 27%– 33%, snout length 9%–10%, eye diameter about 6%, upper jaw length 15%–17%, body depth 20%–24%. Pectoral fin is less than half of head length and has about 15 rays. First dorsal fin originates above angle of gill opening and has 13 to 15 low spines. Second dorsal fin is close to insertion of first dorsal fin, is slightly

shorter than first, and has 15 to 18 rays. Two finlets are located behind second dorsal fin. Pelvic fin is well developed, inserts below pectoral fin base, and consists of 1 spine and 5 rays. Anal fin is similar to second dorsal fin in shape and size and has 15 to 17 rays. Two finlets are located behind anal fin. Lateral line is single and bends ventrally to about midflank at about fourth to sixth dorsal fin spine. Vertebrae number 32: 16 precaudal and 16 caudal. Body is entirely scaled, and scales are interspersed with spinous bony tubercles. Color is brown to dark brown, with tips of pectoral and pelvic fins black. Margins of dorsal and anal fins are white in juveniles. This species occurs worldwide in tropical to warm temperate seas and is benthopelagic between 200 and 700 m. In the western Atlantic it occurs from Newfoundland and Bermuda to northern Brazil, including the Gulf of Mexico. Food consists of squids, crustaceans, and ray-finned fishes. Maximum known size is 300 cm SL, and size is common to 150 cm SL.

references: Bigelow and Schroeder 1953; Fujii 1983e; Robins and Ray 1986; Scott and Scott 1988; Boschung 1992; Nakamura and Parin 1993, 2001b; Smith-Vaniz et al. 1999; Parin et al. 2002a.

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764

TRICHIURIDAE

Cutlassfishes

Cutlassfishes are very elongate and strongly compressed, with a protruding lower jaw, strong jaw teeth, and a splintered opercular gill cover. Snout is long and acute to very acute. Nostril is single on each side and oval to slitlike. Eye is large and laterally located. Mouth is terminal, large, and horizontal to slightly oblique. Upper jaw is not protrusible, and maxilla is covered by suborbital bone. Teeth in front of upper jaw, and often in lower jaw, are fanglike. Vomer and palatine often bear small teeth. Gill opening is very extensive, and gill membranes are separate and free of isthmus. Preopercular and opercular margins lack spines. Gill rakers on first arch are spinous. Pectoral fin is moderately to poorly developed and inserts on lower half of flank. Dorsal fin is continuous but often shallowly notched, runs most of length of body, and consists of anterior spinous and posterior rayed sections. Rayed section is longer than spinous section. Pelvic fin is either absent or represented by scalelike spines located slightly in front to slightly behind pectoral fin base. Caudal fin is absent, represented by a filament, or reduced to a small forked fin. Pre-anal-fin length is less than half of SL. Body is naked, and lateral line is single and located closer to ventral contour than to dorsal contour. Maximum known size is about 225 cm SL. Cutlassfishes occur worldwide in tropical to warm temperate seas from the surface to 2,000 m. They are either benthic or benthopelagic. Food consists of crustaceans, squids, and ray-finned fishes. Eggs and larvae are pelagic. There are 32 species in nine genera, 8 species in six genera in the western Central Atlantic, and 5 species in separate genera in the Gulf of Mexico. references: Nakamura and Parin 1993, 2001c; Nelson 1994; SmithVaniz et al. 1999; Parin et al 2002b.

Key to the Species of the Gulf of Mexico (Adapted from Nakamura and Parin 1993) 1a. Caudal fin present and small and forked; pelvic fins present but reduced or modified into scalelike process with zero to two rays (may be absent in some species) . . . . . . . . . . . . . 2 1b. Caudal fin absent or tapering into hairlike process; pelvic fins absent or modified into scalelike process . . . . . . . . . . . . . . . . . . . . . . . . . Trichiurus lepturus p. 770 2a. Head profile rising very gradually from tip of snout to dorsal fin origin, not forming sagittal crest; notch present between spinous and rayed sections of dorsal fin . . . . . . . 3 2b. Head profile rising abruptly above eye, forming sagittal crest; no notch between spinous and rayed sections of dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

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TRICHIURIDAE 765

3a. Spinous section of dorsal fin about half the length of rayed section . . . . . . . . . . . . . . . . . . Benthodesmus tenuis p. 767 3b. Spinous section of dorsal fin slightly shorter than rayed section . . . . . . . . . . . . . . . . Aphanopus intermedius p. 766 4a. Head about 15% of TL; eye large and located near dorsal contour . . . . . . . . . . . . . . . . . . . Lepidopus altifrons p. 769 4b. Head about 12.5% of TL; eye moderate in size and located laterally . . . . . . . . . . . . . . Evoxymetopon taeniatus p. 768

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766 PERCIFORMES

Aphanopus intermedius Parin, 1983 Black scabbardfish Aphanopus intermedius is very elongate and depressed, with a small, forked caudal fin and a pelvic fin represented by a single spine in juveniles and absent in adults. Head is very pointed, and profile rises very gradually from snout tip to origin of dorsal fin. Nape is flat and lacks sagittal crest. Jaws have short dermal processes at tips, anterior fangs, and slightly compressed lateral teeth. Teeth are lacking on vomer but occur in a single series on palatine. Measurements are expressed as percent of SL: head length 18%–20%, distance from snout to anus 53%–56%, body depth 6%– 8%; and as percent of head length: snout length 38%– 43%, eye diameter 17%–20%, upper jaw length 45%–50%. Pectoral fin has 12 rays. Dorsal fin consists of 40 to 44 spines, 54 to 59 rays, and 96 to 102 total elements, with a deep notch between the two sections. Pelvic fin inserts below pectoral fin base in juveniles but is absent

in adults. Anal fin originates below second to fifth dorsal fin ray and has 2 spines and 46 to 50 rays. Anal spines are close set and free of remainder of fin and only second is visible in adults. Lateral line is relatively straight. Vertebrae number 102 to 107. Color is black. Mouth and gill cavities are black. This species occurs in the North Atlantic in tropical to warm temperate waters. In the western Atlantic it occurs between 800 and 1,300 m in the northern and southern Gulf of Mexico and off Haiti. It is benthopelagic between 200 and 1,600 m but migrates into the water column at night. Juveniles are mesopelagic. Food consists of cephalopods, crustaceans, and ray-finned fishes. Maximum known size is 100 cm SL. references: Nakamura and Parin 1993; Parin et al. 2002b.

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TRICHIURIDAE 767

Benthodesmus tenuis

(Günther, 1877)

Benthodesmus tenuis is very elongate and depressed, with a small, forked caudal fin and pelvic fins represented by scalelike structures. Head is very pointed, and profile rises very gradually from snout to origin of dorsal fin. Nape is flat and lacks sagittal crest. Mouth is large, lower jaw projects beyond upper, and both jaws have short dermal processes at tips. Jaws have anterior fangs and slightly compressed lateral teeth. Teeth are lacking on vomer, and palatine has a single series of teeth. Measurements are expressed as percent of SL: head length 12%–14%, distance from snout to anus 42%– 46%, body depth 3%– 6%; and as percent of head length: snout length 36%– 44%, eye diameter 13%–17%, upper jaw length 36%– 44%. Pectoral fin has 12 rays. Dorsal fin consists of 40 to 42 spines, 83 to 87 rays, and 125 to 129 total elements, with a

moderate notch between the two sections. Pelvic fin inserts under or slightly anterior to pectoral fin base. Anal fin originates below sixth to eleventh dorsal fin ray and has 2 spines and 72 to 75 rays. Lateral line is relatively straight. Vertebrae number 129 to 131. Color is silvery, with jaws and operculum blackish and inside of mouth and gill cavities black. This species occurs worldwide in tropical and warm temperate seas. In the western Atlantic it occurs from Cape Hatteras to southern Brazil, including the northern and eastern Gulf of Mexico. Adults are benthopelagic between 200 and 850 m, and juveniles are mesopelagic. Maximum known size is 720 mm SL references: Tucker 1953, 1955, 1957; Fujii 1983f; Boschung 1992; Nakamura and Parin 1993; Parin et al. 2002b.

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768 PERCIFORMES

Evoxymetopon taeniatus

Gill, 1863

Evoxymetopon taeniatus is elongate, slender, and compressed, with a triangular pectoral fin, a reduced pelvic fin, and a small caudal fin. Anterior section of head is strongly elevated and convex. Nostril is crescent shaped and located in front of eye. Mouth is moderately large, lower jaw projects slightly beyond upper jaw, and both jaws have short dermal processes at tips. Upper jaw has several fangs anteriorly and slightly compressed lateral teeth. Lower jaw has one fang near symphysis. Teeth are lacking on vomer, and palatine has a single series of fine teeth. Measurements are expressed as percent of SL: head length 12%–13%, body depth 8%–9%. Dorsal fin consists of 81 to 88 elements. Pelvic fin inserts slightly posterior to pectoral fin base and consists of a scalelike spine. Anal fin is very short and posteriorly located. Lateral line is straight and located mid-

laterally or slightly closer to ventral than to dorsal contour. Color is silvery white, with a slight reddish brown cast dorsally and several yellow stripes on body. Anterior section of dorsal fin is blackish. This species occurs in the western Atlantic and the western Pacific in tropical to warm temperate seas. In the western Atlantic it occurs from the northern Gulf of Mexico and Bermuda to southern Brazil, including the Bahamas and the Greater Antilles. It is benthopelagic on the slope and occasionally on the outer continental shelf. Maximum known size is 200 cm SL, and size is commonly between 130 and 180 cm SL. references: Tucker 1956, 1957; Boschung 1992; Nakamura and Parin 1993; Smith-Vaniz et al. 1999; Parin et al. 2002b.

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TRICHIURIDAE 769

Lepidopus altifrons

Parin and Collette, 1993

Lepidopus altifrons is elongate, moderately slender, and compressed, with a prominent sagittal crest, eyes laterally located, and no notch between spinous and rayed sections of dorsal fin. Dorsal profile of head to dorsal fin origin is slightly elevated and nearly straight. Frontal crests on cranium converge anterior to orbits, and interorbital space is strongly convex. Sagittal crest on posterodorsal aspect of cranium extends through nape. Nostril is elliptical and located closer to anterior margin of eye than to tip of snout. Mouth is large, lower jaw projects slightly beyond upper jaw, and both jaws have short dermal processes at tips. Upper jaw has several fangs on side of symphysis and 15 to 20 lateral teeth. Lower jaw has 15 to 21 lateral teeth. Teeth are lacking on vomer, and several small teeth occur in palatine. Gill rakers on first arch number 17 or 18. Measurements are expressed as percent of SL: head length 15%–17%, body depth 8%–9%; and as percent of head length: snout length 37%–

40%, eye length 19%–20%, upper jaw length 33%–36%. Pectoral fin is low on flank and has 12 rays. Dorsal fin consists of 90 to 96 elements. Pelvic fin inserts posterior to pectoral fin base and consists of a scalelike spine and 1 or 2 rays. Anal fin has 2 spines, the second of which is flat, triangular, and short, and 52 to 58 rays, with the last 19 to 23 connected by membrane. Caudal fin is small and forked. Lateral line is straight and slightly declined posteriorly. Color is silvery to brownish and slightly darker along lateral line. Gill cavity is black. This species occurs in the western Atlantic from the Scotian Shelf to southern Brazil, including the northern and western Gulf of Mexico. It is benthopelagic between 200 and 500 m. Maximum known size is 660 mm SL. references: Tucker 1957 (as Evoxymetopon taeniatus); Nakamura and Parin 1993 (as Lepidopus sp.); Parin and Collette 1993; Parin et al. 2002b.

11A-T3324 9/13/05 12:03 PM Page 770

770 PERCIFORMES

Trichiurus lepturus Atlantic cutlassfish

Linnaeus, 1758

Trichiurus lepturus is elongate, slender, and compressed, with posterior margin of opercular margin concave and both pelvic and caudal fins absent. Dorsal profile of head to dorsal fin origin is slightly elevated. Frontal crests on cranium converge posterior to orbits, and interorbital space is flat. Sagittal crest on posterodorsal aspect of cranium is prominent and extends to nape. Nostril is elliptical and located closer to anterior margin of eye than to tip of snout. Mouth is large, lower jaw projects beyond upper jaw, and both jaws have short dermal processes at tips. Upper jaw has two or three fangs on side of symphysis and a series of sharppointed lateral teeth. Lower jaw has one fang on side of symphysis and a series of sharppointed, compressed lateral teeth. Teeth are lacking on vomer, and minute teeth occur in palatine. Gill rakers on first arch are very small. Pectoral fin has one spine and 11 to 13 rays. Dorsal fin originates anterior to opercular opening and consists of three spines and 130 to 135 rays. Anal fin is partially embedded and consists of 100 to 105 rays. Lateral line originates above upper corner of opercular opening, descends to lower third of flank behind pectoral fin base, and continues in a straight line to tip of tail. Color in life is steel blue, with

silvery reflections, pectoral fin nearly transparent, and other fins pale yellow. In preservative, body is silvery gray. This species occurs worldwide in tropical and warm temperate seas. In the western Atlantic it occurs between the shoreline and 350 m, from Cape Cod to northern Argentina, including the Gulf of Mexico, the Bahamas, and the Antilles. It is benthopelagic, and large adults feed at the surface during the day and the bottom at night. Food consists of pelagic squids and ray-finned fishes, such as engraulids, clupeids, myctophids, bregmacerotids, atherinids, carangids, sphyraenids, sciaenids, and scombrids. Juveniles and small adults form loose feeding aggregations and feed on euphausids, other small pelagic crustaceans, and small ray-finned fishes near the surface during the night, and descend to about 100 m above the bottom during the day. Maximum known size is 1,200 mm SL. references: Tucker 1956; Dawson 1967a; Fujii 1983f; Robins and Ray 1986; Boschung 1992; Nakamura and Parin 1993, 2001c; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Parin et al. 2002b.

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771

SCOMBRIDAE

Mackerels and tunas

Mackerels and tunas are elongate, fusiform, and moderately to slightly compressed, with dorsal fins depressible into grooves, finlets behind second dorsal and anal fins, and a narrow caudal peduncle. Snout is moderately long to long and acute. Nostril is paired, and nares are oval to slitlike. Eye is large and laterally located, with anterior and posterior corners covered by adipose eyelid in some genera. Mouth is terminal, large, and slightly oblique. Upper jaw is not protrusible. Premaxilla is beaklike, and maxilla is partially to totally covered by lachrymal bone. Jaw teeth are arranged in a row or in a narrow band and are stout and compressed to slender and conical. Vomer and palatine often bear small teeth, and tongue is toothed in some species. Gill membranes are separate and free of isthmus. Preopercular and opercular margins lack spines. Gill rakers are lathlike and rather stout to slender. Pectoral fin is inserted high on flank. First dorsal fin originates well behind head and is distinctly separated from second dorsal fin. Pelvic fin inserts below pectoral fin base and has one spine and six rays. Anal fin is similar to second dorsal fin in size and shape. Caudal fin is strongly forked to semilunate. Caudal peduncle bears a pair of obliquely oriented keels at base of caudal fin (all species) and a medial keel along midlateral section (some species). Body is naked or partially or totally covered with small or large scales. Lateral line is single and unbranched in most species and runs along upper half of flank. Maximum known size is 300 cm FL. Mackerels and tunas occur worldwide in tropical to warm temperate epipelagic seas. Most species are marine, either coastal or oceanic, but some enter estuaries and at least one migrates up tropical rivers. Food consists of zooplankton, squids, crustaceans, and ray-finned fishes. Eggs and larvae are pelagic. There are 51 species in 15 genera, 16 species in 8 genera in the western Atlantic, and 14 or 15 species in 8 genera in the Gulf of Mexico. references: Collette and Nauen 1983; Collette et al. 1984; Johnson 1986; Collette 2002g; Nelson 1994.

Key to the Species of the Gulf of Mexico (Adapted from Collette and Nauen 1983) 1a. Front and rear of eye covered with adipose eyelid; 5 dorsal and 5 ventral finlets behind second dorsal and anal fins; caudal peduncle with two small oblique keels at base of caudal fin but without long horizontal keel along middle of caudal peduncle . . . . . . . . . . . . . . . . Scomber colias p. 780 1b. Front and rear of eye without adipose eyelid; 7 to 10 finlets behind second dorsal fin and 6 to 10 finlets behind

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772 PERCIFORMES

2a. 2b. 3a.

3b.

4a. 4b. 5a. 5b. 6a. 6b. 7a.

7b.

8a.

8b.

9a.

9b. 10a.

anal fin; caudal peduncle with long horizontal keel along middle, in addition to two small keels at base of caudal fin .............................................2 Jaw teeth well developed, compressed, and triangular; corselet of scales not well developed . . . . . . . . . . . . . . . . . 3 Jaw teeth slender, conical, and little compressed; corselet of scales well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Snout about half of head length; gill rakers on first arch absent; first dorsal fin spines 23 to 27; posterior aspect of maxilla covered by suborbital bone when mouth closed . . . . . . . . . . . . . . . . . . . . . Acanthocybium solandri p. 774 Snout much less than half of head length; gill rakers on first arch 1 to 27; first dorsal fin spines 12 to 22; posterior aspect of maxilla exposed when mouth closed . . . . . . . . . 4 Lateral line descending steeply below first dorsal fin; vertebrae 40 to 46 . . . . . . . . . . . Scomberomorus cavalla p. 781 Lateral line straight or descending gradually below first and second dorsal fins; vertebrae 44 to 56 . . . . . . . . . . . . 5 Flank with spots and at least one stripe; stripe(s) short, wavy, or interrupted . . . . . . Scomberomorus regalis p. 783 Flank usually with spots but without stripes . . . . . . . . . . . . . . . . . . . Scomberomorus maculatus p. 782 Dorsal surface of tongue without cartilaginous ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sarda sarda p. 799 Dorsal surface of tongue with two longitudinal ridges . . . 7 Distance between dorsal fins equal to or greater than length of first dorsal fin base; first dorsal fin with 10 to 12 spines; interpelvic process single and equal to length of longest pelvic fin ray. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Distance between dorsal fins equal to eye diameter; first dorsal fin with 12 to 16 spines; interpelvic process bifid and much shorter than pelvic fin rays. . . . . . . . . . . . . . . . 9 Posterior extension of corselet narrow, only 1 to 5 scales wide under origin of second dorsal fin; dorsal area without scales extending anteriorly to tip of pectoral fin; gill rakers on first arch 38 to 42. . . . . . . . . . . . . Auxis thazard p. 776 Posterior extension of corselet wide, usually 10 to 15 scales wide under origin of second dorsal fin; dorsal area without scales not extending anteriorly to tip of pectoral fin; gill rakers on first arch 39 to 49 . . . . . . . . Auxis rochei p. 775 Belly with three to five dark longitudinal stripes; gill rakers on first arch 53 to 63; vertebrae 41 . . . . . . . . . . . . . . . . . . . . . . . . Katsuwonus pelamis p. 778 Belly without longitudinal stripes; gill rakers on first gill arch 19 to 45; vertebrae 37 to 39 . . . . . . . . . . . . . . . . . . 10 Body naked posterior to corselet of thickened scales; usually several dark spots between pectoral and pelvic fin

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SCOMBRIDAE 773

bases; back dark blue-green with complex striped pattern below dorsal fin bases; pectoral fin rays 25 to 29 . . . . . . . . . . . . . . . . . . . . . . . Euthynnus alletteratus p. 777 10b. Body covered with very small scales posterior to corselet of thickened scales; no black spots on body; back blue and without striped pattern below dorsal fin bases; pectoral fin rays 30 to 36 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11a. Ventral surface of liver with prominent striations; center lobe of liver equal to or longer than left or right lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 11b. Ventral surface of liver without striations; right lobe of liver much longer than left or central lobes . . . . . . . . . . 14 12a. Gill rakers on first arch 31 to 43; pectoral fin less than 80% of head length . . . . . . . . . . . Thunnus thynnus p. 788 12b. Gill rakers on first arch 23 to 31; pectoral fin greater than 80% of head length . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13a. Narrow posterior margin of caudal fin white; pectoral fin extending beyond second dorsal fin base; greatest body depth at or anterior to origin of second dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . Thunnus alalunga p. 784 13b. Posterior margin of caudal fin not white; pectoral fin not extending beyond second dorsal fin base (except in small individuals); greatest body depth near middle of first dorsal fin base. . . . . . . . . . . . . . . . . . . . . . Thunnus obesus p. 787 14a. Gill rakers on first arch 26 to 34 (usually 27 or more); length of second dorsal and anal fins more than 20% of fork length in individuals with 120 cm fork length and larger; maximum size greater than 200 cm fork length . . . . . . . . . . . . . . . . . . . . . . . . . Thunnus albacares p. 785 14b. Gill rakers on first arch 19 to 28 (usually fewer than 27); length of second dorsal and anal fins less than 20% of fork length in all individuals; maximum size less than 110 cm fork length. . . . . . . . . . . . . . . . . Thunnus atlanticus p. 786

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774 PERCIFORMES

Acanthocybium solandri Wahoo

(Cuvier, 1832)

Acanthocybium solandri has a long snout, strong jaw teeth, and a relatively long first dorsal fin. Maxilla extends to level of eye. Jaw teeth are stout, triangular, compressed, finely serrated, and arranged in a single series. Gill rakers are absent. Pectoral fin is less than half of head length and moderately falcate and has 22 to 25 rays. First dorsal fin originates over pectoral fin base and has 23 to 27 spines. Second dorsal fin originates about eye diameter behind insertion of first dorsal fin, has a strongly concave margin and 12 to 16 rays, and is followed by seven to nine finlets. Interpelvic process is bilobed. Anal fin is similar in shape and size to second dorsal fin, has 17 to 19 rays, and is followed by nine finlets. Caudal peduncle is slender and has a well-developed lateral keel between two short diagonal keels. Caudal fin is semilunate and rather short and deep. Body is covered with small scales. Lateral line is single and abruptly curves ventrally under first dorsal fin. Gas bladder is well developed. Vertebrae number 62 to 64: 30 to 32 precaudal and 31 to

33 caudal. Color is iridescent bluish green dorsally and silvery on sides and ventrally, with 24 to 30 cobalt blue vertical bars on side. This species occurs worldwide in tropical and subtropical seas. In the western Atlantic it occurs from New Jersey and Bermuda to northern South America, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is an epipelagic oceanic species occurring solitarily or in loose aggregations. Spawning takes place from May to October, and fecundity ranges up to six million eggs. Food consists of squids and a variety of epipelagic fishes, including clupeids, myctophids, exocoetids, carangids, scombrids, and diodontids. Maximum known size is 210 cm FL. references: Iversen and Yoshida 1957; Hoese and Moore 1977, 1998; Collette and Nauen 1983; Collette 1986b, 2001e, 2002g; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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SCOMBRIDAE 775

Auxis rochei (Risso, 1810) Bullet mackerel Auxis rochei has two longitudinal ridges on tongue, widely separated dorsal fins, and a corselet of scales with a broad posterior extension. Snout is about equal to eye diameter. Mouth is moderate in size and slightly oblique, with maxilla extending below eye and mostly exposed when mouth is closed. Anterior naris is slightly closer to anterior margin of eye than to tip of snout, and posterior naris is slitlike and closer to anterior margin of eye than to anterior naris. Jaw teeth are slender, conical, little compressed, and arranged in a single series. Gill rakers on first arch number 39 to 49. Pectoral fin is short, does not extend beyond dorsal section of corselet, and has 23 to 25 rays. First dorsal fin originates over pectoral fin base and has 10 to 12 spines. Second dorsal fin originates more than one first-dorsal-finbase length behind first dorsal fin, has 10 to 12 rays, and is followed by eight finlets. Interpelvic process is unilobed and longer than pelvic fin. Anal fin is similar in shape and size to second dorsal fin, is located behind base of second dorsal fin, has 11 to 14 rays, and is followed by seven finlets. Caudal peduncle has a well-developed lateral keel between short diagonal keels. Caudal fin is semilunate and rather short and deep. Body is naked except for corselet of scales extending behind head to insertion of dorsal fin, along lateral line to slightly beyond level of second dorsal fin, posterolaterally from gill opening to slightly beyond tip of pec-

toral fin, and posteriorly from isthmus to end of pelvic fin base. Posterior extension of corselet of scales along lateral line is about 10 to 15 scales wide at level of second dorsal fin origin. Lateral line is single and more or less straight. Gas bladder is absent. Vertebrae number 39: 20 precaudal and 19 caudal. Color is bluish to nearly black dorsally and white ventrally. Head is darker than body and 15 or more moderately broad, nearly vertical dark bars are located above corselet extension along lateral line. This species occurs worldwide in tropical and subtropical seas. In the western Atlantic it occurs from Nova Scotia to southeastern Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is epipelagic in neritic waters and forms large schools. Populations in the Atlantic and Indo-West Pacific are considered to be a separate subspecies, A. rochei rochei, from the populations in the eastern Pacific, A. rochei eudorax. In the Gulf of Mexico spawning takes place between March and April and between June and August. Food consists of crustaceans, cephalopods, and epipelagic ray-finned fishes, such as engraulids and clupeids. Maximum known size is 50 cm FL. references: Jones 1963 (as A. thynnoides); Collette 1986b, 2001e, 2002g; Uchida 1981; Collette et al. 1984; Robins and Ray 1986; Scott and Scott 1988; Boschung 1992; Collette and Aadland 1996; Smith 1997; Hoese and Moore 1998.

11A-T3324 9/13/05 12:03 PM Page 776

776 PERCIFORMES

Auxis thazard (Lacepède, 1800) Frigate mackerel

Auxis thazard has two longitudinal ridges on tongue, widely separated dorsal fins, and a corselet of scales with a narrow posterior extension. Snout is about equal to eye diameter. Maxilla extends below eye and is mostly exposed when mouth is closed. Jaw teeth are slender, conical, little compressed, and arranged in a single series. Gill rakers on first arch number 36 to 44 (usually 38 to 42). Pectoral fin is short, reaching slightly beyond dorsal section of corselet, and has 23 to 25 rays. First dorsal fin originates over pectoral fin base and has 10 to 12 spines. Second dorsal fin originates more than one first-dorsal-fin-base length behind first dorsal fin, has 10 to 12 rays, and is followed by eight finlets. Interpelvic process is unilobed and longer than pelvic fin. Anal fin is similar in shape and size to second dorsal fin, is located behind base of second dorsal fin, has 11 to 14 rays, and is followed by seven finlets. Caudal peduncle has a well-developed lateral keel between short diagonal keels. Caudal fin is semilunate and rather short and deep. Body is naked except for corselet of scales extending behind head to insertion of dorsal fin, along lateral line to slightly beyond level of second dorsal fin, posterolaterally from gill opening to slightly beyond tip of pectoral fin, and posteriorly from isthmus to end of pelvic fin base. Posterior extension of corselet of scales along lateral line is usually no more than five scales

wide at level of second dorsal fin origin. Lateral line is single and more or less straight. Gas bladder is absent. Vertebrae number 39: 20 precaudal and 19 caudal. Color is bluish to nearly black dorsally and white ventrally. Head is darker than back, and 15 or more moderately narrow, oblique to nearly horizontal wavy lines are located above corselet extension along lateral line. This species occurs worldwide in tropical and subtropical seas. In the western Atlantic it occurs from southern Florida and Bermuda to southeastern Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is epipelagic in neritic and oceanic waters and forms large schools. Populations in the Atlantic and Indo-West Pacific are considered to be a separate subspecies, A. thazard thazard, from populations in the eastern Pacific, A. thazard brachydorax. Fecundity ranges up to 1.37 million eggs. Food consists of crustaceans, cephalopods, and ray-finned fishes. Maximum known size is 58 cm FL, and maturity is reached at about 29 cm FL. references: Hoese and Moore 1977, 1998; Collette 2001e, 2002g; Uchida 1981; Collette and Nauen 1983; Fujii 1983g; Robins and Ray 1986; Boschung 1992; Collette and Aadland 1996; Smith 1997; Smith-Vaniz et al. 1999.

11A-T3324 9/13/05 12:03 PM Page 777

SCOMBRIDAE 777

Euthynnus alletteratus Little tunny

(Rafinesque, 1810)

Euthynnus alletteratus has two longitudinal ridges on tongue, closely spaced dorsal fins, and a naked body except for corselet of scales. Snout is longer than eye diameter. Maxilla extends below eye. Jaw teeth are small, conical, little compressed, and arranged in a single series. Palatine has teeth, but vomer lacks teeth. Gill rakers on first arch number 37 to 45, and gill teeth (short, flattened structures, medial to gill rakers) number 31 or 32. Pectoral fin is moderately short and has 25 to 29 rays. First dorsal fin originates above pectoral fin base, is elevated anteriorly and very short posteriorly, and has 13 to 17 spines. Second dorsal fin is connected to first dorsal fin at base, has 11 to 13 rays, and is followed by 8 to 10 finlets. Interpelvic process is bilobed and shorter than pelvic fin. Anal fin is similar in shape and size to second dorsal fin, is located behind base of second dorsal fin, has 11 to 15 rays, and is followed by 6 to 8 finlets. Caudal peduncle has a well-developed lateral keel between short diagonal keels. Caudal fin is semilunate and rather short and deep. Corselet extends behind head to insertion of dorsal fin, along lateral line to near insertion of first dorsal fin, posterolater-

ally from gill opening to slightly beyond tip of pectoral fin, and posteriorly from isthmus to end of pelvic fin base. Lateral line is single and slightly wavy. Gas bladder is absent. Vertebrae number 39: 20 precaudal and 19 caudal. Color is dark blue dorsally and white ventrally, with oblique to nearly horizontal dark, wavy stripes or bars above lateral line extension of corselet, and four or five dark spots below pectoral fin. This species occurs in the tropical and warm temperate Atlantic Ocean in neritic and oceanic waters. It is epipelagic in neritic waters and forms schools. In the western Atlantic it occurs from Prince Edward Island and Bermuda to northern Brazil, including the entire Gulf of Mexico. Food consists of crustaceans, squids, tunicates, and small ray-finned fishes, such as clupeids and engraulids. Maximum known size is 100 cm FL.

references: de Sylva and Rathjen 1961; Hoese and Moore 1977, 1998; Collette 1986b, 2002g; Collette and Nauen 1983; Fujii 1983g; Robins and Ray 1986; Scott and Scott 1988; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

11B-T3324 9/13/05 11:19 AM Page 778

778 PERCIFORMES

Katsuwonus pelamis Skipjack tuna

(Linnaeus, 1758)

Katsuwonus pelamis has two horizontal ridges on upper surface of tongue, closely spaced dorsal fins, and several longitudinal stripes on belly. Snout is longer than eye diameter. Maxilla extends below eye. Jaw teeth are small, conical, little compressed, and arranged in a single series. Palatine has teeth, but vomer lacks teeth. Gill rakers on first arch number 53 to 63. Pectoral fin is small and has 26 or 27 rays. First dorsal fin originates slightly behind pectoral fin base, is elevated anteriorly and very short posteriorly, and has 14 to 16 spines. Second dorsal fin is separated from first by distance equal to eye length, has 12 rays, and is followed by seven to nine finlets. Interpelvic process is bilobed and shorter than pelvic fin. Anal fin is similar in shape and size to second dorsal fin, is located under midbase of second dorsal fin, has 1 spine and 12 rays, and is followed by seven or eight finlets. Caudal peduncle has a welldeveloped lateral keel between short diagonal keels. Caudal fin is semilunate and rather short and deep. Body is naked except for corselet of scales extending behind head to insertion of second dorsal fin, along lateral line to near insertion of first dorsal fin, posterolaterally from gill opening to slightly beyond tip of pectoral fin, and posteriorly from isthmus to end of pelvic fin base. Lateral line is single and slightly

wavy. Gas bladder is absent. Vertebrae number 41. Color is purplish blue dorsally and silvery on side and belly, with four to six dark stripes on lower side and belly. In life, wavy stripes may appear as a series of dark blotches. This species occurs in tropical and warm temperate seas worldwide. In the western Atlantic it occurs from southern Nova Scotia and Bermuda to southern Brazil, including the entire Gulf of Mexico and the Caribbean Sea. It is an epipelagic schooling species in oceanic waters and concentrates in regions of upwelling, convergences between oceanic currents, and steep thermal gradients. Temperature range is between 14.7C and 30C. Spawning takes place year-round in tropical waters and during spring and summer in warm temperate waters. Food consists of mollusks, crustaceans, and ray-finned fishes. Most feeding activity takes place in early morning and late afternoon. Maximum known size is 108 cm FL.

references: Hoese and Moore 1977, 1998 (as Euthynnus pelamis); Collette 2001e, 2002g; Collette and Nauen 1983; Matsumoto et al. 1984; Robins and Ray 1986 (as E. pelamis); Scott and Scott 1988; Boschung 1992; Smith 1997; SmithVaniz et al. 1999.

11B-T3324 9/13/05 11:19 AM Page 779

SCOMBRIDAE 779

Sarda sarda (Bloch, 1793) Atlantic bonito

Sarda sarda has close-set dorsal fins and body covered with scales but lacks horizontal ridges on upper surface of tongue. Snout is considerably longer than eye diameter. Maxilla extends below eye. Jaw teeth are relatively large, conical, and arranged in a single series. Palatine has a row of 8 to 21 conical teeth, and vomer occasionally has teeth. Gill rakers on first arch number 16 to 23. Pectoral fin is short and has 22 to 27 rays. First dorsal fin originates over pectoral fin base, is elevated anteriorly and has a straight margin, and has 20 to 23 spines. Second dorsal fin is separated from first by distance equal to eye length, has 13 to 18 rays, and is followed by seven to nine (usually eight) finlets. Bilobed process, shorter than pelvic fin, is located between pelvic fins. Anal fin is similar in shape and size to second dorsal fin, is located near insertion of second dorsal fin, has 14 to 17 rays, and is followed by six or seven finlets. Caudal peduncle has a well-developed lateral keel between short diagonal keels. Caudal fin is semilunate and rather short and deep. Body is completely covered with scales, in addition to corselet of scales that extends behind head to about midlength of first dorsal fin and

posterolaterally from gill opening to tip of pectoral fin. Lateral line is single and slightly wavy. Gas bladder is absent. Vertebrae number 50 to 55: 26 to 28 precaudal and 23 to 27 caudal. Liver has two long lateral lobes and a short middle lobe. Color is blue dorsally and silvery on side and ventrally, with oblique stripes on upper half of body. This species occurs in the tropical and warm temperate Atlantic Ocean. In the western Atlantic it occurs from the Gulf of St. Lawrence and Bermuda to northern Argentina, including the entire Gulf of Mexico, but it is rare in the Caribbean Sea. It is an epipelagic schooling species in coastal waters at temperatures between 12 and 27C. Food consists of crustaceans, squids, and small ray-finned fishes, such as clupeids, gadids, and scombrids. Maximum known size is 914 mm FL in the western Atlantic. references: Collette and Chao 1975; Hoese and Moore 1977, 1998; Collette 1986b, 2002g; Yoshida 1980; Collette and Nauen 1983; Robins and Ray 1986; Scott and Scott 1988; Boschung 1992; Schaldach et al. 1997; Smith-Vaniz et al. 1999.

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780 PERCIFORMES

Scomber colias Chub mackerel

Gmelin, 1789

Scomber colias has adipose eyelids, widely separated dorsal fins, and body covered with scales. Snout is pointed and slightly longer than eye diameter. Maxilla extends below eye. Jaw teeth are small, conical, and arranged in a single series. Palatine has one or two rows of conical teeth, and vomer has one row of teeth. Gill rakers on first arch number 29 to 51. Pectoral fin is short and has 19 or 21 rays. First dorsal fin originates behind pectoral fin base, is triangular, and has 9 or 10 spines. Second dorsal fin has 11 or 12 rays and is followed by five finlets. Interpelvic process is unilobed and small. Anal fin is similar in shape and size to second dorsal fin, originates slightly posterior to second dorsal fin origin, has 1 small separate spine and 12 rays, and is followed by five finlets. Caudal peduncle is very slender and lacks lateral keel between upper and lower oblique keels. Caudal fin is strongly forked. Body is completely covered with scales, with those on head and around pectoral fin larger than remainder, and corselet of scales is absent. Lateral line is single and slightly arched anteriorly. Gas bladder is present. Vertebrae number 31:

14 precaudal and 17 caudal. Color is bluish to greenish dorsally, with oblique zigzag and undulating lines, and silvery white ventrally. Lines break into series of dusky spots near lateral line. This species occurs in the tropical to boreal Atlantic Ocean in coastal waters to 300 m. In the western Atlantic it occurs from the Gulf of St. Lawrence and Bermuda to the northern Gulf of Mexico and Cuba and off Venezuela and southern Brazil. It schools by size, and schools move northward with vernal warming and southward with autumnal cooling. Spawning occurs between 15C and 20C. Food consists of copepods, crustaceans, and ray-finned fishes. Maximum known size is 50 cm FL.

references: Bigelow and Schroeder 1953; Hoese and Moore 1977, 1998; Collette 1986b, 2002g; Collette and Nauen 1983; Fujii 1983g; Collette et al. 1984; Robins and Ray 1986; Scott and Scott 1988; Boschung 1992; Schaldach et al. 1997 (all previous, except first and Collette 2002g, as Scomber japonicus); Smith-Vaniz et al. 1999.

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SCOMBRIDAE 781

Scomberomorus cavalla King mackerel

(Cuvier, 1829)

Scomberomorus cavalla has relatively large jaw teeth, closely spaced dorsal fins, and lateral line strongly curved ventrally. Snout is pointed and considerably longer than eye diameter but shorter than remainder of head. Maxilla extends below posterior margin of eye. Jaw teeth are triangular, compressed, and arranged in a single series. Both palatine and vomer have teeth arranged in patches. Gill rakers on first arch are of moderate size and number 1 to 3 on upper limb and 6 to 10 on lower limb. Pectoral fin is moderately short and has 21 or 23 rays. First dorsal fin originates above pectoral fin base, is slightly elevated anteriorly, and has 12 to 18 spines. Second dorsal fin has 15 to 18 rays and is followed by 7 to 10 finlets. Interpelvic process is bifid and small. Anal fin is similar to second dorsal fin in shape and size, originates slightly posterior to second dorsal fin origin, has 16 to 20 (usually 18 or 19) rays, and is followed by 7 to 10 finlets. Caudal peduncle has a long lateral keel between upper and lower keels. Caudal fin is semilunate. Body is completely covered with small scales, and corselet of scales is absent. Lateral line is single and curves abruptly downward below second dorsal fin. Gas bladder is absent. Vertebrae number 41 to 43: 16 or 17 precaudal and 24 to

26 caudal. Color is dark bluish dorsally and silvery to whitish on sides and ventrally. Juveniles are patterned with small, round dark spots that appear gold or yellow in life. This species occurs in neritic waters in the western Atlantic from Nova Scotia and Bermuda to southern Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. Individuals form large schools that in the Northern Hemisphere migrate northward during vernal warming and southward during autumnal cooling. Resident populations may exist off Louisiana and southern Florida. In the western Gulf of Mexico spawning takes place between May and September. Larvae are found in water between 26.3C and 31.0C. Food consists of penaeid shrimps, squids, and rayfinned fishes, such as clupeids (Opisthonema, Harengula, and Brevoortia). Maximum known size is 173 cm FL.

references: Beaumariage 1973; Hoese and Moore 1977, 1998; Manooch et al. 1978; Collette and Russo 1984; McEachran et al. 1980; Fujii 1983g; Robins et al. 1986; Scott and Scott 1988; Boschung 1992; Schaldach et al. 1997; Smith-Vaniz et al. 1999; Collette 2002g.

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782 PERCIFORMES

Scomberomorus maculatus Spanish mackerel

(Mitchill, 1815)

Scomberomorus maculatus has relatively large jaw teeth, closely spaced dorsal fins, and small spots on side of body. Snout is pointed and considerably longer than eye diameter but shorter than remainder of head. Maxilla extends below posterior margin of eye and is exposed when mouth is closed. Jaw teeth are triangular, compressed, and arranged in a single series. Both palatine and vomer have teeth arranged in patches. Gill rakers on first arch are of moderate size and number 10 to 16, with 1 to 4 on upper limb and 8 to 13 on lower limb. Pectoral fin is moderately short and has 20 to 23 rays. First dorsal fin originates above pectoral fin base, is slightly elevated anteriorly, and has 17 to 19 spines. Second dorsal fin has 17 to 20 (usually 18 or 19) rays and is followed by 7 to 9 finlets. Interpelvic process is bifid and small. Anal fin is similar to second dorsal fin in shape and size, originates slightly posterior to second dorsal fin origin, has 17 to 20 rays, and is followed by 7 to 10 finlets. Caudal peduncle has a long lateral keel between upper and lower

keels. Caudal fin is falciform. Body is completely covered with small scales, and corselet of scales is absent. Lateral line is single and gradually slopes downward toward caudal peduncle. Gas bladder is absent. Vertebrae number 51 to 53: 21 or 22 precaudal and 30 to 31 caudal. Color is silvery dorsally and silvery to whitish on sides and ventrally, with three rows of round to elliptical spots on upper side. This species occurs in neritic waters in the western Atlantic from Nova Scotia and Bermuda to southern Florida and the entire Gulf of Mexico. In the western Gulf of Mexico spawning takes place between May and September. Food consists mostly of small fishes. Maximum known size is 77 cm FL.

references: Hoese and Moore 1977, 1998; Manooch et al. 1978; Collette and Russo 1984; McEachran et al. 1980; Robins and Ray 1986; Scott and Scott 1988; Boschung 1992; Banford et al. 1999; Castro-Aguirre et al. 1999; SmithVaniz et al. 1999; Collette 2002g.

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SCOMBRIDAE 783

Scomberomorus regalis Cero

(Bloch, 1793)

Scomberomorus regalis has relatively large jaw teeth, closely spaced dorsal fins, a midlateral stripe, and several rows of yellowish orange streaks on side of body. Snout is pointed and considerably longer than eye diameter but shorter than remainder of head. Maxilla extends below posterior margin of eye and is exposed when mouth is closed. Jaw teeth are triangular, compressed, and arranged in a single series. Both palatine and vomer have teeth arranged in patches. Gill rakers on first arch are of moderate size and number 12 to 18, with 2 to 4 on upper limb and 10 to 14 on lower limb. Pectoral fin is short and has 20 to 24 (usually 21 or 22) rays. First dorsal fin originates above pectoral fin base, is slightly elevated anteriorly, and has 16 to 18 spines. Second dorsal fin has 16 to 19 rays and is followed by 7 to 9 finlets. Interpelvic process is bifid and small. Anal fin is similar to second dorsal fin in shape and size, originates slightly posterior to second dorsal fin origin, has 15 to 20 (usually 18 or 19) rays, and is followed by 7 to 10 finlets. Caudal peduncle has a long lateral keel between upper and lower keels. Caudal fin is semilunate. Body is completely covered with small scales, and corselet of scales is absent. Lateral line is single and gradually slopes downward toward cau-

dal peduncle. Gas bladder is absent. Vertebrae number 47 or 48: 19 or 20 precaudal and 28 or 29 caudal. Intestine has two folds and three lobes. Color is silvery dorsally and silvery to whitish on sides and ventrally, with midlateral stripe, rows of streaks, and small yellow spots on either side of stripe. Anterior third of first dorsal fin is black, and remainder is white. This species occurs in the western Atlantic from Cape Cod to northern South America, including the northern and southern Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is most common in clear water around reefs. Food consists of squids, shrimps, and small schooling ray-finned fishes, such as clupeids (Harengula, Jenkinsia, and Opisthonema) and atherinids (Atherinella). Fecundity ranges from 160,000 to 2,230,000 in females ranging from 38 to 80 cm FL. Maximum known size is 835 mm FL. Males mature between 325 to 349 mm FL, and females mature at about 380 mm FL. references: Hoese and Moore 1977, 1998; Manooch et al. 1978; Collette and Russo 1984; Collette and Nauen 1983; Schaldach et al. 1997; Banford et al. 1999; Castro-Aguirre et al. 1999; Collette 2002g.

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784 PERCIFORMES

Thunnus alalunga (Bonnaterre, 1788) Albacore Thunnus alalunga has very long pectoral fins, closely spaced dorsal fins, and a poorly developed corselet of scales on anterior section of body. Body profile is elliptical, with maximum girth near origin of second dorsal fin. Snout is acute and conical. Jaw teeth are small, conical, and uniserial. Vomer and palatine have or lack teeth. Gill rakers on first arch number 25 to 31. Pectoral fin is falcate, extends beyond anal fin base, and has 30 to 36 rays. First dorsal fin is high anteriorly, has a concave margin, and consists of 11 or 14 spines. Second dorsal fin is sickle shaped and lower than first dorsal fin, has 12 to 16 rays, and is followed by 7 to 10 finlets. Interpelvic process is small and bifid. Anal fin is similar to second dorsal fin in size and shape, has 11 to 16 rays, and is followed by 7 to 10 finlets. Caudal peduncle bears a long midlateral keel between two shorter oblique keels. Body is covered with small scales, and a corselet of large scales extends over nape, anterior section of lateral line, and thorax region. Vertebrae number 39: 18 precaudal and 21 caudal. Gas bladder is present but is poorly developed and is not evident in juveniles smaller than 50 cm FL. Liver in ventral view is striated due to vascular network. Color is iridescent

blue dorsally and whitish on side and ventrally. Dorsal fin is deep yellow, second dorsal and anal fins are light yellow, and finlets are dark. Posterior margin of caudal fin is white. This species occurs worldwide in tropical to temperate seas. It is epipelagic and mesopelagic in oceanic regions and has been captured at 380 m and may occur to 600 m. In the western Atlantic it occurs from Nova Scotia and Bermuda to Argentina. There are no records from the Gulf of Mexico, but it is possible that it occurs there. This species forms large schools that are wide ranging within water masses, and it tends to concentrate along thermal gradients where pelagic prey is abundant. Individuals school by size and often form mixed schools with Katsuwonus pelamis, Thunnus albacares, and T. thynnus. Food consists of squids, pelagic crustaceans, and ray-finned fishes, such as engraulids, sternoptychids, myctophids, and scomberesocids. Maximum known size is 127 cm FL. Males and females mature at about 90 cm FL. references: Collette 1986b, 2001e, 2002g; Collette and Nauen 1983; Scott and Scott 1988; Smith 1997; Smith-Vaniz et al. 1999.

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SCOMBRIDAE 785

Thunnus albacares Yellowfin tuna

(Bonnaterre, 1788)

Thunnus albacares has narrowly separated dorsal fins, moderately long pectoral fins, and a poorly developed corselet of scales on anterior section of body. Snout is acute and conical. Jaw teeth are small, conical, and uniserial. Gill rakers on first arch number 26 to 34 (usually 27 or more). Pectoral fin is falcate, extends beyond anal fin origin, and has 30 to 36 rays. In juveniles to 50 cm FL, pectoral fin is relatively short. First dorsal fin is high anteriorly and has a concave margin and 13 or 14 spines. Second dorsal fin is falcate, has 12 to 16 rays, and is followed by 7 to 10 finlets. Interpelvic process is small and bifid. Anal fin is similar to second dorsal fin in size and shape, has 11 to 16 rays, and is followed by 7 to 10 finlets. Second dorsal and anal fins are greatly elongated in large adult specimens. Caudal peduncle bears long midlateral keel between two shorter oblique keels. Body is covered with small scales, and a corselet of large scales extends over nape, anterior section of lateral line, and thorax region. Vertebrae number 39. Gas bladder is present. Liver in ventral view lacks striations, and right lobe is largest. Color is metallic dark blue dorsally and whitish on side and ventrally. Belly is

often patterned with about 20 broken, strongly oblique to vertical lines. Dorsal and anal fins are bright yellow, and finlets are bright yellow with black margins. This species occurs worldwide in tropical and subtropical seas. It is epipelagic in oceanic regions with temperatures between 18C and 31C and is found above and below the thermocline. In the western Atlantic it occurs from Nova Scotia and Bermuda to southern Brazil, including the entire Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. This species schools in near-surface waters, generally by size, and may school with other epipelagic scombrids of similar size. Spawning takes place between May and September. Food consists of cephalopods, pelagic crustaceans, and ray-finned fishes. Maximum known size is over 200 cm FL. references: Gibbs and Collette 1967; Hoese and Moore 1977, 1998; Collette 1986b, 2001e, 2002g; Collette and Nauen 1983; Robins and Ray 1986; Scott and Scott 1988; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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786 PERCIFORMES

Thunnus atlanticus Blackfin tuna

(Lesson, 1831)

Thunnus atlanticus has narrowly separated dorsal fins, moderately long pectoral fins, and a poorly developed corselet of scales on anterior section of body. Body profile is subelliptical, with maximum girth near first dorsal fin origin. Snout is acute and conical. Jaw teeth are small, conical, and uniserial. Teeth are present or absent in palatine and vomer. Gill rakers on first arch number 19 to 28 (usually fewer than 27). Pectoral fin is falcate, extends almost to anal fin origin, and has 30 to 36 rays. First dorsal fin is high anteriorly and has a concave margin and 13 to 14 spines. Second dorsal fin is sickle shaped, has 12 to 16 rays, and is followed by 7 to 10 finlets. Interpelvic process is small and bifid. Anal fin is similar to second dorsal fin in size and shape, has 11 to 16 rays, and is followed by 7 to 10 finlets. Caudal peduncle bears long midlateral keel between two shorter oblique keels. Body is covered with small scales, and a corselet of large scales extends over nape, anterior section of lateral line, and thorax region. Vertebrae number 39: 19 precaudal and 20 caudal. Gas bladder is small.

Liver in ventral view lacks striations, and right lobe is largest. Color is metallic dark blue dorsally; silvery gray on sides, often with pale streaks and spots partially vertically oriented; and milky white ventrally. Finlets are dusky with some yellow pigment. This species occurs in the western Atlantic from Martha’s Vineyard and Bermuda to Rio de Janeiro, Brazil, including the entire Gulf of Mexico. It is an epipelagic oceanic species that forms mixed schools with Katsuwonus pelamis of the same size. Food consists of squids, stomatopod and decapod crustacean larvae, amphipods, shrimps, crabs, and ray-finned fishes. Maximum known age is five years. Maximum known size is 100 cm FL.

references: de Sylva 1955; Idyll and de Sylva 1963; Beardsley and Simmons 1971; Hoese and Moore 1977, 1998; Collette and Nauen 1983; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Collette 2002g.

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SCOMBRIDAE 787

Thunnus obesus Bigeye tuna

(Lowe, 1839)

Thunnus obesus has narrowly separated dorsal fins, moderately long pectoral fins, and a poorly developed corselet of scales on anterior section of body. Body profile is subelliptical, with maximum girth near middle of first dorsal fin base. Snout is acute and conical. Jaw teeth are small, conical, and uniserial. Teeth are present or absent in vomer and palatine. Gill rakers on first arch number 23 to 31. Pectoral fin is falcate, extends almost to anal fin origin, and has 30 to 36 rays. First dorsal fin is high anteriorly and has a concave margin and 11 to 14 spines. Second dorsal fin is sickle shaped, has 12 to 16 rays, and is followed by 7 to 10 finlets. Interpelvic process is small and bifid. Anal fin is similar to second dorsal fin in size and shape, has 11 to 16 rays, and is followed by 7 to 10 finlets. Caudal peduncle bears a long midlateral keel between two shorter oblique keels. Body is covered with small scales, and corselet of large scales extends over nape, anterior section of lateral line, and thorax region. Vertebrae number 39: 18 precaudal and 21 caudal. Gas bladder is present. Liver in ventral view has striations resulting from blood vascular network, and right

lobe is not longer than other lobes. Color is metallic dark blue dorsally and white ventrally. Live specimens have iridescent blue stripe on side. First dorsal fin is deep yellow, second dorsal and anal fins are light yellow, and finlets are yellow with black margins. This species occurs worldwide in tropical to subtropical seas at epipelagic and mesopelagic depths, between the surface and 250 m. In the western Atlantic it occurs from southern Nova Scotia and Bermuda to Brazil, including the Gulf of Mexico; however, it is rare in the Straits of Florida and in the Gulf of Mexico. Juveniles and small adults school at the surface and may form schools with T. albacares and Katsuwonus pelamis. Spawning may take place year-round in tropical waters. Food consists of cephalopods, crustaceans, and ray-finned fishes. Maximum known age is over three years, and maximum known size is 239 cm FL. references: Collette 1986b, 2001e, 2002g; Collette and Nauen 1983; Robins and Ray 1986; Scott and Scott 1988; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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788 PERCIFORMES

Thunnus thynnus Bluefin tuna

(Linnaeus, 1758)

Thunnus thynnus has narrowly separated dorsal fins, relatively short pectoral fins, and a poorly developed corselet of scales on anterior section of body. Body profile is subelliptical, with maximum girth near middle of first dorsal fin base. Snout is acute and conical. Jaw teeth are small, conical, and uniserial. Teeth are present or absent in vomer and palatine. Gill rakers on first arch number 31 to 43. Pectoral fin is falcate, extends slightly beyond midbase of first dorsal fin, and has 30 to 36 rays. First dorsal fin is high anteriorly and has a concave margin and 13 or 14 spines. Second dorsal fin is higher than first dorsal fin, is sickle shaped, has 12 to 16 rays, and is followed by 7 to 10 finlets. Interpelvic process is small and bifid. Anal fin is similar to second dorsal fin in size and shape, has 11 to 16 rays, and is followed by 7 to 10 finlets. Caudal peduncle bears a long midlateral keel between two shorter oblique keels. Body is covered with small scales, and a corselet of large scales extends over nape, anterior section of lateral line, and thorax region. Vertebrae number 39: 18 precaudal and 21 caudal. Gas bladder is present. Liver in ventral view has striations result-

ing from blood vascular network, and all three lobes are about equal in length. Color is metallic dark blue dorsally and white ventrally. First dorsal fin is yellow or bluish, second dorsal fin is reddish brown, anal fin and finlets are dusky yellow with black margins. Midlateral keel on caudal peduncle is black in adults. This species occurs at epipelagic and mesopelagic depths in the Atlantic Ocean in tropical to temperate seas. A very similar species, T. orientalis, occurs in the Pacific and Indian Oceans. In the western Atlantic it occurs from Labrador and Newfoundland to Brazil, including the Gulf of Mexico and the Caribbean Sea. Food consists of squids, crabs, and ray-finned fishes, such as engraulids, clupeids, osmerids, paralepids, myctophids, scomberesocids, gadids, merluccids, and scombrids. Maximum known age is about 38 years, and maximum known size is 300 cm FL. references: Gibbs and Collette 1967; Hoese and Moore 1977, 1998; Collette 1986b, 2002g; Collette and Nauen 1983; Robins and Ray 1986; Scott and Scott 1988; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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789

XIPHIIDAE

Billfishes

Billfishes are elongate, moderately compressed, and tapering slightly from origin of first dorsal fin, with premaxilla elongated to form a long, pointed rostrum; mouth inferior; and first dorsal fin originating on back of head. Elongated rostrum is either flattened or round in cross section. Nostril is paired, and anterior naris is oval and posterior naris is slitlike. Eye is either relatively small or large and laterally located. Mouth is large and horizontal. Upper jaw is not protrusible, and maxilla is partially covered by lachrymal bone. Lower jaw is elongated but is less than half of upper jaw length. Jaw teeth in adults are either denticle-like or absent. Vomer and palatine often bear small teeth. Gill membranes are united but free of isthmus. Preopercular and opercular margins lack spines. Gill rakers are absent in adults. Pectoral fin inserts on lower flank and is narrow and acutely pointed. First dorsal fin is either short and high or very long and high, at least anteriorly. Second dorsal fin is short, with last ray short or elongated. Pelvic fin is very long, consists of one spine fused with two rays, and inserts below pectoral fin base, or it is absent. If present, pelvic fin fits into a shallow groove on ventral aspect of body. First anal fin is falcate and originates on posterior half of body. Second anal fin is similar in shape and size to second dorsal fin and is located below or in front of second dorsal fin. Caudal peduncle is slender, depressed, and slightly compressed; has shallow notches dorsally and ventrally; and possesses either a single large lateral keel or two keels located above and below midline. Caudal fin is large and falcate. Body is naked or covered with elongate scales bearing one to several points. Vertebrae number 24 or 26: 11 or 12, or 15 or 16, precaudal and 12 or 13, or 10 or 11, caudal. Gas bladder is present and consists of a single chamber or numerous bubblelike chambers. Billfishes occur worldwide in tropical to warm temperate seas at epipelagic and mesopelagic depths. They are capable of making extensive migrations. Food consists of squids and schooling ray-finned fishes. The elongated bill is apparently used to disable prey. There are 11 species in four genera, and 5 species in four genera occur in the western Central Atlantic, including the Gulf of Mexico. references: Heemstra 1986o,p; Nakamura 1985, 1986b,d, 2002a,b; Nelson 1994.

11B-T3324 9/13/05 11:20 AM Page 790

790 PERCIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Nakamura 1985) 1a. Pelvic fin absent; caudal peduncle with large median keel; rostrum depressed in cross section; first dorsal fin short based and widely separated from second dorsal fin; scales and jaw teeth absent in adults . . . . . Xiphias gladius p. 795 1b. Pelvic fin present; caudal peduncle with two relatively short keels on either side of midline; rostrum rounded in cross section; first dorsal fin long based and close to second dorsal fin; scales and jaw teeth present in adults . . . . . . . . . . 2 2a. First dorsal fin sail-like and distinctly higher than body depth at midlength of body; pelvic fin extends nearly to first anal fin origin and has well-developed membrane . . . . . . . . . . . . . . . . . . . . . . . . Istiophorus albicans p. 791 2b. First dorsal fin not sail-like and slightly higher than or lower than body depth at midlength of body; pelvic fin short, not extending to near anal fin origin, and lacks welldeveloped membrane . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. Height of anterior lobe of first dorsal fin equal to or slightly higher than maximum body depth; nape slightly elevated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3b. Height of anterior lobe of first dorsal fin lower than maximum body depth; nape highly elevated . . . . . . . . . . . . . . . . . . . . . . . . . . Makaira nigricans p. 798 4a. Anterior lobe of first dorsal fin slightly higher than remainder of fin, which tapers slightly over length; anus located anterior to first anal fin origin by distance greater than height of fin; pectoral fin length nearly equal to pelvic fin length . . . . . . . . . . . . . . . . . . . Tetrapturus pfluegeri p. 794 4b. Anterior lobe of first dorsal fin distinctly higher than remainder of fin, which is distinctly tapered over length; anus located anterior to first anal fin origin by distance distinctly shorter than height of fin; pectoral fin length greater than pelvic fin length. . . . . . . . . Tetrapturus albidus p. 793

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XIPHIIDAE 791

Istiophorus albicans Atlantic sailfish

(Latreille, 1804)

Istiophorus albicans has a long sail-like first dorsal fin, and pectoral fin and caudal fin lobes are relatively long in juveniles. Rostrum is very long, slender, and round in cross section. Nape is strongly elevated. Pectoral fin is shorter than pelvic fin and has 17 to 20 rays. First dorsal fin is higher than body depth at midlength, with an undulating margin that is highest at midlength of base, and has 42 to 46 rays, some of which are unsegmented and unbranched, thus resembling fin spines. Second dorsal fin is short, with a concave margin and 6 or 7 rays, and resembles a finlet. Pelvic fin extends nearly to anus and has well-developed membrane. First anal fin is pointed anteriorly and has a concave margin and 11 to 14 rays. Second anal fin originates slightly anterior to second dorsal fin, has 6 or 7 rays, and resembles a finlet. Anus is located close to origin of first anal fin. Scales vary with age and in adults are sparse, are embedded in integument, and have a single spine. Lateral line is single and runs along midflank. Vertebrae number 24: 12 precaudal and 12 caudal. Gas bladder consists of numerous bubblelike chambers. Color is dark blue dorsally, light blue blotched with brown laterally, and silver white ventrally. Side has about 20 narrow bands of light blue round spots. First dorsal fin is dark blue to black, with scattered black dots. This species occurs in the tropical to warm

temperate Atlantic Ocean. In the western Atlantic it occurs from about 40N to 40S, including the oceanic regions of the Gulf of Mexico, Bermuda, and the Caribbean Sea. This is an epipelagic species found above the thermocline between 21C and 28C, but it does descend into colder waters. Occasionally it occurs in coastal waters. During summer months it migrates to higher latitudes. Food consists of mollusks (such as Argonauta spp. and squids), pelagic ray-finned fishes (such as Hemiramphus spp., Strongylura spp., Caranx spp., Trichiurus lepturus, and Euthynnus alletteratus), and benthic ray-finned fishes (such as Lagondon rhomboides and triglids). Maximum known size is 315 cm TL. A very similar species, I. platypterus, occurs in the Pacific and Indian Oceans and is distinguished by having relatively shorter pectoral fins and shorter caudal fin lobes as juveniles (body length less than 90 cm) and by reaching a larger size (340 cm TL).

references: Voss 1953 (as I. americanus); Arnold 1955; de Sylva 1957 (as I. americanus); Wise and Davis 1973; Jolley 1974, 1977; Nakamura 1985, 1986b, 2002b; Robins and Ray 1986; Boschung 1992; Smith 1997 (nine previous references, except Nakamura 2002b, as I. platypterus); Schaldach et al. 1997; Smith-Vaniz et al. 1999.

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792 PERCIFORMES

Makaira nigricans Lacepède, 1802 Blue marlin

Makaira nigricans has a long and relatively low first dorsal fin, a highly elevated nape, and a reticulated lateral line. Rostrum is very long, slender, and round in cross section. Nape is strongly elevated. Palatine has teeth resembling denticles. Pectoral fin is longer than pelvic fin, has 19 to 22 rays, and is depressible to side of body. Anterior lobe of first dorsal fin is lower than body depth at midlength and acutely tipped. First dorsal fin has 41 to 43 rays, some of which are unsegmented and unbranched, thus resembling fin spines. Second dorsal fin is short, has a concave margin and 6 or 7 rays, and resembles a finlet. Pelvic fin has poorly developed membrane. First anal fin is pointed anteriorly and has a concave margin and 13 to 15 rays. Second anal fin originates slightly anterior to second dorsal fin, has 6 or 7 rays, and resembles a finlet. Anus is located close to origin of first anal fin. Caudal peduncle is compressed and has poorly developed notch middorsally and midventrally. Scales are elongate and thick and bear one to three points. Lateral line forms network of loops and polygons, and becomes embedded in skin with growth. Vertebrae number 24: 11 precaudal and 13 caudal. Gas bladder is present and is made up of nu-

merous bubblelike chambers. Color is dark bluish black dorsally and silvery white ventrally. First dorsal fin membrane is black or dark blue, and other fins are brownish black and occasionally tinged with dark blue. This species occurs in the tropical to warm temperate Atlantic Ocean between 40– 45N and 30– 40S. In the western Atlantic it occurs from the southern Nova Scotian banks, the Gulf of Maine, and Bermuda to southern Brazil, including the oceanic waters of the Gulf of Mexico and the Caribbean Sea. It is usually found in or above the thermocline at temperatures ranging from 22C–31C; however, it does feed in colder waters. Food consists of pelagic ray-finned fishes, such as Pseudoscopelus spp., Coryphaena spp., Auxis spp., and other scombrids. Maximum known size is 450 cm TL.

references: Krumholtz and de Sylva 1958; Erdman 1968; Mather et al. 1972; Rivas 1975; Heemstra 1986p; Nakamura 1985, 1986b, 2002b; Robins and Ray 1986; Scott and Scott 1988; Boschung 1992; Schaldach et al. 1997; Smith 1997; SmithVaniz et al. 1999.

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XIPHIIDAE 793

Tetrapturus albidus White marlin

Poey, 1860

Tetrapturus albidus has a long and relatively high first dorsal fin, a moderately elevated nape, and a simple lateral line. Rostrum is very long, slender, and round in cross section. Palatine bone has teeth resembling denticles. Pectoral fin is longer than pelvic fin, has 18 to 21 rays and a rounded tip, and is depressible to side of body. Anterior lobe of first dorsal fin is distinctly higher than body depth at midlength and bluntly tipped. First dorsal fin has 38 to 46 rays, some of which are unsegmented and unbranched, thus resembling fin spines. Second dorsal fin is short, has a concave margin and 5 or 6 rays, and resembles a finlet. Pelvic fin has poorly developed membrane and is slightly shorter than pectoral fin. First anal fin has a broadly rounded margin and 12 to 17 rays. Second anal fin originates slightly anterior to second dorsal fin, has 5 or 6 rays, and resembles a finlet. Anus is located slightly anterior to origin of first anal fin. Caudal peduncle is compressed and has shallow notch mid-dorsally and midventrally. Scales are elongate and thick and bear one or two points. Lateral line is distinct. Vertebrae number 24: 12 precaudal and 12 caudal. Gas bladder is present and is made up of numerous bubblelike chambers. Color is dark bluish black dorsally and silvery white

ventrally, with obscure to distinct vertical lines on flank. First dorsal fin membrane is dark blue with black spots, second dorsal fin is dark blue, and pectoral and caudal fins are blackish brown. This species occurs in the tropical to warm temperate oceanic waters of the Atlantic Ocean. In the western Atlantic it occurs from southern Nova Scotia, the Gulf of Maine, and Bermuda to southern Brazil, including the oceanic waters of the Gulf of Mexico. It is limited to water temperatures over 22C and is found at high latitudes only during warm months of the year. Food consists of squids, crabs, and ray-finned fishes, especially Etrumeus teres, Coryphaena hippurus, and Caranx crysos, but also other clupeids, echeneids, sphyraenids, other carangids, trichiurids, balistids, and tetraodontids. Individuals spawn in pairs, and spawning may occur year-round. Maximum known size is 280 cm TL. references: Gibbs 1957; de Sylva 1963a; Mather et al. 1972, 1975; Nakamura 1985, 1986b, 2002b; Heemstra 1986p; Robins and Ray 1986; Scott and Scott 1988; Boschung 1992; Schaldach et al. 1997; Smith-Vaniz et al. 1999.

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794 PERCIFORMES

Tetrapturus pfluegeri Robins and de Sylva, 1963 Longbill spearfish

Tetrapturus pfluegeri has a long and relatively high first dorsal fin, a nearly straight nape, and a simple lateral line. Rostrum is very long, slender, and round in cross section. Palatine bone has teeth resembling denticles. Pectoral fin is long and broad, has 18 to 21 rays and a rounded tip, and is depressible to side of body. Anterior lobe of first dorsal fin is slightly higher than body depth at midlength and bluntly tipped anteriorly. First dorsal fin has 44 to 50 rays, some of which are unsegmented and unbranched, thus resembling fin spines. Second dorsal fin is short, has a concave margin and 6 or 7 rays, and resembles a finlet. Pelvic fin has poorly developed membrane and is about equal to length of pectoral fin. First anal fin has a broadly rounded margin and 12 to 17 rays. Second anal fin originates distinctly anterior to second dorsal fin, has 6 or 7 rays, and resembles a finlet. Anus is located distinctly anterior to origin of first anal fin. Caudal peduncle is compressed and has a shallow notch mid-dorsally and midventrally. Scales are elongate and thick and bear two to five posterior

points. Lateral line is distinct. Vertebrae number 24: 12 precaudal and 12 caudal. Gas bladder is present and is made up of numerous bubblelike chambers. Color is dark bluish black dorsally and silvery white ventrally. First dorsal fin membrane is dark blue but without black spots, second dorsal fin is dark blue, and pectoral fin is blackish brown. This species occurs in oceanic waters of the tropical to warm temperate Atlantic Ocean. In the western Atlantic it occurs from New Jersey and Bermuda to Venezuela, including the oceanic waters of the Gulf of Mexico, but it is apparently absent in the Antilles. Food consists of squids and pelagic ray-finned fishes, such as engraulids, Alepisaurus spp., Coryphaena spp., and Vomer spp. Maximum known size is 250 cm TL. references: Robins and de Sylva 1963; Ovchinnikov 1970; Robins 1975; Heemstra 1986b; Nakamura 1986b; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith-Vaniz et al. 1999.

11B-T3324 9/13/05 11:20 AM Page 795

XIPHIIDAE 795

Xiphias gladius Linnaeus, 1758 Swordfish

Xiphias gladius has a high and short first dorsal fin and lacks pelvic fins. Body tapers slightly from origin of first dorsal fin. Elongated rostrum is flattened in cross section. Eye is relatively large and laterally located. Maxilla is partially covered by lachrymal bone. Lower jaw is elongated but is less than half of upper jaw length. Jaw teeth are absent in adults. Pectoral fin is falcate, narrow, and acutely pointed and has 16 to 18 rays. First dorsal fin has 34 to 49 rays and originates above or in front of gill opening. Second dorsal fin is very small, has 4 to 6 rays, and is widely separated from first dorsal fin. First anal fin is falcate, originates on posterior half of body, and has 13 or 14 rays. Second anal fin is very small, is usually located in front of second dorsal fin, and has 3 or 4 rays. Caudal peduncle is slender and possesses a large lateral keel and deep notches dorsally and ventrally. Caudal fin is large and semilunate. Anus is located anterior to first anal fin origin. Body is naked in adults, but spinelike scales occur in juveniles to 100 cm TL. Lateral line is present in juveniles to about 100 cm TL but is absent in larger specimens.

Vertebrae number 26: 15 or 16 precaudal and 10 or 11 caudal. Gas bladder consists of a single chamber. This species occurs worldwide in tropical to warm temperate oceanic seas. In the western Atlantic it occurs from Newfoundland, the Grand Banks, and Bermuda to southern Brazil, including the oceanic section of the Gulf of Mexico. It is found from the surface to 630 m in waters above 13C, usually between 18C and 22C. Food consists of pelagic squids and ray-finned fishes, including clupeids, engraulids, Alepisaurus spp., Coryphaena spp., Sphyraena spp., scombrids, Thunnus spp., and Gempylus sp. Adults make major descents into deep water often to feed on benthic fishes such as macrourids, physids, scorpaenids, and trichiurids. Maximum known size is 450 cm TL. references: Arata 1954; Arnold 1955; Eschmeyer 1963; Nakamura 1985, 1986d, 2002a; Heemstra 1986o; Robins and Ray 1986; Scott and Scott 1988; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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796

CENTROLOPHIDAE

Medusafishes

Centrolophids are moderately slender to deep bodied and moderately compressed to compressed, with a single dorsal fin, pelvic fins in adults, and toothed pharyngeal sacs in esophagus. Dorsal profile of head is strongly rounded, with snout moderately blunt to very blunt and covered with pores. Nostril is paired, and nares are close set. Eye is large, is set low on head, and lacks conspicuous adipose tissue. Mouth is terminal, large, and horizontal to moderately oblique, and lower jaw is often included in upper jaw. Maxilla is covered by lachrymal bone when mouth is closed. A single supramaxilla, located on dorsal margin of maxilla, is present in all but one genus. Jaw teeth are small and conical and usually uniserial. Teeth are absent in vomer and palatine. Posterior margin of preoperculum is usually denticulate but is spinulose in juveniles of many species and in adults of one genus. Operculum has two flat spines on posterior margin. Gill rakers on first arch number 17 to 25. Branchiostegal rays number 7. Pectoral fin is well developed, inserts on lower flank, and has 18 to 23 rays. Dorsal fin is continuous and consists of either five weak spines and 31 to 52 rays or five to nine stout, short spines and 19 to 39 rays. Pelvic fin is present in adults, inserts below or behind pectoral fin base, and folds into a broad, shallow groove. Anal fin has zero to three spines and 15 to 35 rays. Caudal fin is shallowly to deeply forked. Scales are cycloid or occasionally ctenoid, are deciduous in all but one genus, and cover body but not head. Head is covered with small pores. Lateral line is single and continuous. Vertebrae number 25 to 60. Anterior section of esophagus is enlarged and specialized to form pharyngeal sac that has irregularly shaped papillae in 10 to 20 longitudinal bands. Maximum known size is 137 cm FL. Centrolophids occur worldwide in tropical to temperate seas, with exception of the tropical eastern Pacific. Most species are epipelagic to demersal over continental shelves, but some are pelagic on the high seas. Juveniles are often associated with floating objects, including jellyfishes, at the surface. Food includes jellyfishes, salps, crustaceans, and small ray-finned fishes. There are 27 species in seven genera, 5 species in two genera in the western Central Atlantic, and 2 or 3 species in one or two genera in the Gulf of Mexico. references: Haedrich 1967, 2002a; Horn 1984; Nelson 1994; SmithVaniz et al. 1999; McDowall 2001.

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CENTROLOPHIDAE 797

Key to the Species of the Gulf of Mexico (Adapted from Haedrich 1967) 1a. Dorsal fin spines fewer than six, weakly developed, and gradually increasing in length over course of fin to length of dorsal fin rays . . . . . . . . . . . . Centrolophus niger p. 798 1b. Dorsal fin spines seven to nine, stout, and distinctly shorter than dorsal fin rays. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Dorsal fin with seven or eight spines and 22 to 25 rays; lateral line scales fewer than 89 . . . . . . . . . . . . . . . . . . . . . . Hyperoglyphe bythites p. 799 2b. Dorsal fin with eight or nine spines and 19 to 21 rays; lateral line scales 89 to 95 . . . . . . . . . . . . . . . . . . . . Hyperoglyphe perciformis p. 800

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798 PERCIFORMES

Centrolophus niger Black ruff

(Gmelin, 1789)

Centrolophus niger is moderately slender and moderately compressed, with small pores on the head and weak dorsal fin spines. Dorsal profile of head is convex, and snout is very blunt. Nares are located near tip of snout, with anterior naris round and posterior naris slitlike. Eye lacks adipose tissue. Mouth is slightly oblique, with lower jaw included in upper jaw and upper jaw extending to below eye. Preoperculum is thin and finely denticulate and slightly bulging posteriorly at corner. Operculum is thin, with a denticulate margin, and has two weak, flat spines. Gill rakers on first arch are stout and moderately long, with 5 or 6 on upper limb and 13 to 16 on lower limb. Pectoral fin is rounded in juveniles and pointed in adults and has 19 to 22 rays. Dorsal fin originates slightly behind pelvic fin insertion and has five spines and 32 to 36 rays. Anal fin has three weak spines and 20 to 23 rays. Caudal fin is broad and moderately forked. Caudal pe-

duncle is deep. Body is covered with small cycloid scales. Skin is thick and contains an extensive subdermal canal system leading to surface through small pores. Lateral line is slightly arched anteriorly. Vertebrae number 25: 10 precaudal and 15 caudal. Color is brown to black, with median fins and pelvic fin darker. This species occurs in the North Atlantic, southern Pacific and eastern Indian Oceans. In the western Atlantic it occurs from Nova Scotia and the Grand Banks to New Jersey. It has been reported from the northern Gulf of Mexico (Hoese and Moore 1977, 1998), but this record has not been confirmed. Juveniles are associated with jellyfishes and molids. Adults are deep dwelling. Maximum known size is 110 cm TL. references: Haedrich 1967, 1986b; Hoese and Moore 1977, 1998; Scott and Scott 1988; Robins and Ray 1986.

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CENTROLOPHIDAE 799

Hyperoglyphe bythites Black driftfish

(Ginsburg, 1954)

Hyperoglyphe bythites is moderately deep bodied and moderately compressed, with short, stout dorsal fin spines; a spinous preopercular margin; and a pelvic fin that inserts under pectoral fin base. Dorsal profile of head is strongly convex, and snout is very blunt. Nares are located near tip of snout, with anterior naris round and posterior naris slitlike. Eye lacks adipose tissue. Mouth is moderately oblique, with lower jaw included in upper jaw and upper jaw extending to below eye. Preoperculum is striated, and margin has many small spines extending from striations. Opercular margin is finely denticulate or entire. Gill rakers on first arch number 6 or 7 on upper limb, 1 in corner, and 15 or 16 on lower limb. Measurements are expressed as percent of SL: head length 31%– 34%, snout length 7%–9%, eye diameter 7%– 8%, upper jaw length 11%–12%, pectoral fin length 23%–27%, pelvic fin length 17%–19%, caudal peduncle depth 10%–12%, body depth 43%– 47%. Pectoral fin has 20 or 21 rays. Dorsal fin originates over or slightly behind pectoral fin base and has seven or eight spines and 22 to 25 rays. Anal fin has three spines and

16 or 17 rays. Body, operculum, and bases of median fins are covered with moderately large cycloid scales. Lateral line is arched anteriorly to midlength of anal fin base. Lateral line scales number about 87. Vertebrae number 25: 10 precaudal and 15 caudal. Color is greenish gray or bluish gray to reddish brown, and darker dorsally than ventrally. Median fins are usually darker than body. This species occurs between 30 and 200 m in the western Atlantic from southern Florida to the Gulf of Mexico, or possibly to Brazil. In the Gulf of Mexico it is known from the west coast of Florida and from Alabama, Louisiana, and Veracruz. Juveniles are found at the surface and are associated with flotsam. Adults are found near the bottom and form large schools. Maximum known size is 600 mm TL. The status of this species is uncertain. references: Ginsburg 1954 (as Palinurichthys bythites); Dawson 1972a; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Haedrich 2002a.

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800 PERCIFORMES

Hyperoglyphe perciformis Barrelfish

(Mitchill, 1818)

Hyperoglyphe perciformis is moderately deep bodied and compressed, with short, stout dorsal fin spines; a spinous preopercular margin; and a pelvic fin that inserts under pectoral fin base. Dorsal profile of head is strongly convex, and snout is very blunt. Nares are located near tip of snout, with anterior naris round and posterior naris slitlike. Eye lacks adipose tissue. Mouth is moderately oblique, with lower jaw included in upper jaw and upper jaw extending below eye. Preoperculum is striated, and margin has many small spines extending from striations. Opercular margin is finely denticulate or entire. Gill rakers on first arch number 6 or 7 on upper limb, 1 in corner, and 15 to 17 on lower limb. Measurements are expressed as percent of SL: head length 30%–34%, snout length 7%– 8%, eye diameter 6%–9%, upper jaw length 11%–12%, pectoral fin length 22%–25%, pelvic fin length 16%–21%, caudal peduncle depth 10%–12%, body depth 35%– 43%. Pectoral fin has 21 to 23 rays. Dorsal fin originates slightly behind pectoral fin base and has eight or nine short spines and

19 to 21 rays. Anal fin has three spines and 16 or 17 rays. Body, operculum, and bases of median fins are covered with moderately large cycloid scales. Lateral line is arched anteriorly to midlength of anal fin base. Lateral line scales number 89 to 95. Vertebrae number 25: 10 precaudal and 15 caudal. Color is greenish gray or bluish gray to reddish brown, often with a mottled pattern, and darker dorsally than ventrally. Median fins are usually darker than body. This species occurs in the North Atlantic. In the western Atlantic it occurs from Nova Scotia and Bermuda to the northern Gulf of Mexico. Adults occur near the bottom between 60 and 121 m. Juveniles are associated with drifting seaweed. Food consists of mollusks, squids, crustaceans, and small ray-finned fishes. Maximum known size is 910 mm TL. The status of this species is uncertain. references: Ginsburg 1954; Haedrich 1967, 1986b, 2002a; Dawson 1972a; Scott and Scott 1988; Boschung 1992; Smith-Vaniz et al. 1999.

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801

NOMEIDAE

Driftfishes

Driftfishes are slender to deep bodied and compressed, with two dorsal fins and pelvic fins present in adults. Dorsal profile of head is moderately to strongly convex, with snout moderately blunt to very blunt. Nostril is paired, and nares are close set. Eye is large and laterally located and has inconspicuous adipose tissue. Mouth is terminal, small, and moderately oblique. Supramaxilla is absent. Upper jaw is not protrusible, and maxilla is partially to totally covered by lachrymal bone. Jaw teeth are small, conical or multicuspid, and usually uniserial. Teeth also occur in vomer, palatine, and basibranchs. Posterior margin of preoperculum is usually entire or fleshy. Operculum is thin and has 2 flat, weak spines on posterior margin. Gill rakers are slender and number 23 to 29. Branchiostegal rays number 6. Pectoral fin is well developed, inserts on lower flank, and has 16 to 23 rays. First dorsal fin consists of 9 to 12 slender to stout spines and folds into a groove. Second dorsal fin consists of 1 or 2 weak spines and 15 to 30 rays. Pelvic fin inserts below or behind pectoral fin base and folds into a narrow groove, with last ray attached to abdomen by membrane. Anal fin is similar in size and shape to second dorsal fin and has 1 to 3 spines and 15 to 30 rays. Caudal fin is deeply forked, and upper and lower lobes overlap in a scissorlike manner when depressed. Scales are deciduous and cycloid or weakly ctenoid, and cover body and often sides of head. Lateral line is single, runs close to dorsal profile, and may not extend to caudal peduncle. Vertebrae number 30 to 42. Anterior section of esophagus is enlarged and specialized to form pharyngeal sac that has papillae with stellate bases arranged in five broad bands. Maximum known size is about 100 cm TL. Nemeids occur worldwide in tropical to subtropical oceanic waters. Adults inhabit midwaters or are demersal over continental slopes. Juveniles are found near the sea surface and are often associated with jellyfishes and siphonophores. There are 15 species in three genera, and 7 species in three genera occur in the western Central Atlantic and the Gulf of Mexico. references: Haedrich 1967, 1986b, 2002b; Horn 1984; Agafonova 1994; Nelson 1994; Smith 1997; Smith-Vaniz et al. 1999; Last 2001b.

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802 PERCIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Haedrich 1967; Haedrich and Horn 1972; Butler 1979; Agafonova 1994) 1a. Maximum body depth less than 35% of SL; dorsal fin origin over or behind pectoral fin base . . . . . . . . . . . . . . . . . 2 1b. Maximum body depth greater than 40% of SL; dorsal fin origin anterior to or over pectoral fin base . . . . . . . . . . . . 5 2a. Anal fin with one to three spines and 14 to 25 rays; pelvic fin base on a vertical with or behind pectoral fin base; tongue with oval patch of knoblike teeth . . . . . . . . . . . . . 3 2b. Anal fin with one or two spines and 24 to 29 rays; pelvic fin base anterior to or under pectoral fin base; tongue without tooth patch. . . . . . . . . . . . . . . Nomeus gronovii p. 806 3a. Vomerine teeth in oval patch . . . . . . . . . . . . . . . . . . . . . . 4 3b. Vomerine teeth in a single row . . . . . . . . . . . . . . . . . . . . . . . . . . Cubiceps capensis p. 803 4a. Anal fin with three spines and 19 to 22 rays; dorsal fin rays 21 to 24; pelvic fin base below or slightly behind pectoral fin base; no bony keel on breast . . . . . . . . . . . . . . . . . . . . . . . . . . . Cubiceps gracilis p. 804 4b. Anal fin with two spines and 14 to 17 rays; dorsal fin rays 15 to 18; pelvic fin base behind pectoral fin base; bony keel on breast . . . . . . . . . . . . . . . Cubiceps pauciradiatus p. 805 5a. Lower jaw teeth long, knifelike, compressed, close set, and different from those of upper jaw . . . . . . . . . . . . . . . . . . . 6 5b. Lower jaw teeth short, not knifelike, round in cross section, spaced, and like those in upper jaw . . . . . . . . . . . . . . . . . . . . . . . . . . Psenes cyanophrys p. 808 6a. Second dorsal fin spines and rays 28 to 34; anal fin spines and rays 29 to 34 . . . . . . . . . . . . . Psenes pellucidus p. 809 6b. Second dorsal fin spines and rays 20 to 23; anal fin spines and rays 23 or 24 . . . . . . . . . . . . Psenes arafurensis p. 807

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NOMEIDAE 803

Cubiceps capensis

(Smith, 1845)

Cubiceps capensis is slender, fusiform, and compressed, with winglike pectoral fins, a relatively long second dorsal fin, and scales on much of head. Dorsal profile of head is moderately convex, and snout is moderately pointed. Nares are porelike and located near tip of snout. Adipose tissue around eye is well developed. Maxilla extends to anterior margin of eye. Jaw teeth are slightly recurved. Vomer has a single row, palatine has one to several rows, and tongue has a single row of small teeth. Gill rakers on first arch are long and slender and number 7 to 11 on upper limb, 1 in corner, and 16 to 20 on lower limb. Measurements are expressed as percent of SL: head length 27%– 29%, pre-dorsal-fin length 32%–38%, preanal-fin length 57%– 62%, pectoral fin length 27%–36%, pelvic fin length about 17%, body depth 26%–28%; and as percent of head length: snout length 27%–30%, orbit length 23%–25%, interorbital length 25%–39%, upper jaw length 30%–35%. Pectoral fin has 20 to 24 rays. First dorsal fin has 11 spines, and second dorsal fin has 1 spine and 20 to 22 rays. Pelvic fin inserts slightly behind pectoral fin base. Anal fin has 3 spines and 19 to 22 rays. Upper procurrent caudal fin rays number 9 to

11, and lower procurrent caudal fin rays number 9 to 11. Body and head, except for tip of snout and preorbital space, are covered with deciduous cycloid scales. Lateral line terminates at end of second dorsal fin base. Lateral line scales number 62 to 71. Vertebrae number 31: 12 precaudal and 19 caudal. Color in preservative is grayish brown, with belly nearly black and fins grayish black. In life, head is dark blue. This species occurs worldwide in subtropical to warm temperate seas in both oceanic and neritic waters. In the western Atlantic it occurs from Brown’s Bank off Nova Scotia and from Bermuda to northern Argentina, including the northern Gulf of Mexico. This species approaches the sea surface at night, but its daytime depth is unknown. Large specimens sometimes wash ashore. Maximum known size is 87 cm SL.

references: Haedrich 1967, 1972, 1986, 2002b; Haedrich and Horn 1972 (all as C. caeruleus); Ahlstrom et al. 1976 (in part as C. caeruleus); Butler 1979; Arndt 1981 (as C. baxteri); Scott and Scott 1988; Agafonova 1994; Smith-Vaniz et al. 1999; Last 2001b.

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804 PERCIFORMES

Cubiceps gracilis (Lowe, 1843)

Cubiceps gracilis is slender, fusiform, and compressed, with winglike pectoral fins, a relatively long second dorsal fin, and scales on much of head. Dorsal profile of head is moderately convex, and snout is moderately blunt. Nares are porelike and located near tip of snout. Adipose tissue around eye is well developed. Maxilla extends to anterior margin of eye. Jaw teeth are slightly recurved. Vomer has an oval patch, palatine has a single series, and tongue has a patch of small, blunt teeth. Gill rakers on first arch are long and slender and number 6 to 10 on upper limb and 16 to 18 on lower limb. Measurements are expressed as percent of SL: head length 28%–32%, pre-dorsal-fin length 36%–39%, pre-anal-fin length 59%– 63%, pectoral fin length 25%–31%, pelvic fin length 8%–11%, body depth 23%–28%; and as percent of head length: snout length 23%–31%, orbit length 25%–30%, interorbital length 20%–34%, upper jaw length 24% to 31%. Pectoral fin has 21 to 24 rays. First dorsal fin has 11 or 12 spines, and second dorsal fin has 1 spine and 21 to 24 rays. Pelvic fin inserts be-

low to slightly behind pectoral fin base. Anal fin has 3 spines and 19 to 22 rays. Upper procurrent caudal fin rays number 7 to 10, and lower procurrent caudal fin rays number 7 to 10. Body and head, including tip of snout, preorbital space, and operculum, are covered with deciduous cycloid scales. Lateral line terminates at end of second dorsal fin base. Lateral line scales number 59 to 69. Color in preservative is grayish brown or steely gray, with bases of dorsal and anal fins darker than those of other fins. This species occurs in the northern subtropical and warm temperate Atlantic Ocean. It is a mesopelagic species that rises toward the surface at night. In the western Atlantic it occurs in deep water on the Grand Banks, off Virginia, and in the northern Gulf of Mexico. Maximum known size is 72 cm SL. references: Haedrich 1967, 1986d, 2002b; Haedrich and Horn 1972; Hoese and Moore 1977; Scott and Scott 1988; Agafonova and Kukuyev 1990; Agafonova 1994.

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NOMEIDAE 805

Cubiceps pauciradiatus Günther, 1872

Cubiceps pauciradiatus is moderately elongate and compressed, with winglike pectoral fins, a bony keel on breast, and scales on much of head. Dorsal profile of head is slightly convex, and snout is moderately blunt. Nares are porelike and located near tip of snout. Adipose tissue around eye is well developed. Maxilla extends to anterior margin of eye. Jaw teeth are slightly recurved. Vomer has a patch of knobby teeth, palatine has a single series of about three teeth, and tongue has a small patch of teeth. Gill rakers on first arch are long and slender and number 7 to 9 on upper limb and 16 to 19 on lower limb. Measurements are expressed as percent of SL: head length 28%–33%, pre-dorsal-fin length 35%– 40%, pre-anal-fin length 66%–73%, pectoral fin length 28%– 37%, pelvic fin length 11%–14%, body depth 21%–28%; and as percent of head length: snout length 25%–34%, orbit length 25%– 32%, interorbital length 26%–34%, upper jaw length 25%–32%. Pectoral fin has 17 to 20 rays. First dorsal fin has 10 or 11 spines, and second dorsal fin has 1 spine and 15 to 18 rays. Pelvic fin inserts behind pectoral fin base. Anal fin has 2 spines and 14 to 17 rays. Upper pro-

current caudal fin rays number 8 to 10, and lower procurrent caudal fin rays number 8 to 10. Scales are cycloid and deciduous and cover body and most of head, including upper head, cheeks, and operculum. Lateral line terminates at end of second dorsal fin base. Lateral line scales number 46 to 60. Vertebrae number 30 or 31: 13 or 14 precaudal and 17 or 18 caudal. Color in preservative is gray to grayish brown, with bases of fins relatively darker than remainder of fins. This species occurs worldwide in tropical and subtropical seas between 40N and 40S. In the western Atlantic it occurs between 384 and 823 m, from the Scotian Shelf to northern Argentina, including the eastern and northern Gulf of Mexico. Maximum known size is 141 mm SL.

references: Haedrich 1965, 1967, 1986d, 2002b; Haedrich and Horn 1972 (all as C. athenae); Butler 1979; Shimizu 1983c; Robins and Ray 1986 (as C. athenae); Scott and Scott 1988; Salekhov 1989; Agafonova and Poluyaktov 1992; Agafonova 1994; Hoese and Moore 1998; Last 2001b.

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806 PERCIFORMES

Nomeus gronovii Man-of-war fish

(Gmelin, 1789)

Nomeus gronovii is elongate, moderately slender, and moderately compressed, with large fanlike pelvic fins and a blotched and spotted color pattern. Dorsal profile of head is evenly convex, and snout is blunt. Nares are at tip of snout, with anterior naris round and posterior naris slitlike. Eye is of moderate size and surrounded by well-developed adipose tissue. Mouth is moderately oblique, with maxilla terminating below anterior margin of eye. Jaw teeth are slightly recurved. Vomer and palatine have a single series of small, recurved teeth. Gill rakers on first gill arch are slender and number 8 or 9 on upper limb, 1 in corner, and 15 to 18 on lower limb. Measurements (of adults) are expressed as percent of SL: head length 29%– 30%, snout length 6%–7%, eye length 8%– 9%, upper jaw length 6%–11%, pectoral fin length 26%–33%, pelvic fin length 22%–25%, body depth 25%–27%. Juveniles have a relatively larger eye, shorter pectoral fin, longer pelvic fin, and deeper body than adults. Pectoral fin is rounded in juveniles and pointed in adults and has 21 to 23 rays. First dorsal fin has 8 to 11 spines, and second dorsal fin has 1 spine and 24 to 28 rays. Pelvic fin inserts anterior to pectoral fin base and is connected to body by membrane along its length. Anal fin is similar to second dorsal fin in shape and size and has 2 spines and 24 to 29 rays. Caudal peduncle is slender and compressed. Caudal fin is

deeply forked, and lobes are long. Body is covered with small cycloid scales, and scales extend forward on nape to anterior margin of eye. Snout and top of head are covered with adipose tissue. Lateral line runs along dorsal contour and terminates at level of end of base of second dorsal fin. Vertebrae number 41: 14 precaudal and 27 caudal. Color of juveniles in life is silvery, with bright blue blotches and spots scattered over body. Pelvic fin rays and membranes are bright blue. In preservative, color is tan, with dark brown blotches and spots. Adults are dark bluish or brown. This species occurs worldwide in tropical and warm temperate seas except for the eastern Atlantic. In the western Atlantic it occurs from Newfoundland, the Grand Banks, and Bermuda to Brazil, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is oceanic, and juveniles are associated with the Portuguese man-of-war (Physalia). Large specimens also occur independently of Physalia in deep water. Maximum known size is 400 mm TL.

references: Haedrich 1967, 1986d, 2002b; Böhlke and Chaplin 1968; Haedrich and Horn 1972; Ahlstrom et al. 1976; Hoese and Moore 1977, 1998; Horn 1984; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Last 2001b.

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NOMEIDAE 807

Psenes arafurensis Silver driftfish

Günther, 1889

Psenes arafurensis is relatively deep bodied and compressed, with knifelike teeth in lower jaw, pelvic fins in adults, and mottled or banded color pattern. Dorsal profile of head is moderately elevated, and snout is truncate. Nares are located on tip of snout, with anterior naris porelike and posterior naris slitlike. Eye is relatively large and surrounded by adipose tissue. Maxilla extends almost to center of eye. Teeth in upper jaw are slender, pointed, widely spaced, and uniserial. Teeth in lower jaw are long, compressed, and close set. Vomer has several series, and palatine has a single series of sharp-pointed teeth. Gill rakers on first arch are slender and number 9 on upper limb, 1 in corner, and 15 on lower limb. Measurements are expressed as percent of SL: head length 33%–35%, snout length 7%– 8%, eye length 9%–12%, upper jaw length 10%–12%, pre-dorsal-fin length 33%–36%, pre-anal-fin length 59%– 62%, pectoral fin length 31%– 32%, pelvic fin length 18%–24%, body depth about 40%. Pectoral fin is rounded and has 20 to 22 rays. First dorsal fin originates anterior to pectoral fin base and has 10 or 11 spines. Second dorsal fin has 1 or 2 spines and 19 to

21 rays. Pelvic fin inserts under posterior section of pectoral fin base. Anal fin is slightly higher than second dorsal fin and has 3 spines and 20 or 21 rays. Caudal peduncle is slender. Caudal fin is deeply forked, and upper and lower lobes are sharply rounded. Body, including bases of unpaired fins, is covered with small scales. Scales are deciduous and possess several weak ctenii. Lateral line follows dorsal contour and terminates at end of second dorsal fin. Vertebrae number 35: 12 precaudal and 23 caudal. Color is tan, with series of discontinuous dark brown bands on sides. Adults are plain colored, but juveniles are banded and have black pelvic fins. This species occurs worldwide in tropical to temperate seas. In the western Atlantic it occurs from the southern Scotian banks to northern South America, including the northern Gulf of Mexico. In the Gulf of Mexico it occurs off Alabama. Individuals are associated with jellyfishes. Maximum known size is 250 mm TL. references: Shimizu 1983c; Robins and Ray 1986; Scott and Scott 1988; Boschung 1992; Smith 1997 (all as P. maculatus); Last 2001b; Haedrich 2002b.

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808 PERCIFORMES

Psenes cyanophrys Freckled driftfish

Valenciennes, 1833

Psenes cyanophrys is relatively deep bodied and compressed, with longitudinal streaks on side. Dorsal profile of head is moderately elevated, and snout is truncate. Nares are located on tip of snout, with anterior naris porelike and posterior naris slitlike. Eye is moderately large and surrounded by adipose tissue. Maxilla extends to below center of eye. Jaw teeth are slender, pointed, and widely spaced. Vomer has a small patch of teeth, and palatine has a single series of teeth. Gill rakers on first arch are slender and number 11 on upper limb and 19 on lower limb. Measurements are expressed as percent of SL: head length 30%– 35%, snout length 7%– 8%, eye length 9%– 11%, upper jaw length 9%–11%, pectoral fin length 27%–31%, pelvic fin length 13%–19%, body depth 44%–51%. Pectoral fin is rounded and has 17 to 20 rays. First dorsal fin originates anterior to pectoral fin base and has 9 to 11 spines. Second dorsal fin has 1 spine and 24 to 28 rays. Pelvic fin inserts under posterior section of pectoral fin base. Anal fin is slightly higher than second dorsal fin and has 3 spines

and 24 to 28 rays. Caudal peduncle is slender. Caudal fin is deeply forked. Body, including bases of unpaired fins, is covered with small scales. Scales on top of head anterior to posterior margin of orbits are relatively small. Scales are deciduous and possess several weak ctenii. Lateral line follows dorsal contour and terminates at end of second dorsal fin. Vertebrae number 31: 12 or 13 precaudal and 18 or 19 caudal. Color is brown to yellowish in preservative, with small spots on scales forming series of stripes on flanks. This species occurs worldwide in tropical and subtropical seas. In the western Atlantic it occurs from Massachusetts and Bermuda to northern South America, including the northern Gulf of Mexico. Maximum known size is 230 mm TL. references: Haedrich 1967, 1986d, 2002b; Haedrich and Horn 1972; Ahlstrom et al. 1976; Hoese and Moore 1977, 1998; Shimizu 1983c; Horn 1984; Robins and Ray 1986; Boschung 1992; Smith-Vaniz et al. 1999; Last 2001b.

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NOMEIDAE 809

Psenes pellucidus Bluefin driftfish

Lütken, 1880

Psenes pellucidus is relatively deep bodied and compressed, with knifelike teeth in lower jaw, bases of median fins translucent, and pelvic fins in adults. Dorsal profile of head is moderately elevated, and snout is truncate. Nares are located on tip of snout, with anterior naris porelike and posterior naris slitlike. Eye is relatively large and surrounded by adipose tissue. Maxilla extends almost to center of eye. Teeth in upper jaw are slender, pointed, and widely spaced. Teeth in lower jaw are long, compressed, and close set. Vomer has a small patch of teeth, and palatine has a single series of teeth. Gill rakers on first arch are slender and number 8 or 9 on upper limb, 1 in corner, and 14 to 16 on lower limb. Measurements are expressed as percent of SL: head length 33%– 34%, snout length 8%–9%, eye length 10%– 13%, upper jaw length 13%–15%, pectoral fin length 27%–28%, pelvic fin length about 19%, body depth 54%–55%. Pectoral fin is rounded and has 18 to 20 rays. First dorsal fin originates anterior to pectoral fin base and has 10 or 11 spines. Second dorsal fin has 1 or 2 spines and 27 to 32 rays. Pelvic fin inserts under posterior

section of pectoral fin base. Anal fin is slightly higher than second dorsal fin and has 3 spines and 26 to 31 rays. Caudal peduncle is slender. Caudal fin is deeply forked. Body, including bases of unpaired fins, is covered with small scales. Scales are cycloid and deciduous and possess several weak ctenii. Lateral line follows dorsal contour and terminates at end of second dorsal fin. Vertebrae number 40 to 42: 13 or 14 precaudal and 27 to 29 caudal. Color is dark brown in preservative. Adults are plain colored, but juveniles are banded. This species occurs worldwide in tropical and subtropical seas, with exception of the eastern Pacific. In the western Atlantic it occurs from Georges Bank and Bermuda, possibly to South America, including the northern Gulf of Mexico. Maximum known size is 80 cm TL. references: Haedrich 1967, 1986d, 2002b; Haedrich and Horn 1972; Hoese and Moore 1977, 1998; Shimizu 1983c; Horn 1984; Robins and Ray 1986; Scott and Scott 1988; Smith 1997; SmithVaniz et al. 1999; Last 2001b.

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810

ARIOMMATIDAE

Ariommatids are either slender and rounded in cross section or deep bodied and compressed, with two dorsal fins, pelvic fins present in adults, and caudal peduncle square in cross section. Dorsal and ventral profiles of body are symmetrical. Profile of head is evenly convex, and snout is moderately to strongly blunt. Nostril is paired, and nares are close set. Eye is moderately large to large, laterally located, and surrounded by adipose tissue. Mouth is terminal, small, and moderately oblique. Maxilla is partially or totally covered by lachrymal bone. Supramaxilla is absent. Jaw teeth are small, simple or tricuspid, and uniserial. Teeth are absent in vomer, palatine, and basibranchs. Posterior margin of preoperculum is usually entire or fleshy. Operculum is thin. Gill rakers on first arch are slender and number 20 to 30. Branchiostegal rays number 6. Pectoral fin is well developed and inserts on lower flank. First dorsal fin consists of 10 to 13 slender spines and folds into a groove. Second dorsal fin consists of 1 spine and 13 to 16 rays. Pelvic fin inserts below pectoral fin base and folds into a groove, with last ray attached to body by membrane. Anal fin is similar in size and shape to second dorsal fin and has 1 to 3 spines and 13 to 18 rays. Caudal fin is deeply forked. Caudal peduncle is very slender, with a weak keel above and below caudal fin base. Scales are cycloid and deciduous. Lateral line is single and high, follows dorsal profile, and does not extend onto caudal peduncle. Branch of lateral line extends over eye in bony canal. Vertebrae number 29 to 32. Anterior section of esophagus is enlarged and specialized to form pharyngeal sac that has papillae in upper end of sac. Papillae have rounded bases and are not arranged in bands. Maximum known size is 20 cm TL. Ariommatids occur worldwide in tropical to subtropical oceanic waters except for the eastern Pacific. They live in deep water near continental and insular shelves and are thought to occur near the bottom to a depth of 750 m. There are six species in a single genus, and three species occur in the western Central Atlantic and the Gulf of Mexico. references: Haedrich 1967, 2002c; Nelson 1994; Smith 1997; Last 2001c.

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ARIOMMATIDAE 811

Key to the Species of the Gulf of Mexico (Adapted from Haedrich and Horn 1972; Haedrich 2002c) 1a. Body depth greater than 33% of SL; body with spots . . . . . . . . . . . . . . . . . . . . . . . . . . . Ariomma regulus p. 814 1b. Body depth less than 28% of SL; body without spots. . . . 2 2a. Body uniformly dark brown to blackish; peritoneum dark, with many melanophores; lateral line scales 50 to 65; scales on top of head extend to anterior margin of eye . . . . . . . . . . . . . . . . . . . . . . . . . Ariomma melanum p. 813 2b. Body pale brown to blue dorsally and silvery ventrally; peritoneum pale, with few melanophores; lateral line scales 30 to 45; scales on top of head extend to pupil of eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ariomma bondi p. 812

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812 PERCIFORMES

Ariomma bondi Silver rag

Fowler, 1930

Ariomma bondi is elongate, slender, and moderately compressed, with relatively large eyes, relatively large scales, and a pale peritoneum. Snout is moderately blunt. Anterior naris is porelike, and posterior naris is slitlike. Jaw teeth are conical and slightly recurved. Gill rakers are slender and number 8 to 10 on upper limb and 15 to 18 on lower limb. Measurements are expressed as percent of SL: head length 27%–31%, snout length 7%– 8%, eye length 9%–10%, upper jaw length 6%–7%, pectoral fin length 21%–23%, pelvic fin length 13%–15%, body depth 24%–27%. Pectoral fin has 20 to 23 rays. First dorsal fin is triangular and has 10 or 11 spines, and second dorsal fin has 1 spine and 14 to 17 rays. Anal fin has 3 spines and 12 to 16 rays. Scales cover

body and extend anteriorly on top of head to level of anterior margin of pupil. Lateral line scales number 30 to 45. Vertebrae number 30 or 31. Color is brown to bluish brown dorsally and silvery or pale ventrally. This species occurs in the tropical and warm temperate Atlantic Ocean. In the western Atlantic it occurs between 25 and 640 m, from Nova Scotia to Uruguay, including the northern and western Gulf of Mexico and the Caribbean Sea. Maximum known size is 300 mm TL. references: Ginsburg 1954 (as Cubiceps nigriargenteus); Haedrich 1967, 2002c; Haedrich and Horn 1972; Hoese and Moore 1977, 1998; Shimizu 1983d; Robins and Ray 1986; Scott and Scott 1988; Boschung 1992.

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ARIOMMATIDAE 813

Ariomma melanum Brown driftfish

(Ginsburg, 1954)

Ariomma melanum is elongate, slender, and moderately compressed, with relatively small eyes, relatively small scales, and a dark peritoneum. Snout is moderately blunt. Anterior naris is porelike, and posterior naris is slitlike. Jaw teeth are conical and slightly recurved. Gill rakers are slender and number 9 to 11 on upper limb and 18 to 20 on lower limb. Measurements are expressed as percent of SL: head length 30%–32%, snout length 8%–10%, eye length 8%–9%, upper jaw length 8%–9%, pectoral fin length 21%–22%, pelvic fin length 11%–15%, body depth 24%–25%. Pectoral fin has 21 to 23 rays. First dorsal fin is triangular and has 10 or 11 spines, and second dorsal fin has 1 spine and 15 to 18 rays. Anal fin

has 3 spines and 13 to 16 rays. Scales cover body and extend onto top of head to level of anterior margin of eye. Lateral line scales number 50 to 65. Vertebrae number 30 or 31. Color is dark brown to blackish. Peritoneum is darkly pigmented. This species occurs in the tropical and warm temperate Atlantic Ocean. In the western Atlantic it occurs from New York to the Caribbean Sea, including the northern Gulf of Mexico. Maximum known size is 300 mm TL.

references: Ginsburg 1954; Haedrich 1967, 1986d, 2002c; Haedrich and Horn 1972; Shimizu 1983d; Robins and Ray 1986; Boschung 1992.

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814 PERCIFORMES

Ariomma regulus Spotted driftfish

(Poey, 1868)

Ariomma regulus is deep bodied and compressed, with relatively small eyes, relatively small scales, and a dark peritoneum. Snout is blunt. Anterior naris is porelike, and posterior naris is slitlike. Jaw teeth are conical and slightly recurved. Gill rakers are slender and number 7 on upper limb and 16 on lower limb. Measurements are expressed as percent of SL: head length 32%–34%, snout length 7%–9%, eye length 9%–10%, upper jaw length 8%– 10%, pectoral fin length 25%–27%, pelvic fin length 16%–18%, body depth 39%– 45%. Pectoral fin has 21 to 23 rays. First dorsal fin is broadly rounded and has 10 or 11 spines, and second dorsal fin has 1 spine and 14 or 15 rays. Anal fin is similar in shape and size to sec-

ond dorsal fin and has 3 spines and 14 or 15 rays. Scales cover body and extend onto top of head to level of orbit. Vertebrae number 30 to 32. Juveniles are light colored, with irregular blotches on side. Adults have dark spots, smaller than eye diameter, on upper half of body. This species occurs in the western Atlantic from North Carolina to Guyana, including the northern Gulf of Mexico and the Caribbean Sea. Maximum known size is 230 mm TL. references: McKinney 1961; Haedrich 1967, 2002c; Haedrich and Horn 1972; Hoese and Moore 1977, 1998; Shimizu 1983d; Robins and Ray 1986; Boschung 1992; Smith 1997.

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815

TETRAGONURIDAE

Tetragonurids are elongate, slender, and almost round in cross section, with first dorsal fin originating behind pectoral fin base, base of first dorsal fin longer than base of second dorsal fin, and two keels on long caudal peduncle. Dorsal profile of head is moderately convex, and snout is blunt. Nostril is paired, and nares are widely separated and located on upper section of snout. Eye is moderately large and laterally located, with series of grooves on posterior rim but without adipose tissue. Mouth is terminal and slightly oblique, with maxilla extending below eye and largely covered by lachrymal bone when mouth is closed. Supramaxilla is absent. Upper jaw is not protrusible. Jaw teeth are small, simple or tricuspid, and uniserial. Upper jaw teeth are small, pointed, recurved, and widely spaced. Lower jaw teeth are large, knifelike, and close set. Head of vomer has well-developed, recurved teeth. Shafts of vomer and palatine have a single series of teeth. Small teeth are present on basibranchs and are scattered over tongue. Posterior margins of preoperculum and operculum are spinulose in juveniles but entire in adults. Operculum has 2 weak spines on posterior margin. Gill rakers are flattened and number 14 to 20. Branchiostegal rays number 5 or 6. Pectoral fin is small and inserts on lower flank. First dorsal fin consists of 10 to 20 stout spines and folds into a groove. Second dorsal fin consists of 0 to 1 spine and 10 to 17 rays. Pelvic fin is present in adults, inserts behind pectoral fin base, and folds into a groove. Anal fin originates slightly posterior to origin of second dorsal fin and has 1 or 2 spines and 9 to 15 rays. Caudal peduncle is long and thick and has two keels composed of modified scales. Caudal fin is forked. Body and cheeks are covered with moderate-sized thick scales possessing longitudinal keels. Small scales are located at bases of median fins. Lateral line is single, runs near dorsal profile, and extends onto caudal peduncle to origin of keels. Vertebrae number 43 to 61. Anterior section of esophagus is enlarged and specialized to form pharyngeal sac. Maximum known size is 50 cm TL. Tetragonurids occur worldwide in tropical and subtropical oceanic waters at epipelagic and mesopelagic depths. Juveniles are associated with tubelike salps. Food consists of ctenophores and jellyfishes. There are three species in a single genus, one or two species occur in the western Central Atlantic, and one species occurs in the Gulf of Mexico. references: Grey 1955a; Haedrich 1967, 1986e, 2002d; Nelson 1994; Last 2001d.

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816 PERCIFORMES

Tetragonurus atlanticus Bigeye squaretail

Lowe, 1839

Tetragonurus atlanticus has a relatively long dorsal fin and a relatively large head. Interorbital width is about equal to eye diameter. Snout is slightly longer than eye diameter. Jaw teeth increase in number with growth. Number of upper jaw teeth ranges from 6 to 14 in specimens from 15 to 25 mm SL, 10 to 17 in specimens from 50 to 100 mm SL, and 27 to 40 in specimens greater than 200 mm SL. Lower jaw teeth range from 10 to 16 in specimens from 15 to 25 mm SL to 50 to 60 in specimens greater than 200 mm SL. Teeth are present or absent on side of tongue. Gill rakers on first arch number 6 on upper limb and 11 on lower limb. Measurements are expressed as percent of SL: head length 29%– 42%, snout length 8%–13%, eye length 8%–12%, interorbital width 7%–14%, pectoral fin length 18%– 23%, body depth 22%–30%. Head, snout, eye, interorbital, pectoral fin, and body depth proportions decrease with growth. Pectoral fin

has 14 to 18 rays. First dorsal fin has 13 to 16 spines, and second dorsal fin has 1 spine and 10 to 13 rays. Anal fin has 1 spine and 9 to 12 rays. Scales are oblong in shape and possess parallel ridges that increase in number with growth. Scales in lateral series from upper opercular margin to origin of caudal keels number 83 to 95. Vertebrae number 45 to 51. This species occurs worldwide in tropical and subtropical seas but is rare outside the Atlantic Ocean. In the western Atlantic it occurs from Nova Scotia and Bermuda to Panama, including the eastern Gulf of Mexico and the Greater Antilles. It is generally limited to oceanic epipelagic waters between 20C and 25C. Specimens may penetrate greater depths as they grow. Maximum known size is 500 mm SL. references: Grey 1955a; Haedrich 1967, 1986e, 2002d; Haedrich and Horn 1972; Robins and Ray 1986; Scott and Scott 1988; Last 2001d.

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817

STROMATEIDAE Butterfishes

Butterfishes are deep bodied and compressed, with a single dorsal fin and pelvic fins absent in adults. Dorsal profile of head is moderately to strongly elevated, and snout is moderately blunt to very blunt. Nostril is paired, and nares are close set. Eye is moderate to rather small and laterally located, with inconspicuous adipose tissue. Mouth is terminal to subterminal, small, and moderately oblique. Supramaxilla is absent. Jaw teeth are small, laterally flattened, and unicuspid to tricuspid. Upper jaw is not protrusible. Teeth are absent from vomer, palatine, and basibranchs. Posterior margins of preoperculum and operculum are entire. Branchiostegal rays number 5 or 6. Gill membranes are free of isthmus except for one genus. Pectoral fin is long and pointed and inserts on lower flank, with base moderately to strongly oblique. Dorsal fin originates over or behind pectoral fin base, is moderately to strongly falcate, and has 38 to 53 elements. Pelvic fin is present only in juveniles of one genus (Stromateus). Anal fin is similar in shape and size to dorsal fin and has 34 to 47 elements. Caudal fin is shallowly to deeply forked. Scales are cycloid and deciduous, and cover body but not head. Lateral line is single and continuous and follows dorsal profile. Anterior section of esophagus is enlarged and specialized to form pharyngeal sac that has papillae in upper and lower halves. Papillae have stellate bases, with teeth located along central stalk. Vertebrae number 29 to 48. Stromateids occur worldwide in tropical to temperate seas along continental margins, with exception of the southwestern Pacific and southern Indian Oceans. There are about 13 species in three genera, 3 species in a single genus in the western Central Atlantic, and two species in the Gulf of Mexico. references: Haedrich 1967, 1986c, 2002e; Nelson 1994; Smith 1997; Last 2001e.

Key to the Species of the Gulf of Mexico (Adapted from Haedrich and Horn 1972; Haedrich 2002e) 1a. Dorsal and anal fins only slightly falcate; row of about 17 to 25 relatively large pores below anterior half of dorsal fin; premaxillary teeth usually tricuspid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Peprilus burti p. 818 1b. Dorsal and anal fins moderately to strongly falcate; no row of pores below anterior half of dorsal fin; premaxillary teeth simple or unicuspid . . . . . . . . . . Peprilus paru p. 819

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818 PERCIFORMES

Peprilus burti Fowler, 1944 Gulf butterfish

Peprilus burti has relatively large eyes, two to four bladelike spines anterior to rays of dorsal and anal fins, and a row of 17 to 25 relatively large pores immediately below anterior half of dorsal fin. Dorsal and ventral profiles of body are strongly convex and similar, and snout is blunt. Anterior naris is round, and posterior naris is slitlike. Mouth is terminal, with upper jaw extending to anterior margin of eye. Branchiostegal rays number 6. Gill rakers on first arch number 21 to 26. Measurements are expressed as percent of SL: head length 26%– 39%, snout length 5%–10%, eye length 7%– 14%, upper jaw length 7%–13%, pectoral fin length 23%–39%, body depth 46%– 64%. Pectoral fin inserts just below midline, is long and winglike, and has 19 to 23 rays. Dorsal fin is moderately falcate and has 38 to 48 rays. Pelvic fin is absent, but spinelike element on pelvic girdle projects through skin midventrally below pectoral fin base. Anal fin is similar in shape and size to dorsal fin and has 35 to 42 rays. Caudal peduncle is short, slender, and compressed. Caudal fin is deeply forked. Scales

are small and very deciduous, and cover body and side of head. Lateral line scales number 90 to 100. Gas bladder is apparently functional only in juveniles less than 80 to 100 mm SL. Vertebrae number 29 to 31: 13 precaudal and 16 to 18 caudal. Color is iridescent bluish or greenish silver, without dark spots on upper part of body. This species occurs in the western Atlantic from Virginia to Florida and the Gulf of Mexico. In the Gulf it is widespread along the continental shelf from Tampa Bay, FL, to Yucatán. Maximum known size is 200 mm TL, and maximum known age is about two and onehalf years.

references: Fowler 1944; Caldwell 1961; Collette 1963; Haedrich 1967, 2002e; Horn 1970; Haedrich and Horn 1972; Hoese and Moore 1977, 1998; Perschbacher et al. 1979; Castro-Aguirre and Márquez-Espinoza 1981; Robins and Ray 1986; Murphy and Chittenden 1991; Boschung 1992; Schaldach et al. 1997 (as P. triacanthus); Smith 1997; Castro-Aguirre et al. 1999.

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STROMATEIDAE 819

Peprilus paru (Linnaeus, 1758) Harvestfish

Peprilus paru has extremely falcate dorsal and anal fins and two to four bladelike spines anterior to rays of dorsal and anal fins. Dorsal and ventral profiles of body are slightly convex and similar, and snout is blunt. Anterior naris is round, and posterior naris is slitlike. Mouth is terminal, with upper jaw extending to anterior margin of eye. Branchiostegal rays number 6. Gill rakers on first arch number 20 to 23. Measurements are expressed as percent of SL: head length 25%– 40%, snout length 4%– 8%, eye length 6%–15%, upper jaw length 7%–11%, pectoral fin length 27%– 45%, body depth 57%– 88%. Pectoral fin inserts just below midline, is long and winglike, and has 18 to 24 rays. Dorsal fin has 38 to 47 rays. Pelvic fin is absent, but spine on pelvic girdle projects through skin midventrally below pectoral fin base. Anal fin is similar in shape and size to dorsal fin, though slightly shorter, and has 35 to 45 rays. Caudal peduncle is short, slender, and compressed. Caudal fin is deeply forked.

Scales are small and very deciduous and cover body and side of head. Lateral line scales number 80 to 95. Gas bladder is apparently functional only in juveniles less than 80 to 100 mm SL. Vertebrae number 29 to 31: 12 or 13 precaudal and 16 to 18 caudal. Color is iridescent bluish or greenish silver, without dark spots on upper part of body. This neritic species occurs in the western Atlantic from Chesapeake Bay to northern Argentina, including the entire Gulf of Mexico and the Greater Antilles, except for Cuba and Jamaica. Maximum known size is 280 mm SL.

references: Mansueti 1963; Haedrich 1967, 2002e; Horn 1970; Haedrich and Horn 1972; Hoese and Moore 1977; Shimizu 1983e; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Hoese and Moore 1998; CastroAguirre et al. 1999 (last eight references as P. alepidotus).

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820

PLEURONECTIFORMES Pleuronectiforms are a highly corroborated monophyletic group derived from an unknown taxon that is thought to be currently classified in Perciformes. The order consists of about 14 families, and 6 of these occur in the Gulf of Mexico.

Key to the Families of the Gulf of Mexico (Adapted from Gutherz 1967, in part) 1a. Margin of preoperculum free of skin and scales . . . . . . . . 2 1b. Margin of preoperculum covered with skin and scales . . . 5 2a. Eyes on right side of head. . . . . . . . Poecilopsettidae p. 861

2b. Eyes on left side of head . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. Bases of both pelvic fins short and nearly symmetrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralichthyidae p. 832

3b. Bases of both pelvic fins long or base of pelvic fin on ocular side long and base of pelvic fin on blind side short; in either case, pelvic fin bases moderately to strongly asymmetrical .............................................4 4a. Bases of both pelvic fins long . . . . . Scophthalmidae p. 830

4b. Base of pelvic fin on ocular side long and base of pelvic fin on blind side short. . . . . . . . . . . . . . . . . . . Bothidae p. 821

5a. Eyes on right side of head. . . . . . . . . . . . . Achiridae p. 864

5b. Eyes on left side of head . . . . . . . . . . Cynoglossidae p. 872

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821

BOTHIDAE

Lefteye flounders

Lefteye flounders are asymmetrical and greatly compressed, with both eyes on the left side of the head, dorsal fin originating on neurocranium, and pelvic fins asymmetrical. Head profile is convex to deeply concave. Eyes protrude slightly to markedly above body surface, with lower eye located at about mid-depth of head and moderately to distinctly separated from and anterior to upper eye. Nostrils are paired and located on each side of head. Mouth is moderate to very large, with one to several rows of small to moderate-sized teeth. Jaws are either equally developed on each side or asymmetrical, and teeth are either equally developed in jaws or better or exclusively developed on blind side. Supramaxilla is absent. Teeth are lacking in vomer and palatine. Preoperculum is free of skin and scales. Gill membranes are joined at ventral margin of head. One or both pectoral fins are present (fin on blind side is absent in some species), and fin rays are unbranched. Dorsal fin originates on blind side of head in front of right eye. Base of pelvic fin on ocular side is located on ventral midline and is longer and slightly to distinctly anterior to fin on blind side. Pelvic fin lacks spine and has 6 unbranched rays. Dorsal and anal fins are separate from caudal fin. Caudal fin generally has 17 rays, and 9 to 13 of these are branched. Lateral line is present on ocular side and present or absent on blind side. On ocular side, lateral line does not extend onto head below left eye. Urinary papilla is located on ocular side at level of first anal fin ray. Anus is located on blind side. Lefteye flounders are found worldwide in tropical to warm temperate seas. They are benthic and usually limited to soft bottoms on continental and insular shelves. Only a few species occur deeper than 200 m. There are about 115 species in 20 genera, 14 or 15 species in 5 genera in the western Central Atlantic, and 7 species in 5 genera in the Gulf of Mexico. references: Norman 1934; Nelson 1994; Hensley 1995; Hensley and Amaoka 2001; Munroe 2002a.

Key to the Species of the Gulf of Mexico (Adapted from Gutherz 1967) 1a. Pectoral fin absent on blind side of body . . . . . . . . . . . . . . . . . . . . . . . Monolene sessilicauda p. 828 1b. Pectoral fin present on blind side of body . . . . . . . . . . . . . 2 2a. Ocular-side pelvic fin base considerably longer than blindside pelvic fin base, with anterior rays distinctly anterior to those of blind-side fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2b. Ocular-side pelvic fin base about same length as blind-side

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822 PLEURONECTIFORMES

3a.

3b.

4a.

4b.

5a.

5b.

6a.

6b.

pelvic fin base, with anterior rays not anterior to those of blind-side fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Body depth 50% of SL or more; maxilla not extending to vertical line through middle of lower eye; interorbital space greater than eye diameter . . . . . . . . . . . . . . . . . . . . . . . . . 4 Body depth less than 40% of SL; maxilla extending to or beyond vertical line through posterior margin of lower eye; interorbital space less than eye diameter . . . . . . . . . . . . . . . . . . . . . Chascanopsetta lugubris p. 826 Body depth greater than 60% of SL; eye diameter more than 23% of head length; dorsal fin rays 76 to 91; anal fin rays 58 to 68 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Body depth 60% of SL or less; eye diameter less than 23% of head length; dorsal fin rays 90 to 105; anal fin rays 70 to 80 . . . . . . . . . . . . . . . . . . . . . . . . Bothus lunatus p. 823 Caudal fin with two large spots horizontally arranged on median rays; body spotting and mottling indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bothus robinsi p. 825 Caudal fin with no spots or with two spots vertically arranged; body spotting and mottling distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . Bothus ocellatus p. 824 Maxilla not extending beyond vertical line through anterior margin of eye; upper jaw about 25% of head length; interorbital ridge with spines; tentacle on eyes of both males and females (may be absent on large males) . . . . . . . . . . . . . . . . . . . . . . . . . . Engyophrys senta p. 827 Maxilla extending beyond vertical line through anterior margin of eye; upper jaw 32% to 45% of head length; interorbital ridge without spines; eyes without tentacles . . . . . . . . . . . . . . . . . . . . . . . Trichopsetta ventralis p. 829

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BOTHIDAE 823

Bothus lunatus

(Linnaeus, 1758)

Bothus lunatus is very deep bodied, with a small mouth, widely separated eyes, and distinct blue rings and curved spots on body. Snout has a notch in front of lower eye. Mouth is downturned, with upper jaw extending posteriorly to or slightly beyond anterior margin of lower eye. Jaw teeth are small and sharp pointed and arranged in two rows. Distance between eyes is about one eye diameter in both sexes. Anterior margin of upper eye is on a vertical line with posterior margin of lower eye in specimens larger than 100 mm SL, and on a vertical with middle of lower eye in specimens of about 50 mm SL. Eye tentacles are poorly developed and not present on specimens smaller than 100 mm SL. Gill rakers on first arch of ocular side are short and number 8 to 10 on lower limb. Measurements are expressed as percent of SL: head length 26%–31%, body depth 54%– 59%; and as percent of head length: eye diameter 16%–22%, upper jaw length 30%–35%. Ocular-side pectoral fin has 10 to 12 rays and is longer than that on blind side. Ocular-side pectoral fin rays are elongated in mature males. Dorsal fin has 91 to 99 rays. Base of ocularside pelvic fin extends to tip of urohyal. Anal

fin originates anterior to pectoral fin base and has 71 to 76 rays. Caudal fin is bluntly pointed. Scales are ctenoid on ocular side and cycloid on blind side. Lateral line is arched over pectoral fin on ocular side, and lateral line scales number 83 to 95. Color on ocular side is grayish brown, with numerous blue rings and curved spots over entire surface and three diffuse dark spots on straight part of lateral line. Ocular-side pectoral fin is banded in large individuals. This species occurs from shore to about 65 m in the western Atlantic from the east coast of Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it occurs along the northern coast of Yucatán. It is most common on sandy bottoms near coral reefs, but it also inhabits sea-grass beds and mangrove habitats. Food consists of octopods, crustaceans, and small ray-finned fishes. Maximum known size is 450 mm TL. references: Norman 1934; Gutherz 1967; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999; Munroe 2002a.

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824 PLEURONECTIFORMES

Bothus ocellatus (Agassiz, 1831) Ocular flounder

Bothus ocellatus is very deep bodied, with a small mouth, widely separated eyes, and distinct spotting and mottling on body. Mouth is downturned, with upper jaw extending posteriorly to or slightly beyond anterior margin of lower eye. Jaw teeth are small and sharp pointed and arranged in two rows. Males have a spine at tip of snout and a bony protuberance above lower eye. Distance between eyes is about equal to eye diameter in adult females and greater than eye diameter in adult males but less than eye diameter in juveniles. Upper eye possesses fleshy ridge along posterodorsal margin, and lower eye is anterior to upper eye and possesses fleshy ridge along posteroventral margin. Gill rakers on first arch of ocular side are short and number 0 to 6 on upper limb and 7 to 10 on lower limb. Measurements are expressed as percent of SL: head length 24%– 26%, snout length 5%– 6%, eye diameter 5%– 6%, upper jaw length 6%–7%, ocular-side pectoral fin length 16%–25%, body depth 61%– 67%. Pectoral fin on ocular side has 8 to 10 rays and is longer than blind-side fin. Uppermost ocular-side pectoral fin rays are elongated in mature males. Dorsal fin has 76 to 91 rays. Base of ocular-side pelvic fin extends

to tip of urohyal. Anal fin originates anterior to pectoral fin base and has 58 to 68 rays. Caudal fin is bluntly pointed. Scales are ctenoid on ocular side and cycloid on blind side. Lateral line is arched over pectoral fin on ocular side, and lateral line scales number 70 to 78. Precaudal vertebrae number 10, and caudal vertebrae number 25 to 27. Color on ocular side is pale brown to gray, with dark mottling or spotting. Caudal fin often has two vertically arranged dark spots on base above and below median rays. This species occurs between 18 and 91 m in the western Atlantic from Long Island and Bermuda to Rio de Janeiro, Brazil, including the eastern and southern Gulf of Mexico, the Florida Keys, the Bahamas, and the Antilles. It occurs on soft bottoms. Food consists of polychaetes and crustaceans. Maximum known size is 160 mm SL. references: Norman 1934; Gutherz 1967; Böhlke and Chaplin 1968; Topp and Hoff 1972; Moore 1975a; Robins and Ray 1986; Boschung 1992; Cervigón 1993a; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Munroe 2002a.

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BOTHIDAE 825

Bothus robinsi Topp and Hoff, 1972 Two-spot flounder Bothus robinsi is deep bodied, with a small mouth, widely separated eyes, and two horizontally arranged spots on caudal fin. Mouth is downturned, with upper jaw extending posteriorly to or slightly beyond anterior margin of lower eye. Jaw teeth are small and sharp pointed and arranged in two rows. Males have a sharp-pointed spine on snout. Distance between eyes is about equal to eye width in adult females and about 2 times eye width in adult males, but less than eye width in juveniles. Eyes lack fleshy ridges. Gill rakers on first arch of ocular side are short and number two to seven on upper limb and five to nine on lower limb. Measurements are expressed as percent of SL: head length 23%–28%, snout length 4%– 6%, eye diameter 6%–7%, upper jaw length 5%– 8%, ocular-side pectoral fin length 21%–32%, body depth 65%–76%. Pectoral fin on ocular side has 8 to 11 rays and is longer than blindside fin. Uppermost pectoral fin rays on ocular side are elongated in mature males. Dorsal fin has 78 to 90 rays. Base of pelvic fin on ocular side extends to tip of urohyal. Anal fin originates anterior to pectoral fin base and has 59 to 68 rays. Caudal fin is bluntly pointed. Scales are ctenoid on ocular side and cycloid on blind side. Lateral line is arched over pectoral fin on

ocular side, and lateral line scales number 70 to 77. Precaudal vertebrae number 10, and caudal vertebrae number 26 to 28. Color on ocular side is light tan, with brown spotting and mottling, or dark brown, without spotting and mottling. Caudal fin has two horizontally arranged dark spots along median rays. This species occurs between 18 and 91 m in the western Atlantic from North Carolina and Bermuda to Brazil, including the entire Gulf of Mexico, the Florida Keys, the Bahamas, and the Antilles. It occurs on soft bottoms. Feeding takes place during daylight, and food consists of bryozoans, brachiopods, ophiuroids, chaetognaths, scaphopods, gastropods, bivalves, polychaetes, isopods, shrimps (carideans and penaeids), and crabs (majids, galatheids, and pagurids). Maximum known size is 222 mm SL, but specimens greater than 150 mm are rare.

references: Gutherz 1967 (as Bothus sp.); Böhlke and Chaplin 1968 (as Bothus sp.); Topp and Hoff 1972; Hoese and Moore 1977, 1998; Matsuura 1983p (as Bothus sp.); Robins and Ray 1986; Boschung 1992; Cervigón 1993a; Smith-Vaniz et al. 1999; Munroe 2002a.

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826 PLEURONECTIFORMES

Chascanopsetta lugubris Pelican flounder

Alcock, 1894

Chascanopsetta lugubris is elongate and tapering, with upper jaw extending beyond posterior margin of lower eye and base of pelvic fin on ocular side extending forward onto urohyal. Tip of lower jaw projects beyond upper jaw. Eyes are close set and equally spaced in males and females. Jaws and jaw teeth are about equally developed on each side. Jaw teeth are slender, curved, depressible, and arranged in a single row. Gill rakers on first arch are absent or consist of one or two rudiments. Measurements are expressed as percent of SL: head length 20%–21%, snout length 4%– 6%, eye diameter 4%–5%, upper jaw length 16%– 19%, ocular-side pectoral fin length 10%– 15%, body depth 27%–31%. Pectoral fin on ocular side is larger than fin on blind side, and each fin has 14 to 17 rays. Dorsal fin has 112 to 122 rays, with anterior rays connected by membrane only at their bases. Base of pelvic fin

on ocular side extends to tip of urohyal. Anal fin has 77 to 85 rays. Lateral line is arched over pectoral fin base. Lateral line scales number about 190. Body is covered with cycloid scales. Color of ocular side is grayish to yellowish brown, with dusky fin membranes and with or without dark spots and blotches. Peritoneum is black and is visible through abdominal flesh. Blind side is white. This species occurs in the tropical Atlantic, western Pacific, and Indian Oceans. In the western Atlantic it occurs between 229 and 549 m, from the east coast of Florida to Brazil, including the entire Gulf of Mexico. Maximum known size is 300 mm TL.

references: Norman 1934; Gutherz 1967; Matsuura 1983p; Robins and Ray 1986; Cervigón 1993a; Smith 1997; Munroe 2002a.

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BOTHIDAE 827

Engyophrys senta Spiny flounder

Ginsburg, 1933

Engyophrys senta is ovate in profile, with eyes separated by a narrow ridge and three to five spines on interorbital ridge. Upper orbit bears spine along anterodorsal margin, and lower orbit bears spine along anterior margin. Posterior regions of eyes bear moderately long tentacles in juveniles and females. Maxilla extends to anterior margin of lower eye. Jaws on each side are equally developed, but small teeth are mostly confined to blind side. Gill rakers on first arch are very short and number zero to three on upper limb and four to seven on lower limb. Measurements are expressed as percent of SL: head length 21%–24%, snout length 3%–5%, eye diameter 6%– 8%, upper jaw length 4%–5%, ocular-side pectoral fin length 12%–15%, body depth 53%–59%. Pectoral fin on ocular side is longer than fin on blind side and has 8 to 10 (usually 9) rays. Dorsal fin has 74 to 83 rays. Caudal fin is bluntly pointed. Scales are ctenoid on ocular side and cycloid on blind side. Lateral line is strongly arched over pectoral fin. Lateral line scales number

about 50. Precaudal vertebrae number 10, and caudal vertebrae number 27 or 28. Ocular side is tan to brownish, with three dark blotches along lateral line and along dorsal and ventral margins of body. Blotches occur at beginning of straight part of lateral line, on caudal peduncle, and slightly posterior to mid-distance between them. Blotches along dorsal and ventral margins often appear as narrow dark rings. Blind side is white in juveniles and females but is dusky in males, with three to seven curved bars on anterior third of body. This species occurs between 37 and 183 m in the western Atlantic from North Carolina and the Bahamas to Brazil, including the northern and southern Gulf of Mexico, the Florida Keys, and Trinidad. Maximum known size is 100 mm TL.

references: Gutherz 1967; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Munroe 2002a.

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828 PLEURONECTIFORMES

Monolene sessilicauda Deepwater flounder

Goode, 1880

Monolene sessilicauda is oval shaped and elongate, with eyes narrowly separated and with pectoral fin on ocular side only. Maxilla extends slightly beyond anterior margin of lower jaw. Jaws and jaw teeth are about equally developed on each side. Jaw teeth are very small and arranged in a single row. Gill rakers are moderately developed and number about 4 on upper limb and 8 to 10 on lower limb. Measurements are expressed as percent of SL: head length 17%–20%, snout length 3%– 4%, eye diameter 5%– 6%, upper jaw length 4%–5%, ocular-side pectoral fin length 10%–11%, body depth 30%–33%. Pectoral fin has 11 to 14 rays. Dorsal fin has 92 to 107 rays. Caudal fin is bluntly pointed. Scales are ctenoid on ocular side and cycloid on blind side. Lateral line is strongly elevated over pectoral fin. Lateral line scales number 88 to 94. Precaudal verte-

brae number 10 or 11, and caudal vertebrae number 36. Ocular side is tan to brownish, with darker blotches arranged into bands. Pectoral fin has a clear base and dark spots and blotches forming bands on distal half of fin. Caudal fin may have a large black blotch. Blind side is white. This species occurs in the western Atlantic from New England to Brazil, including the northern and southern Gulf of Mexico and the Caribbean Sea, between 78 and 549 m. Monolene antillarum Norman is apparently a junior synonym of this species. Maximum known size is 130 mm SL. references: Norman 1934 (as M. antillarum, in part); Gutherz 1967; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Munroe 2002a.

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BOTHIDAE 829

Trichopsetta ventralis Sash flounder

(Goode and Bean, 1885)

Trichopsetta ventralis is ovate and relatively deep, with eyes separated by a narrow ridge bearing a narrow groove, pectoral fin on blind side longer than fin on ocular side, and base of pelvic fin on ocular side slightly longer than its antimere on blind side. Posterior naris of blind side lacks membranous valve. Lower eye is anterior to upper eye. Maxilla extends to pupil of lower eye. Jaws and teeth are about equally developed on each side. Jaw teeth are small, pointed, slender, and uniserial in both jaws. Gill rakers on first arch are relatively long and slender, are absent on upper limb, and number 9 to 11 on lower limb. Measurements are expressed as percent of SL: head length 26%– 29%, snout length 5%– 6%, eye diameter 6%– 7%, upper jaw length 11%–13%, ocular-side pectoral fin length 16%–18%, body depth 41%–50%. Pectoral fin on ocular side has 12 or 13 rays, and blind-side pectoral fin has 7 to 10 rays. Dorsal fin has 89 to 95 rays. Blind-

side pelvic fin is considerably longer than ocular-side pelvic fin in mature males. Anal fin has 69 to 75 rays. Caudal fin is broadly rounded to bluntly pointed. Scales are ctenoid on upper side and cycloid on lower side. Lateral line is strongly arched over pectoral fin. Lateral line scales number 63 to 68. Color on ocular side is brown, with darker spots and speckles. Blotches are often present at beginning of straight portion of lateral line and on caudal peduncle. Males have a dark blotch on anterior section of anal fin. Blind side is white. This species occurs between 30 and 400 m in the western Atlantic from the northern and southern Gulf of Mexico and Suriname. Maximum known size is 200 mm TL. references: Norman 1934; Gutherz 1967; Anderson and Gutherz 1967; Hoese and Moore 1977, 1998; Matsuura 1983p; Robins and Ray 1986; Boschung 1992; Munroe 2002a.

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830

SCOPHTHALMIDAE

Scophthalmids are moderately to very deep bodied, with both eyes on the left side of the head, dorsal fin originating on neurocranium, ocular-side pelvic fin located on ventral midline, and both pelvic fin bases elongated. Head profile is convex to moderately concave. Eyes are located on same vertical line, or lower eye is slightly anterior to upper eye. Nostrils are paired and located on either side of head. Mouth is terminal and large to very large, with one to several rows of small to moderate-sized teeth. Lower jaw is generally prominent. Jaws are equally developed on each side, and teeth are equally developed in jaws. Teeth are present in vomer. Preoperculum is free of skin and scales. Gill membranes are connected. Both pectoral fins are present, and generally some fin rays of ocular-side fin are branched. Dorsal fin originates on blind side of head at about level of blind-side nares. Pelvic fins extend anteriorly onto urohyal, lack spines, and have unbranched rays. Dorsal and anal fins are separate from caudal fin, and dorsal fin occasionally inserts on blind side of caudal peduncle. Scales are present or are replaced by bony tubercles. Lateral line is present on both sides. Scophthalmids occur in the North Atlantic in warm temperate and boreal seas. They are benthic and generally associated with soft bottoms. There are 8 species in five genera, and 1 species occurs in the western Central Atlantic and possibly the Gulf of Mexico. references: Norman 1934; Gutherz 1967; Nielsen 1986; Nelson 1994; Munroe 2002b.

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SCOPHTHALMIDAE 831

Scophthalmus aquosus Windowpane

(Mitchill, 1815)

Scophthalmus aquosus is deep bodied, with branchial septum entire, anterior rays of dorsal fin branched, and small cycloid scales. Anterior nares on both sides of head have membranous flaps on posterior margin. Maxilla extends to middle or posterior section of eye. Anterior end of ocular-side maxilla possesses a bony tubercle. Jaw teeth are small, curved, and arranged in narrow bands. Gill rakers on first arch are slender and of moderate size and number 8 on upper limb and 22 to 26 on lower limb. Measurements are expressed as percent of SL: head length 28%–30%, snout length 6%–7%, orbit diameter 6%–7%, upper jaw length 12%–15%, ocular-side pectoral fin length 19%–22%, body depth 61%– 68%. Pectoral fin on ocular side has 11 rays. Dorsal fin has 64 to 71 rays, with anterior rays relatively long, branched, and free of membrane over greater part of length. Pelvic fin on

ocular side originates slightly anterior to its antimere and has 6 rays. Anal fin has 48 to 55 rays. Scales are small and cycloid. Lateral line is equally developed on each side and is strongly arched over pectoral fin. Precaudal vertebrae number 11, and caudal vertebrae number 23 to 25. Ocular side is brown, with many dark and lighter spots on body and fins. This species occurs in the western Atlantic from the Gulf of St. Lawrence to Florida to a depth of 46 m. There is a single record of this species from the Gulf of Mexico (Baughman 1950), off Texas, but this record has not been verified. This species is found on soft bottoms. Maximum known size is 450 mm TL.

references: Norman 1934; Baughman 1950; Gutherz 1967; Hoese and Moore 1977, 1998; Robins et al. 1986; Munroe 2002b.

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832

PARALICHTHYIDAE

Paralichthyids are greatly compressed, with both eyes usually on the left side of the head, dorsal fin originating on neurocranium, and pelvic fin bases about equal in length. Head profile is convex to deeply concave. Eyes protrude slightly above body surface, with lower eye located at about mid-depth of head and moderately to widely separated from and anterior to upper eye. Nostrils are paired and located on each side of head, with anterior naris bearing flap on posterior margin. Mouth is terminal and moderate to very large, with one to several rows of small to moderatesized teeth. Lower jaw is more or less prominent. Jaws are either equally developed or asymmetrical, and teeth are either equally developed in jaws or better or exclusively developed on blind side. Supramaxilla is absent. Teeth are lacking in vomer and palatine. Preoperculum is free of skin and scales. Gill membranes are connected across isthmus. Both pectoral fins are usually present, though the one on blind side is occasionally absent, and fin rays are branched. Dorsal fin originates on blind side of head above or anterior to upper eye. Pelvic fins are symmetrical or asymmetrical, with bases on either side of ventral midline or with base of ocular-side fin on midline and base of blind-side fin on side of midline. Pelvic fin lacks spine and has six or, rarely, five unbranched rays. Dorsal and anal fins are separate from caudal fin. Lateral line is present on ocular side and present or absent on blind side. On ocular side, lateral line extends anteriorly onto head, to above lower eye in some species. Urinary papilla is located either on ocular or blind side, opposite first anal ray, and anus is located on blind side. Paralichthyids occur worldwide in tropical to temperate seas and occasionally in freshwater. All species are benthic and are usually associated with soft bottoms. There are 85 species in 16 genera worldwide, 33 species in 8 genera in the western Central Atlantic, and 22 or 23 species in 8 genera in the Gulf of Mexico. references: Norman 1934; Nelson 1994; Munroe 2002c.

Key to the Species of the Gulf of Mexico (Adapted from Munroe 2002c) 1a. Lateral line on ocular side with high arch over pectoral fin base; pelvic fins symmetrically arranged or nearly symmetrically arranged on either side of midventral line; branched caudal fin rays 13 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Lateral line on ocular side without high arch over pectoral fin base; pelvic fins asymmetrically arranged, base of ocular-side pelvic fin on midventral line; branched caudal fin rays 11 (rarely 10 or 12). . . . . . . . . . . . . . . . . . . . . . . . . . 8

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PARALICHTHYIDAE 833

2a. Ocular-side pelvic fin rays longer than blind-side pelvic fin rays in adults; anterior dorsal fin rays elongated in adults; first gill arch with 6 to 9 gill arches on lower limb; dorsal fin rays 58 to 79 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2b. Ocular-side pelvic fin rays not longer than blind-side pelvic fin rays in adults; anterior dorsal fin rays not elongated in adults; first gill arch with 7 to 12 gill rakers on lower limb; dorsal fin rays 71 to 95 . . . . . . . . . . . . . . . . . . . . . . . . . . 5 3a. Origin of dorsal fin distinctly anterior to eyes; dorsal profile of head evenly convex; scales on ocular side cycloid and embedded; dorsal fin rays 58 to 65; three ocelli on ocular side, with one above pectoral fin, and two at midbody, on either side on midline . . . . . . Gastropsetta frontalis p. 853 3b. Origin of dorsal fin over or slightly anterior to front of eyes; dorsal profile of head concave in front of upper eye; scales on ocular side ctenoid; dorsal fin rays 67 to 79; three or four ocelli on ocular side but not arranged as above (3a) . . . . . 4 4a. Four ocellated spots on ocular side, anterior spot above curved section of lateral line, posterior spots arranged in triangle, with two on vertical line at midbody and one near caudal peduncle and on midline . . . . . . . . . . . . . . . . . . . Ancylopsetta quadrocellata p. 838 4b. Three ocellated spots on ocular side arranged in triangle posterior to curved section of lateral line . . . . . . . . . . . . . . . . . . . . . . . . Ancylopsetta dilecta p. 837 5a. Distinct ocellated spots on ocular side . . . . . . . . . . . . . . . 6 5b. No distinct ocellated spots on ocular side. . . . . . . . . . . . . 7 6a. Eyes close set, separated by narrow ridge; first gill arch with 7 to 10 gill rakers on lower limb; lateral line scales 63 to 95; four ocellated spots on ocular side of body arranged in a trapezoid pattern; anal fin rays 58 to 72 . . . . . . . . . . . . . . . . . . . . . . Hippoglossina oblonga p. 854 6b. Eyes separated by flat interspace, without ridge; first gill arch with 8 to 12 gill rakers on lower limb; lateral line scales 78 to 117; three to five ocellated spots on ocular side but not arranged in trapezoid pattern; anal fin rays 53 to 63 . . . . . . . . . . . . . . . . . . . Paralichthys albigutta p. 855 7a. Body depth greater than 47% of SL; lateral line scales 104 to 117; blind side dusky in large specimens . . . . . . . . . . . . . . . . . . . . Paralichthys squamilentus p. 857 7b. Body depth 47% of SL or less; lateral line scales 85 to 100; blind side immaculate or dusky . . . . . . . . . . . . . . . . . . . . . Paralichthys lethostigma p. 856 8a. Maxilla 24% to 29% of head length, nearly reaching vertical line through anterior margin of eye; blind-side jaws arched; no enlarged teeth . . . . . . . . . . . . . . . . . . . . . . . . . 9 8b. Maxilla greater than 29% of head length, reaching vertical line through mideye; jaws on blind side not arched; front teeth in jaws enlarged. . . . . . . . . . . . . . . . . . . . . . . . . . . 11

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834 PLEURONECTIFORMES

9a. Accessory scales absent; first gill arch with six to nine (modally seven) gill arches on lower limb; snout without scales . . . . . . . . . . . . . . . . . . . . . . Etropus crossotus p. 850 9b. Accessory scales present; first gill arch with three to six (rarely seven) gill rakers on lower limb; snout with scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10a. Snout scaled anterior to line connecting ocular- and blindside nares in specimens greater than 30 mm SL; ctenii on snout scales highly modified, especially in large males; primary scales on blind side ctenoid (ctenii visible only on specimens greater than 50 mm SL; dark circles on upper side absent . . . . . . . . . . . . . . . . . . . Etropus rimosus p. 852 10b. Snout not scaled anterior to line connecting ocular- and blind-side nares (rarely one or two scales anterior to line in large specimens); ctenii on snout scales simple; primary scales on blind side cycloid; often with four to six dark circles on upper side arranged above and below lateral line (may be indistinct in darkly pigmented specimens . . . . . . . . . . . . . . . . . . . . . . . Etropus cyclosquamus p. 851 11a. Upper jaw with two rows of immovable teeth . . . . . . . . 12 11b. Upper jaw with single row of immovable teeth. . . . . . . . 19 12a. Body depth usually 48% of SL or greater; dorsal fin rays 74 to 85; anal fin rays 59 to 68 . . . Syacium gunteri p. 858 12b. Body depth usually 45% of SL or less; dorsal fin rays 82 to 94; anal fin rays 64 to 75 . . . . . . . . . . . . . . . . . . . . . . . . 13 13a. Specimens greater than 120 mm SL . . . . . . . . . . . . . . . . 14 13b. Specimens less than 120 mm SL . . . . . . . . . . . . . . . . . . . 18 14a. Interorbital width greater than 75% of lower eye diameter . . . . . . . . . . . . . . . . . . Syacium papillosum (males) p. 860 14b. Interorbital width less than 75% of lower eye diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 15a. Ocular-side pectoral fin rays not elongate, less than 25% of SL (females) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 15b. Ocular-side pectoral fin rays elongate, greater than 25% of SL (males) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16a. Interorbital width 25% to 35% of lower eye diameter in specimens 120 to 150 mm SL, and 60% in specimens about 220 mm SL; ocular side plain brown, with little or no mottling . . . . . . . . . . . . . . . . . . Syacium papillosum p. 860 16b. Interorbital width about 20% of lower eye diameter in specimens 120 to 150 mm SL, and 27% in specimens to 195 mm SL; ocular side light tan to brown, with mottling on body and fins and large black spots on lateral line . . . . . . . . . . . . . . . . . . . . . . . . . Syacium micrurum p. 859 17a. Interorbital width 30% to 70% of lower eye diameter in specimens 120 to 150 mm SL, 50% to 90% in specimens 150 to 180 mm SL, and greater than 75% in specimens greater than 180 mm SL. . . . . . Syacium papillosum p. 860

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PARALICHTHYIDAE 835

17b. Interorbital width less than 35% of lower eye diameter in specimens 120 to 150 mm SL, less than 50% in specimens 150 to 180 mm SL, and less than 75% in specimens greater than 180 mm SL . . . . . . Syacium micrurum p. 859 18a. Snout length 54% to 74% of shortest distance between tip of snout and orbit of upper eye; interorbital width generally greater than 15% of lower eye diameter . . . . . . . . . . . . . . . . . . . . . . . . Syacium papillosum p. 860 18b. Snout length 80% to 92% of shortest distance between tip of snout and orbit of upper eye; interorbital width generally less than 15% of lower eye diameter . . . . . . . . . . . . . . . . . . . . . . . . . Syacium micrurum p. 859 19a. Scales ctenoid; gill rakers on first arch slender and moderately long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 19b. Scales cycloid; gill rakers on first arch stout and short. . . 27 20a. Snout with osseous protuberance; upper jaw length less than 33% of head length (slightly less in some specimens of C. spilopterus); body depth 34% to 43% (usually less than 40%) of SL. . . . . . . . . Citharichthys arctifrons p. 840 20b. Snout without osseous protuberance (males of some species may have labial or cephalic spination); upper jaw length usually greater than 33% of head length; body depth greater than 40% of SL . . . . . . . . . . . . . . . . . . . . . . . . . 21 21a. Dorsal fin rays 88 or more; anal fin rays 68 or more; lower jaw distinctly included in upper jaw when mouth is closed; several large canines overhanging lower jaw; caudal fin with or without two large spots, if spots present, arranged one above and one below median rays . . . . . . . . . . . . . . . . . . . . . . Citharichthys dinoceros p. 842 21b. Dorsal fin rays fewer than 88; anal fin rays fewer 68; lower jaw not distinctly included in upper jaw when mouth is closed; without conspicuous canines overhanging lower jaw; caudal fin without large spots . . . . . . . . . . . . . . . . . 22 22a. Ocular side of body and fins densely covered with regularly arranged spots and blotches (if scales absent, body markings obscure) . . . . . . . . . . . . Citharichthys macrops p. 845 22b. Ocular side of body and fins not profusely covered with regularly arranged spots and blotches . . . . . . . . . . . . . . 23 23a. Eye diameter usually 30% of head length or greater; cephalic spines present in males but absent in females . . . . . . 24 23b. Eye diameter 25% of head length or less; no cephalic spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 24a. Snout partially covered with scales; ocular-side pelvic fin with six rays; lateral line scales 40 or more; axil of pectoral fin with small dark spot . . . . Citharichthys cornutus p. 841 24b. Snout naked; ocular-side pelvic fin with five rays; lateral line scales fewer than 40; axil of pectoral fin without dark spot. . . . . . . . . . . . . . . . Citharichthys gymnorhinus p. 843

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836 PLEURONECTIFORMES

25a. Dorsal fin rays about 68; anal fin rays about 52; lateral line scales 52 to 55 . . . . . . . . . . . . . Citharichthys uhleri p. 847 25b. Dorsal fin rays 68 to 84; anal fin rays 48 to 63; lateral line scales 42 to 50 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 26a. Body depth usually less than 45% of SL; interorbital space narrow, consists solely of narrow bony ridge; first gill arch with three to five gill rakers on upper limb; caudal vertebrae 23 to 25 . . . . . . . . . . Citharichthys spilopterus p. 846 26b. Body depth usually greater than 45% of SL; interorbital space relatively wide, not completely filled by bony ridge; first gill arch with five to eight gill rakers on upper limb; caudal vertebrae 21 to 23. . . . Citharichthys abbotti p. 839 27a. Large black spot on center of caudal fin; large black blotch on distal margin of ocular-side pectoral fin, but no black blotch on body under ocular-side pectoral fin; ocular-side pectoral fin rays 11 or 12. . . . Cyclopsetta fimbriata p. 849 27b. No large black spot on center of caudal fin; no black blotch on distal margin of ocular-side pectoral fin, but with large black blotch on body under ocular-side pectoral fin; ocular-side pectoral fin rays 14 to 16 . . . . . . . . . . . . . . . . . . . . . . Cyclopsetta chittendeni p. 848

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PARALICHTHYIDAE 837

Ancylopsetta dilecta Three-eye flounder

(Goode and Bean, 1883)

Ancylopsetta dilecta is moderately deep bodied, with several anterior dorsal fin rays elongated, pelvic fin on ocular side longer than fin on blind side, and three ocellated spots on ocular side. Lower eye is located directly below upper eye. Maxilla extends to center of pupil. Jaws and teeth are equally developed on each side. Jaw teeth are caninelike, larger anteriorly than posteriorly, and in a single row. Gill rakers on first gill arch are moderately long and slender and number one to three on upper limb and six to nine on lower limb. Measurements are expressed as percent of SL: head length 25%–31%, snout length 5%– 6%, eye diameter 6%– 8%, upper jaw length 11%–14%, ocular-side pectoral fin length 17%–20%, body depth 45%–53%. Pectoral fin on ocular side has 10 to 12 rays. Dorsal fin has 68 to 79 rays, with second through sixth or seventh elongated and second or third the longest. Anteriormost dorsal fin rays bear terminal fleshy filaments. Pelvic fin bases are short and insert on

either side of ventral midline. Anal fin has 53 to 60 rays. Caudal fin margin is obtusely angled. Scales on both sides of body are ctenoid. Lateral line is strongly arched over pectoral fin base. Lateral line scales number 73 to 82. Vertebrae number 35 to 37: 10 precaudal and 25 to 27 caudal. Ocular side is tan to pale brown, with numerous spots and blotches and three ocellated spots. Ocelli are in a triangular pattern, with anterior two symmetrically arranged on either side of the lateral line slightly posterior to midlength, and posterior ocellus located on caudal peduncle. Blind side is white. This species occurs between 59 and 366 m in the western Atlantic from North Carolina to Suriname, including the entire Gulf of Mexico. Maximum known size is 250 mm TL. references: Gutherz 1966, 1967; Hoese and Moore 1977, 1998; Matsuura 1983p; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Munroe 2002c.

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838 PLEURONECTIFORMES

Ancylopsetta quadrocellata Ocellated flounder

Gill, 1864

Ancylopsetta quadrocellata is relatively deep bodied, with several anterior dorsal fin rays slightly elongated, pelvic fin on ocular side slightly longer than one on blind side, and four ocellated spots on upper side. Lower eye is slightly anterior to upper eye. Maxilla extends slightly beyond center of pupil. Jaws and teeth are equally developed on each side. Jaw teeth are caninelike, larger anteriorly than posteriorly, and arranged in a single row. Gill rakers on first gill arch are short and blunt and number two or three on upper limb and six or seven on lower limb. Measurements are expressed as percent of SL: head length 25%–32%, snout length 4%– 6%, eye diameter 5%– 6%, upper jaw length 10%–12%, ocular-side pectoral fin length 16%–19%, body depth 53%– 63%. Pectoral fin on ocular side has 10 to 12 rays. Dorsal fin has 67 to 76 rays, with second through fifth to seventh slightly elongated. Pelvic fin bases are short and insert on either side of ventral midline. Anal fin has 54 to 61 rays. Caudal fin margin is obtusely angled. Scales on both sides of body are ctenoid. Lateral line is strongly arched over pectoral fin base. Lateral line scales number 80 to 90. Vertebrae number 35 to 37: 10 precaudal and 25 to 27 caudal (At-

lantic specimens have 11 precaudal vertebrae). Ocular side is brown, with four ocellated spots possessing white centers. Anteriormost ocellus is located over curved section of lateral line, and the other three are in a triangular pattern, with anterior two symmetrically arranged on either side of the lateral line slightly posterior to midlength, and posterior ocellus located on caudal peduncle. Blind side is uniformly white. This species occurs in the western Atlantic from North Carolina to Jupiter Inlet, FL, and in the Gulf of Mexico from the Dry Tortugas to Campeche, at depths less than 91 m. Food consists of crustaceans, including stomatopods, ostracods, mysids, amphipods, and brachyurans (Heterocrypta, Portunus, and Parthenops), and ray-finned fishes (Myrophis punctatus). Spawning occurs in late fall and winter in the Gulf of Mexico. Maximum known size is 250 mm TL.

references: Hildebrand 1955; Springer and Woodburn 1960; Gutherz 1966, 1967; Topp and Hoff 1972; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Munroe 2002c.

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PARALICHTHYIDAE 839

Citharichthys abbotti

Dawson, 1969

Citharichthys abbotti is moderately elongate and oval, with a smooth snout and smooth orbital rims, snout longer than eye, and ocular side densely speckled with small brown blotches. Upper eye is slightly anterior to lower eye. Lower jaw projects slightly beyond upper jaw, and maxilla extends to posterior section of lower eye. Interorbital distance is slightly more than half of eye diameter. Jaws are symmetrical, and jaw teeth are symmetrically arranged. Upper and lower jaws have an outer row of recurved fixed teeth and an inner one or two rows of depressible teeth. Teeth decrease in size posteriorly. Gill rakers on first arch are moderately long and slender and number 5 to 8 on upper limb and 13 to 16 on lower limb. Measurements are expressed as percent of SL: head length 27%–31%, ocular-side pectoral fin length 12%–15%, upper pelvic fin length 10%–13%, caudal peduncle depth 12%–14%, body depth 44%–51%; and as percent of head length: snout length 18%–21%, orbit diameter 17%–24%, upper jaw length 35%–39%. Pec-

toral fin on ocular side has 9 to 11 rays. Dorsal fin originates on blind side anterior to posterior naris and has 72 to 81 rays. Pelvic fin has 6 rays. Anal fin has 52 to 60 rays. Caudal fin margin is obtusely angled. Most scales on ocular side are ctenoid, and those on blind side are cycloid. Lateral line is slightly elevated above pectoral fin, and pored scales number 42 to 49. Generally there are two or three secondary scales between pored lateral line scales. Vertebrae number 31 to 33: 9 to 11 precaudal and 21 to 23 caudal. Ocular side is light sandy tan and densely speckled with small brown spots or blotches. Blind side is creamy white. This species occurs between the shoreline and 2 m in the western Atlantic in the southern Gulf of Mexico from Boca del Río, Veracruz, to Zacatal, Campeche. It is benthic on fine sandy to muddy bottoms. Maximum known size is 143 mm SL. references: Dawson 1969a; Schaldach et al. 1997; Castro-Aguirre et al. 1999; Munroe 2002c.

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840 PLEURONECTIFORMES

Citharichthys arctifrons Gulf stream flounder

Goode, 1880

Citharichthys arctifrons is moderately elongate and oval in profile, with a blunt, bony protuberance projecting forward in front of lower eye and eyes separated by a bony ridge. Lower eye is slightly anterior to upper eye. Maxilla extends to below middle of lower eye. Jaws and teeth are equally developed on each side. Jaw teeth are small and uniserial. Gill rakers on first arch are short and stout and number five on upper limb and six to eight on lower limb. Measurements are expressed as percent of SL: head length 20%–25%, snout length 3%– 4%, eye diameter 5%–7%, upper jaw length 6%– 8%, ocular-side pectoral fin length 14%–19%, body depth 34%– 43%. Pectoral fin on ocular side has 10 or 11 rays. Dorsal fin has 74 to 83 rays. Pelvic fin bases are short, with that of ocular-side fin on ventral midline and slightly posterior to that of blind-side fin. Ocular-side pelvic fin has 6 rays. Anal fin has 58 to 67 rays. Caudal fin is bluntly rounded. Body is covered

with cycloid or weakly ctenoid scales. Lateral line is developed on both sides and is not strongly curved over pectoral fin. Lateral line scales number 37 to 43. Precaudal vertebrae number 10 or 11, and caudal vertebrae number 26 to 28. Color of ocular side is tan to brown. Blind side is white. This species occurs between 46 and 366 m in the western Atlantic from Georges Bank off Massachusetts to Yucatán, including the Gulf of Mexico. It has occasionally been captured as shallow as 22 m. In the Gulf of Mexico it is known from the west coast of Florida and off Alabama and Veracruz. This species is associated with soft bottoms. Maximum known size is 180 mm TL. references: Norman 1934; Gutherz 1967; Robins and Ray 1986; Boschung 1992; Smith 1997; Castro-Aguirre et al. 1999; Klein-MacPhee 2002b; Munroe 2002c.

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PARALICHTHYIDAE 841

Citharichthys cornutus Horned whiff

(Günther, 1880)

Citharichthys cornutus is moderately deep bodied and oval in profile, with a relatively large mouth and a strong spine projecting from snout in males. Mature males have two small spines in front of upper eye, one or two small spines in front of lower eye, one small spine in front of nares above upper lip, and one spine projecting posteroventrally from symphysis of lower jaw. Eyes are close set in juveniles and adult females but are separated by distance equal to half the diameter of eye in adult males. Lower eye is slightly anterior to upper eye in females and juveniles and distinctly anterior to upper eye in mature males. Upper jaw extends to middle of lower eye. Jaw teeth are small and arranged in a single row but are slightly larger anteriorly than posteriorly. Gill rakers on first arch are moderately long and slender and number 3 to 5 on upper limb and 11 to 15 on lower limb. Measurements are expressed as percent of SL: head length 27%–28%, snout length 5%– 6%, eye diameter about 9%, upper jaw length 12%–13%, ocular-side pectoral fin length 18%–23%, body depth 46%–51%. Pectoral fin on ocular side has 11 or 12 rays, and dorsalmost pectoral ray of mature males is elongated

as a filament. Dorsal fin has 74 to 84 rays. Pelvic fin bases are short and that of ocular-side fin is on ventral midline and slightly posterior to that of blind-side fin. Ocular-side pelvic fin has 6 rays. Anal fin has 59 to 67 rays. Caudal fin is rounded. Ocular side of body is covered with weakly ctenoid scales, and blind side is covered with cycloid scales. Lateral line is developed on both sides and is not strongly curved over pectoral fin. Lateral line scales number 40 to 45. Precaudal vertebrae number 10, and caudal vertebrae number 25 or 26. Color of ocular side is tan to brown, with dark blotch on axil of pectoral fin. Blind side is white. This species occurs between 27 and 366 m in the western Atlantic from North Carolina and the Bahamas to Brazil, including the northern and southern Gulf of Mexico and the Greater Antilles. It is associated with soft bottoms. Maximum known size is 100 mm TL. references: Norman 1934; Gutherz 1967; Topp and Hoff 1972; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; CastroAguirre et al. 1999; Munroe 2002c.

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842 PLEURONECTIFORMES

Citharichthys dinoceros Goode and Bean, 1886

Citharichthys dinoceros is moderately elongate and oval in profile, with a rostral spine overhanging upper lip, a short spine in front of upper eye, and a relatively long pectoral fin. Lower eye is slightly anterior to upper eye. Eyes are close set and separated by a sharp interorbital ridge. Upper jaw extends beyond middle of lower eye. Jaw teeth are small and arranged in a single row but are larger anteriorly than posteriorly, with those in upper jaw overhanging those of lower jaw. Gill rakers on first arch are moderately long and slender and number 3 on upper limb and 7 to 10 on lower limb. Measurements are expressed as percent of SL: head length 27%–29%, snout length 6%–7%, eye diameter 7%– 8%, upper jaw length 12%– 14%, ocular-side pectoral fin length 30%– 36%, body depth 41%– 44%. Pectoral fin on ocular side has 9 or 10 rays. Dorsal fin has 90 to 95 rays. Pelvic fin bases are short, and that of ocular-side fin is on ventral midline and is

slightly posterior to that of blind-side fin. Ocular-side pelvic fin has 6 rays. Anal fin has 70 to 76 rays. Caudal fin is rounded. Body is covered with cycloid scales. Lateral line is developed on both sides and is not strongly curved over pectoral fin. Lateral line scales number 43 to 53. Precaudal vertebrae number 10 or 11, and caudal vertebrae number 26 to 29. Color of ocular side is pale to dark brown, usually with large dark spots on dorsal and anal fins. Caudal fin often has two vertically arranged dark spots. Blind side is white. This species occurs between 183 and 1,829 m in the western Atlantic from the east coast of Florida to Nicaragua, including the eastern Gulf of Mexico off Florida and Alabama, the Greater Antilles, and Barbados. Maximum known size is 120 mm SL. references: Norman 1934; Gutherz 1967; Robins and Ray 1986; Boschung 1992; Munroe 2002c.

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PARALICHTHYIDAE 843

Citharichthys gymnorhinus Anglefin whiff

Gutherz and Blackman, 1970

Citharichthys gymnorhinus is moderately elongate and oval in profile, with well-developed cephalic spines in males, ocular-side pelvic fin with 5 rays, and snout naked. Adult males have one spine projecting anteriorly from ethmoid bone of skull, two subequal spines on anterior margin of upper eye, two spines on anterior dorsal margin of lower eye, and several small spines on lachrymal bone below lower eye. Upper eye is directly above lower eye or lower eye is slightly anterior to upper eye, and eyes are separated by a narrow bony ridge. Maxilla extends to near middle of eye. Adult males have an anterolaterally directed spine on articular head of maxilla and a forwardly directed spine at symphysis of lower jaw. Jaws are symmetrical, and jaw teeth are small, symmetrically distributed in jaws, and arranged in a single row. Gill rakers on first arch are moderately long and slender and number 3 to 5 on upper limb and 9 to 11 on lower limb. Measurements are expressed as percent of SL: head length 25%–29%, snout length 5%–7%, orbit diameter 8%–11%, upper jaw length 10%–

13%, ocular-side pectoral fin length 22%– 32% in males and 22%–31% in females, body depth 42%–50%. Pectoral fin on ocular side has 8 to 11 (usually 9 or 10) rays, with the second ray greatly elongated in mature males. Dorsal fin has 70 to 76 rays, with about the twenty-sixth and several following rays elongated in mature males, giving fin margin an angled appearance. Anteriormost dorsal fin rays are also elongated in mature males. Anal fin has 56 to 61 rays, with about thirteenth and several following rays elongated in mature males, giving fin margin an angled appearance. Caudal fin margin is acutely rounded. Scales on ocular side are large, ctenoid, and deciduous, and scales on blind side are cycloid, large, and deciduous. Head anterior to posterior margin of eyes lacks scales. Lateral line is slightly arched over pectoral fin and has 33 to 37 pored scales. Ocular side is straw colored in preservative, with dark blotches symmetrically arranged on dorsal and ventral contours and along midline of body. Two dark blotches are located near caudal fin base. Males have large

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844 PLEURONECTIFORMES

dark blotches on extended sections of dorsal and anal fins. This species occurs between 35 and 200 m in the western Atlantic from the Florida Keys and the Bahamas to Guyana, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it is known from

Alabama and the west coast of Florida from Tampa Bay to the Dry Tortugas. Maximum known size is 60 mm SL. references: Gutherz and Blackman 1970; Topp and Hoff 1972; Robins and Ray 1986; Boschung 1992; Munroe 2002c.

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PARALICHTHYIDAE 845

Citharichthys macrops Spotted whiff

Dresel, 1885

Citharichthys macrops is relatively deep bodied and oval in profile, with a relatively large mouth, leaflike cirri on margin of lower operculum, and body and fins densely covered with regularly arranged dark spots and blotches. Eyes are close set and separated by a low, narrow concave ridge, and are on same vertical line. Upper jaw extends nearly to middle of lower eye. Jaw teeth are very small and arranged in a single row. Gill rakers on first arch are moderately long and slender and number 5 or 6 on upper limb and 13 to 16 on lower limb. Measurements are expressed as percent of SL: head length 24%–27%, snout length 4%– 6%, orbit diameter 6%–7%, upper jaw length 9%– 10%, ocular-side pectoral fin length 15%– 17%, body depth 50%–53%. Pectoral fin on ocular side has 9 to 12 rays. Dorsal fin has 79 to 82 rays, with anterior rays more or less free of membranes and first ray longer than other anterior rays. Pelvic fin bases are short, and that of ocular-side fin is on ventral midline and slightly posterior to that of blind-side fin.

Ocular-side pelvic fin rays number 6. Anal fin has 59 to 63 rays. Caudal fin is bluntly pointed. Body is covered with cycloid or weakly ctenoid scales. Lateral line is developed on both sides and is not strongly curved over pectoral fin. Lateral line scales number 37 to 44 (usually 37 to 40). Precaudal vertebrae number 10, and caudal vertebrae number 24 or 25. Color of ocular side is tan to dark brown, with numerous dark spots and blotches on body and fins. Blind side is white. This species occurs from the shoreline to 15 m, and occasionally to 91 m, in the western Atlantic from North Carolina to Honduras, including the northern and southern Gulf of Mexico. It is associated with hard sand bottoms. Maximum known size is 200 mm TL. references: Norman 1934; Hildebrand 1955; Springer and Woodburn 1960; Gutherz 1967; Topp and Hoff 1972; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; CastroAguirre et al. 1999; Munroe 2002c.

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846 PLEURONECTIFORMES

Citharichthys spilopterus Bay whiff

Günther, 1862

Citharichthys spilopterus is moderately elongate and oval in profile, with a relatively large mouth and a distinctly concave anterior profile. Eyes are close set and separated by a low, narrow concave ridge, with upper eye slightly anterior to lower eye. Upper jaw extends nearly to posterior region of lower eye. Jaw teeth are small to moderate in size, increase in size anteriorly, and are arranged in a single row. Gill rakers on first arch are moderately long and stout and number 4 or 5 on upper limb and 9 to 15 on lower limb. Measurements are expressed as percent of SL: head length 26%–28%, snout length 4%– 6%, eye diameter 3%– 4%, upper jaw length 9%–11%, ocular-side pectoral fin length 13%–16%, body depth 43%– 49%. Pectoral fin on ocular side has 9 or 10 rays. Dorsal fin has 75 to 84 rays. Pelvic fin bases are short, and that of ocularside fin is on ventral midline and slightly posterior to that of blind side. Ocular-side pelvic

fin has 6 rays. Anal fin has 56 to 63 rays. Caudal fin is rounded. Scales are weakly ctenoid on upper side and cycloid on lower side. Lateral line is developed on both sides and is not strongly curved over pectoral fin. Lateral line scales number 41 to 49. Precaudal vertebrae number 10, and caudal vertebrae number 23 to 25. Color of ocular side is brown, with or without darker spots. Blind side is white. This species occurs from the shoreline to 75 m in the western Atlantic from New Jersey to Brazil, including the entire Gulf of Mexico and the Greater Antilles. It is associated with muddy bottoms. Maximum known size is 200 mm TL. references: Gutherz 1967; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Castro-Aguirre et al. 1999; Munroe 2002c.

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PARALICHTHYIDAE 847

Citharichthys uhleri Jordan, 1889

Citharichthys uhleri is relatively deep bodied and oval in profile, with a relatively large mouth and white spots on ocular side. Eyes are close set; separated by a low, narrow concave ridge; and on same vertical line. Upper jaw extends nearly to middle of lower eye. Jaw teeth are very small and arranged in a single row. Gill rakers on first arch are moderately short and slender and number 5 on upper limb and 12 or 13 on lower limb. Measurements are expressed as percent of SL: head length 26%– 28%, snout length about 5%, orbit diameter 4%– 6%, upper jaw length 10%–11%, ocularside pectoral fin length 12%–15%, body depth 45%–50%. Pectoral fin on ocular side has 10 rays. Dorsal fin has 68 rays. Pelvic fin bases are short, and base of ocular-side fin is on ventral midline and slightly posterior to base of

blind-side fin. Ocular-side pelvic fin rays number 6. Anal fin has 52 rays. Caudal fin is convex. Body is covered with cycloid or weakly ctenoid scales. Lateral line is developed on both sides and is not strongly curved over pectoral fin. Lateral line scales number 52 to 55. Color of ocular side is dark brown with white spots. Fins are mottled. Blind side is white. This species occurs in the western Atlantic from the southern Gulf of Mexico to Costa Rica, including the Greater and Lesser Antilles. In the Gulf of Mexico it is known from Veracruz. This species may be a junior synonym of C. arenaceus Evermann and Marsh (Munroe 2002c). Maximum known size is 110 mm SL. references: Parr 1931; Norman 1934; Gutherz 1967; Castro-Aguirre et al. 1999; Munroe 2002c.

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848 PLEURONECTIFORMES

Cyclopsetta chittendeni Mexican flounder

Bean, 1895

Cyclopsetta chittendeni is moderately elongate and oval in profile, with lateral line only slightly arched anteriorly and large round spots on ocular side of dorsal, anal, and caudal fins. Eyes are close set and separated by a moderately narrow concave ridge, and upper eye is directly above lower eye. Mouth is large, with upper jaw extending to posterior margin of lower eye and lower jaw possessing angular process on ventral aspect of symphysis. Jaws are symmetrical, and teeth are relatively large and symmetrically arranged. Jaw teeth are slender, sharp pointed, larger in anterior section than in posterior section of jaws, and arranged in a single row. Gill rakers on first arch are short, broad based, and covered with teeth, and number three to five on upper limb and eight or nine on lower limb. Measurements are expressed as percent of SL: head length 26%– 29%, snout length 5%– 6%, orbit diameter 5%– 6%, upper jaw length 14%–15%, ocularside pectoral fin length 13%–15%, body depth 45%– 49%. Pectoral fin on ocular side has oblique distal margin and 14 to 16 rays. Dorsal fin has 82 to 90 rays, with first several rays slightly longer than more posterior rays and

partially free of membranes. Pelvic fin bases are short, and that of ocular-side fin inserts on midline and is slightly posterior to that of blind-side fin. Anal fin has 63 to 69 rays. Caudal fin margin is obtusely angled. Scales on both sides are cycloid and adherent. Lateral line scales number 74 to 80. Ocular side is brown, with a large black blotch under pectoral fin, three more or less evenly spaced dark spots on dorsal fin, two more or less evenly spaced dark spots on anal fin, and three more or less evenly spaced dark spots on distal margin of caudal fin. Blind side is white. This species occurs between 18 and 137 m in the western Atlantic from the northern Gulf of Mexico to Brazil. It has been captured in both the northern and southern Gulf of Mexico, but it is apparently absent off the Florida gulf coast and in the Antilles. Maximum known size is 320 mm TL. references: Gunter 1946 (as C. decussata); Hildebrand 1955; Gutherz 1967, Dawson 1968; Hoese and Moore 1977, 1998; Matsuura 1983p; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Munroe 2002c.

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PARALICHTHYIDAE 849

Cyclopsetta fimbriata Spotfin flounder

(Goode and Bean, 1885)

Cyclopsetta fimbriata is moderately elongate and oval in profile, with lateral line only slightly arched anteriorly and large round spots on ocular side of dorsal, anal, and caudal fins. Eyes are close set and separated by a moderately narrow concave ridge, and upper eye is directly above lower eye. Mouth is large, with upper jaw extending to posterior margin of lower eye and lower jaw possessing an angular process on ventral aspect of symphysis. Jaws are symmetrical, and teeth are symmetrically arranged. Jaw teeth are slender and sharp pointed, larger in anterior sections of jaws, and arranged in a single row. Gill rakers on first arch are short, broad based, and covered with teeth, and number 3 or 4 on upper limb and 9 or 10 on lower limb. Measurements are expressed as percent of SL: head length 28%–30%, snout length 5%– 6%, orbit diameter 6%–7%, upper jaw length 14%–15%, ocular-side pectoral fin length 15%–18%, body depth 45%– 47%. Pectoral fin on ocular side has a truncate distal margin and 11 or 12 rays. Dorsal fin has 78 to 87 rays, with anteriormost slightly longer than more posterior rays and partially free of membranes. Pelvic fin bases are short, and that of ocular-

side fin inserts on midline and slightly posterior to that of blind-side fin. Anal fin has 59 to 67 rays. Caudal fin margin is obtusely angled. Scales on both sides are cycloid and adherent. Lateral line is very slightly elevated over pectoral fin base. Lateral line scales number 65 to 75. Ocular side is brown. Distal section of pectoral fin is dark. Posterior half of dorsal fin has two more or less evenly spaced dark spots. Posterior two-thirds of anal fin has two more or less evenly spaced dark spots. A large spot is located on center of caudal fin, and three more or less evenly spaced dark spots occur on distal margin. Blind side is white. This species occurs between 118 and 228 m in the western Atlantic from North Carolina to Guyana, including the Gulf of Mexico and the Antilles. In the Gulf of Mexico it is most common east of Pensacola and is rare on Campeche Bank. Maximum known size is 330 mm TL. references: Longley and Hildebrand 1941; Hildebrand 1955; Gutherz 1967; Topp and Hoff 1972; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Munroe 2002c.

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850 PLEURONECTIFORMES

Etropus crossotus Fringed flounder

Jordan and Gilbert, 1882

Etropus crossotus is deep bodied, with a small mouth, a low interorbital ridge, and no secondary squamation. Anterior profile of head is straight in juveniles and concave in adults. Eyes are close set and separated by a slightly elevated interorbital ridge, and lower eye is slightly anterior to upper eye. Maxilla extends to anterior section of lower eye. Mandible is asymmetrical and has symphyseal knob. Jaw teeth are small, conical, blunt, and arranged in a single row, and are larger on blind side than on ocular side. Gill rakers are short and stout and number three to six on upper limb and six to nine on lower limb. Measurements are expressed as percent of SL: head length 20%– 25%, snout length 3%– 4%, eye diameter 4%– 5%, upper jaw length 4%– 6%, ocular-side pectoral fin length 14%–16%, caudal peduncle depth 12%–14%, body depth 53%–59%. Pectoral fin on ocular side has 8 to 10 rays. Dorsal fin has 73 to 84 rays. Pelvic fin bases are short, and base of ocular-side fin is on ventral midline and slightly posterior to base of blind-side fin. Anal fin has 57 to 67 rays. Caudal fin is rounded. Scales are weakly ctenoid on ocular

side and cycloid on blind side. Scales are lacking on snout. Lateral line is developed on both sides and is not arched over pectoral fin. Lateral line scales number 38 to 45. Precaudal vertebrae number 10, and caudal vertebrae number 24 to 26. Color of ocular side is uniformly dark olive brown, often with dark margin on caudal fin. Specimens from Campeche Bank are whitish on ocular side (Hildebrand 1955). Blind side is white. This species occurs in the western Atlantic and eastern Pacific. In the western Atlantic it occurs between the shoreline and 30 m, and occasionally to 91 m, from Virginia to possibly northern South America, including the northern and southern Gulf of Mexico and the Antilles. Maximum known size is 200 mm TL.

references: Norman 1934 (as Etropus intermedius, in part); Hildebrand 1955; Springer and Woodburn 1960; Gutherz 1967; Hoese and Moore 1977, 1998; Robins and Ray 1986; Leslie and Stewart 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Munroe 2002c.

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PARALICHTHYIDAE 851

Etropus cyclosquamus Shelf flounder

Leslie and Stewart, 1986

Etropus cyclosquamus is deep bodied and elliptical in profile, with a small mouth, a low interorbital ridge, and secondary squamation covering most scales on ocular side. Eyes are close together and separated by a slightly elevated interorbital ridge, and lower eye is usually directly below upper eye. Mandible is asymmetrical and has symphyseal knob. Jaw teeth are small, conical, blunt, and arranged in a single row, and are larger on blind side than on ocular side. Gill rakers are short and stout and number four to eight on upper limb and three to six on lower limb. Measurements are expressed as percent of SL: head length 22%– 26%, snout length 4%–5%, orbit diameter 7%– 8%, upper jaw length 6%–7%, ocularside pectoral fin length 16%–24%, caudal peduncle depth 12%–16%, body depth 47%– 61%. Pectoral fin on ocular side has 8 to 11 (usually 9 or 10) rays. Dorsal fin has 70 to 82 (usually 73 to 80) rays. Pelvic fin bases are short, and that of ocular-side fin is on ventral midline and slightly posterior to that of blindside fin. Anal fin has 54 to 64 (usually 56 to 62) rays. Caudal fin is rounded. Scales are weakly ctenoid on ocular side and cycloid on blind side. Primary scales on ocular side are covered

with secondary scales except for narrow posterior margin. Scales are lacking on snout anterior to line connecting ocular- and blind-side nares (large specimens may have several scales anterior to nares). Ctenii on snout scales are unmodified. Lateral line is developed on both sides and is not arched over pectoral fin. Lateral line scales number 36 to 45 (usually 38 to 42). Precaudal vertebrae number 10, and caudal vertebrae number 23 to 25. Ocular side is dusky gray to dusky brown, with dark spots and blotches. Some specimens have a row of four to six dark rings in place of blotches above and below lateral line. Caudal fin lacks dark margin. Blind side is white. This species occurs between 18 and 36 m in the western Atlantic from just north of Cape Hatteras to the eastern Gulf of Mexico. In the Gulf of Mexico it has been captured between Fort Myers, FL, and Mississippi Sound. Maximum known size is 104 mm SL. references: Gutherz 1967; Robins and Ray 1986 (both previous references as E. microstomus, in part); Leslie and Stewart 1986; Retzer 1990; Boschung 1992; Hoese and Moore 1998; Munroe 2002c.

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852 PLEURONECTIFORMES

Etropus rimosus Gray flounder

Goode and Bean, 1885

Etropus rimosus is deep bodied and elliptical in profile, with a small mouth, a low interorbital ridge, and secondary squamation covering most of scales on ocular side. Eyes are close together and separated by a strongly elevated interorbital ridge, and lower eye is slightly anterior to upper eye. Maxilla extends to anterior section of lower eye. Mandible is asymmetrical and has symphyseal knob. Jaw teeth are small, conical, blunt, and arranged in a single row, and are larger on blind side than on ocular side. Gill rakers are short and stout and number five to eight on upper limb and four to six on lower limb. Measurements are expressed as percent of SL: head length 22%–26%, snout length about 3%, eye diameter about 5%, upper jaw length about 5%, ocular-side pectoral fin length 12%–24%, caudal peduncle depth 13%–16%, body depth 51%–59%. Pectoral fin on ocular side has 10 or 11 rays. Dorsal fin originates on blind side at level of anterior margin of upper eye and has 70 to 83 (usually 75 to 80) rays. Pelvic fin bases are short, and that of ocular-side fin is on ventral midline and slightly posterior to that of blind-side fin. Anal fin has 53 to 64 (usually 59 to 62) rays. Cau-

dal fin is rounded. Scales are weakly ctenoid; however, ctenii may not be visible on specimens less than 50 mm SL. Primary scales on ocular side are covered with secondary scales except for narrow posterior margin. Scales occur on snout anterior to line connecting ocularand blind-side nostrils. Ctenii on snout scales are highly modified. Lateral line is developed on both sides and is not arched over pectoral fin. Lateral line scales number 36 to 45 (usually 38 to 42). Precaudal vertebrae number 10, and caudal vertebrae number 24 or 25. Ocular side is dusky gray. Caudal fin lacks dark margin. Blind side is white. This species occurs between 7 and 60 m in the western Atlantic from just north of Cape Hatteras to the eastern Gulf of Mexico. In the Gulf of Mexico it has been captured from the Dry Tortugas to the Florida Big Bend (2901N). Maximum known size is 111 mm SL. references: Norman 1934; Gutherz 1967 (possibly as E. cyclosquamus); Robins and Ray 1986 (possibly as E. cyclosquamus) ; Retzer 1990; Boschung 1992; Munroe 2002c.

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PARALICHTHYIDAE 853

Gastropsetta frontalis Shrimp flounder

Bean, 1895

Gastropsetta frontalis is moderately deep bodied, with several anterior dorsal fin rays elongated, ocular-side pelvic fin longer than blindside pelvic fin, and three ocellated spots on ocular side. Margin of head is nearly vertical. Lower eye is located directly below upper eye. Maxilla extends to anterior margin of pupil of lower eye. Jaws and teeth are equally developed on each side. Jaw teeth are moderate in size, caninelike, and arranged in a single row. Gill rakers on first gill arch are short and thick and number three to six on upper limb and six to eight on lower limb. Measurements are expressed as percent of SL: head length 23%– 30%, snout length 4%–5%, eye diameter 5%– 6%, upper jaw length 8%–9%, ocular-side pectoral fin length 16%–17%, body depth 45%–55%. Pectoral fin on ocular side has 10 or 11 rays. Dorsal fin has 58 to 65 rays with first through fifth or sixth elongated and third or fourth the longest. Anteriormost dorsal fin rays bear terminal fleshy filaments. Bases of pelvic fins are short and insert on either side of ventral midline. Anal fin has 46 to 53 rays. Caudal fin margin is rounded. Scales are small

and cycloid. Lateral line is strongly arched over pectoral fin base. Lateral line scales number 95 to 112. Vertebrae number 35 to 38: 10 precaudal and 25 to 28 caudal. Ocular side is tan to dark brown, with three ocellated spots. Ocelli are in a triangular pattern, with anterior one over arched section of lateral line and posterior two symmetrically arranged on either side of lateral line slightly posterior to midlength. Blind side is white. This species occurs between 35 and 183 m in the western Atlantic from North Carolina to Panama, including the Gulf of Mexico and the Bahamas. It occurs throughout the Gulf of Mexico but is more abundant in the eastern than in the western Gulf. Food consists of crustaceans and synodontid fishes. Maximum known size is 250 mm TL. references: Longley and Hildebrand 1941; Springer and Bullis 1956; Bullis and Thompson 1965; Gutherz 1966, 1967; Topp and Hoff 1972; Robins and Ray 1986; Boschung 1992; Munroe 2002c.

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854 PLEURONECTIFORMES

Hippoglossina oblonga Fourspot flounder

(Mitchill, 1815)

Hippoglossina oblonga is relatively elongate and oval in profile, with large eyes, lateral line strongly arched over pectoral fin base, and four ocellated spots on ocular side of body. Eyes are close set and separated by a narrow interorbital ridge, and lower eye is directly below upper eye. Maxilla extends to posterior margin of pupil of lower eye. Mandible is symmetrical, and jaw teeth are symmetrically arranged in jaws. Teeth are caninelike, larger anteriorly than posteriorly, and arranged in a single row. Gill rakers on first arch are moderately long and moderately stout and number 3 to 6 on upper limb and 7 to 10 on lower limb. Measurements are expressed as percent of SL: head length 24%–27%, snout length 5%– 6%, orbit diameter 5%–7%, upper jaw length 11%– 12%, ocular-side pectoral fin length 13%– 15%, body depth 38%– 44%. Pectoral fin on ocular side has 10 to 12 rays. Dorsal fin has 71 to 86 rays, with anteriormost ray elongated and only basally connected to fin membrane.

Pelvic fin bases are symmetrical. Anal fin has 58 to 72 rays. Lateral line scales number 63 to 95. Scales are cycloid and deciduous on both sides. Vertebrae number 41 or 42: 11 precaudal and 30 to 31 caudal. Ocular side is brown, with four ocelli arranged in a trapezoid pattern. Anterior pair of ocelli is located on either side of lateral line just posterior to midlength, and posterior pair is located on either side of lateral line just anterior to caudal peduncle. Blind side is white. This species occurs in the western Atlantic from Georges Bank to the Dry Tortugas. In the Gulf of Mexico it is known only from the Dry Tortugas. It occurs from 27 to about 400 m in the northern part of its range, but it occurs as deep as 274 m in the southern part of its range. Maximum known size is about 450 mm TL. references: Ginsburg 1952b; Gutherz 1967; Robins and Ray 1986; Murdy et al. 1997; KleinMacPhee 2002b; Munroe 2002c.

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PARALICHTHYIDAE 855

Paralichthys albigutta Gulf flounder

Jordan and Gilbert, 1882

Paralichthys albigutta is moderately elongate and oval in profile, with lateral line strongly arched over pectoral fin base, pelvic fin bases symmetrical and of equal length, and three ocelli arranged in a triangular pattern on ocular side. Eyes are moderately separated, with lower eye directly below upper eye. Maxilla extends beyond level of lower eye. Jaws are symmetrical, and jaw teeth are symmetrically arranged. Jaw teeth are caninelike, larger anteriorly than posteriorly, and arranged in a single row. Some jaw teeth are fixed, but others are depressible. Gill rakers on first arch are moderately long and moderately stout and number 4 to 7 on upper limb and 9 to 12 on lower limb. Measurements are expressed as percent of SL: head length 27%–30%, snout length 6%–7%, orbit diameter 4%– 6%, upper jaw length 14%–16%, ocular-side pectoral fin length 14%–15%, body depth 39%– 42%. Pectoral fin on ocular side has 10 to 12 rays. Dorsal fin has 71 to 85 rays. Anal fin has 53 to 63 rays. Scales are cycloid and deciduous. Lateral line has 78 to 117 scales. Vertebrae num-

ber 37: 10 precaudal and 27 caudal. Ocular side is light to dark brown, with numerous spots and blotches and three relatively small ocelli arranged in a triangular pattern: two ocelli just behind arched section of lateral line and a third ocellus on posterior section of lateral line. This species occurs from 19 to 128 m in the western Atlantic from North Carolina to Panama, including the entire Gulf of Mexico and the Bahamas. It is generally found on hard or sandy bottoms. In the northern Gulf of Mexico spawning takes place between November and February. Food consists of penaeid shrimps, portunid crabs, and ray-finned fishes such as Anchoa sp., Fundulus similis, Syngnathus sp., Haemulon sp., and Gobiosoma robustum. Maximum known size is 710 mm TL. references: Ginsburg 1952b; Gutherz 1967; Böhlke and Chaplin 1968; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Castro-Aguirre et al. 1999; Munroe 2002c.

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856 PLEURONECTIFORMES

Paralichthys lethostigma Southern flounder

Jordan and Gilbert, 1884

Paralichthys lethostigma is relatively elongate and oval in profile, with lateral line strongly arched over pectoral fin base, pelvic fin bases symmetrical and of equal length, and ocular side plain colored or with spots or blotches. Eyes are distinctly separated, with lower eye directly below upper eye. Maxilla extends beyond level of lower eye. Jaws are symmetrical. Jaw teeth are caninelike, equally developed on each side, larger anteriorly than posteriorly, and arranged in a single row. Some jaw teeth are fixed, but others are depressible. Gill rakers on first arch are moderately long and moderately stout and number 2 or 3 on upper limb and 8 to 11 on lower limb. Measurements are expressed as percent of SL: head length 24%– 30%, snout length 5%–7%, orbit diameter 3%–5%, upper jaw length 12%–15%, ocularside pectoral fin length 12%–15%, body depth 41%– 47%. Pectoral fin on ocular side has 11 to 13 rays. Dorsal fin has 80 to 95 rays. Anal fin has 63 to 74 rays. Scales are cycloid and deciduous. Lateral line has 85 to 100 scales. Ver-

tebrae number 37 to 39: 10 or 11 precaudal and 27 or 28 caudal. Ocular side is light to dark brown, with or without diffuse spots and blotches. Larger specimens may lack spots and blotches. Blind side is white. This species occurs between the shoreline and 40 m in the western Atlantic from North Carolina to the Gulf of Mexico. It occurs throughout the Gulf of Mexico except for the southwestern coast of Florida. This species is associated with muddy bottoms along the coast, estuaries and bays, and freshwater with access to the sea. Food consists of shrimps, crabs, and ray-finned fishes. Juveniles feed on soft-bodied invertebrates. Maximum known size is 750 mm TL. references: Ginsburg 1952b; Springer and Woodburn 1960; Gutherz 1967; Topp and Hoff 1972; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Castro-Aguirre et al. 1999; Munroe 2002c.

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PARALICHTHYIDAE 857

Paralichthys squamilentus Broad flounder

Jordan and Gilbert, 1882

Paralichthys squamilentus is relatively deep bodied and oval in profile, with lateral line strongly arched over pectoral fin base, pelvic fin bases of equal length and symmetrical, and ocular side plain colored or spotted. Eyes are distinctly separated, with lower eye directly below upper eye. Maxilla extends beyond level of lower eye. Jaws are symmetrical, and jaw teeth are symmetrically arranged. Jaw teeth are caninelike, larger anteriorly than posteriorly, and arranged in a single row. Some jaw teeth are fixed, but others are depressible. Gill rakers on first arch are moderately long and moderately stout and number 3 to 5 on upper limb and 9 to 12 on lower limb. Measurements are expressed as percent of SL: head length 26%– 27%, snout length 5%– 6%, orbit diameter 5%– 6%, upper jaw length 12%–13%, ocularside pectoral fin length 15%–17%, body depth

50%–54%. Pectoral fin on ocular side has 11 to 13 rays. Dorsal fin has 76 to 85 rays. Anal fin has 59 to 65 rays. Scales are cycloid and deciduous. Lateral line has 104 to 117 scales. Vertebrae number 37 to 39: 10 precaudal and 27 to 29 caudal. Ocular side is brown, with or without diffuse spots and blotches. Blind side is white in small specimens and dusky in large specimens. This species occurs from near shore to 229 m in the western Atlantic from North Carolina to Florida and throughout the Gulf of Mexico. Small juveniles occur in shallow water, and large juveniles and adults occur in deep water. Maximum known size is 460 mm TL. references: Ginsburg 1952b; Gutherz 1967; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Munroe 2002c.

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858 PLEURONECTIFORMES

Syacium gunteri Shoal flounder

Ginsburg, 1933

Syacium gunteri is relatively deep bodied and oval in profile, with a comparatively broad interorbital space and a relatively straight lateral line. Lower eye is distinctly anterior to upper eye. Interorbital width varies between sexes and with size within sexes. In males between 80 and 90 mm SL, interorbital width is 25% to 40% of lower eye diameter; between 91 and 100 mm SL, 25% to 40%; and between 101 mm and maximum SL, 35% to 55%. In females between 80 and 90 mm SL, interorbital width is 20% to 35% of lower eye diameter, and for those between 91 and 98 mm SL, it is 25% to 40%. Maxilla extends to below middle of lower eye. Jaws and teeth are equally developed on each side. Jaw teeth are conical and curved, biserial in upper jaw, and uniserial in lower jaw. Gill rakers on first arch are relatively long and thick and number two to four on upper limb and six to eight on lower limb. Measurements are expressed as percent of SL: head length 25%–29%, snout length 5%– 6%, orbit diameter 5%– 6%, upper jaw length 10%–11%, ocular-side pectoral fin length 27%–37% for males and 18%–23% for females, body depth 50%–56%. Pectoral fin on

ocular side has 9 to 11 rays. Dorsal fin has 74 to 85 rays. Pelvic fin bases are short and asymmetrical. Anal fin has 59 to 68 rays. Caudal fin is bluntly rounded. Body is covered with ctenoid scales on ocular side and cycloid scales on blind side. Lateral line scales number 46 to 55. Vertebrae number 33 to 36: 10 precaudal and 23 to 26 caudal. Ocular side is tan, with or without darker spots and blotches on body and median fins. Blind side is white. This species occurs between 17 and 126 m in the western Atlantic from the northern and southern Gulf of Mexico and the Antilles. It is more abundant in the northwestern than in the northeastern Gulf of Mexico. It is generally found on sandy and mud bottoms, and it is relatively less abundant on calcareous bottoms. Food consists of annelids, crustaceans, and ray-finned fishes. Maximum known size is 200 mm TL. references: Hildebrand 1955; Gutherz 1967; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Munroe 2002c.

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PARALICHTHYIDAE 859

Syacium micrurum Ranzani, 1842 Channel flounder

Syacium micrurum is moderately elongate and oval in profile, with a relatively straight lateral line and a relatively narrow interorbital space. Lower eye is distinctly anterior to upper eye. Interorbital distance varies slightly between sexes and moderately with size within sexes. Interorbital width is less than 15% of lower eye diameter in males and females less than 120 mm SL, less than 35% in males and about 20% in females between 120 and 150 mm SL, less than 50% in males and about 27% in females between 150 and 180 mm SL, and less than 75% in males greater than 180 mm SL. Maxilla extends to below midsection of lower eye. Jaws and teeth are equally developed on each side. Jaw teeth are conical and curved, biserial in upper jaw, and uniserial in lower jaw. Gill rakers on first arch are short and stout and number two on upper limb and seven to nine on lower limb. Measurements are expressed as percent of SL: head length 26%– 28%, snout length 5%– 6%, eye diameter 5%– 7%, upper jaw length 10%–11%, ocular-side

pectoral fin length about 46% for males, body depth 40%– 43%. Pectoral fin on ocular side has 10 or 11 rays. Dorsal fin has 83 to 92 rays. Anal fin has 64 to 74 rays. Caudal fin is bluntly rounded. Body is covered with ctenoid scales on ocular side and cycloid scales on blind side. Lateral line scales number 54 to 68. Precaudal vertebrae number 10, and caudal vertebrae number 24 or 25. Ocular side is tan to brown, with or without darker spots and blotches on body and median fins. Blind side is white. This species occurs in the western Atlantic from the east coast of Florida to Rio de Janeiro, Brazil, including the Bahamas, the Greater Antilles, and possibly the northern Gulf of Mexico. Records from the Gulf of Mexico have not been confirmed. It has been captured to a depth of 412 m but is generally limited to 91 m. Maximum known size is 300 mm TL. references: Norman 1934; Gutherz 1967; Böhlke and Chaplin 1968; Robins and Ray 1986; Boschung 1992; Munroe 2002c.

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860 PLEURONECTIFORMES

Syacium papillosum Dusky flounder

(Linnaeus, 1758)

Syacium papillosum is moderately elongate and oval in profile, with a comparatively broad interorbital space and a relatively straight lateral line. Lower eye is distinctly anterior to upper eye. Interorbital width varies between sexes and with size within sexes. In males between 80 and 90 mm SL, interorbital width is 16% to 29% of lower eye diameter; between 91 and 100 mm SL, 30% to 33%; and between 101 mm and maximum SL, 45% to 102%. In females between 80 and 90 mm SL, interorbital width is 11% to 16% of lower eye diameter; between 91 and 98 mm SL, 25% to 29%; and between 101 mm and maximum SL, 41% to 42%. Maxilla extends to below midsection of lower eye. Jaws and teeth are equally developed on each side. Jaw teeth are conical and curved, biserial in upper jaw, and uniserial in lower jaw. Gill rakers on first arch are short and stout and number two on upper limb and eight or nine on lower limb. Measurements are expressed as percent of SL: head length 26%– 28%, snout length 5%– 6%, orbit diameter 5%–7%, ocular-side pectoral fin length 22%– 62% for males and about 24% for females, body depth 40%– 44%. Pectoral fin on ocular side has 11 or 12 rays. Dorsal fin has 82

to 94 rays. Pelvic fin bases are short, and that of ocular-side fin is on ventral midline and slightly posterior to that of blind-side fin. Anal fin has 64 to 75 rays. Caudal fin is bluntly rounded. Body is covered with ctenoid scales on ocular side and cycloid scales on blind side. Lateral line scales number 47 to 60. Vertebrae number 35 or 36: 10 precaudal and 25 or 26 caudal. Ocular side is brown, with few or no darker spots and blotches and three or four dark bands on pectoral fin. Blind side is dusky in males and white to slightly dusky in females. This species occurs between 9 and 137 m in the western Atlantic from North Carolina to Brazil, including the entire Gulf of Mexico and the Antilles. It is more abundant in the northeastern than in the northwestern Gulf of Mexico. This species is associated with both calcareous and muddy bottoms but is relatively more common on calcareous bottoms. Maximum known size is 250 mm TL. references: Longley and Hildebrand 1941; Hildebrand 1955; Gutherz 1967; Hoese and Moore 1977, 1998; Matsuura 1983p; Robins and Ray 1986; Boschung 1992; Cervigón 1993a; Schaldach et al. 1997; Smith 1997; Munroe 2002c.

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861

POECILOPSETTIDAE

Poecilopsettids are moderately deep bodied, with both eyes on the right side of the head, dorsal fin originating on neurocranium, and lateral line obsolete on blind side. Head profile is convex to deeply concave. Eyes are on same vertical line or lower eye is slightly anterior to upper eye, and upper eye is close to dorsal margin of head. Interorbital space is narrow to moderately broad. Nostrils are paired and located on each side of head. Mouth is small to very large and symmetrical or asymmetrical. Jaw teeth are variable in size and shape and are equally developed on each side or better developed in blind-side jaws. Palatine bone lacks teeth. Supramaxilla is absent. Preoperculum is free of skin and scales. Gill membranes are connected. Both pectoral fins are present, and fin rays are branched or unbranched. Dorsal fin originates on blind side of head at level of eyes or more anteriorly. Pelvic fins are symmetrical or asymmetrical. Pelvic fin has six unbranched rays. Dorsal and anal fins are separate from caudal fin. Lateral line has a high arch over ocular-side pectoral fin and is rudimentary or absent on blind side. Poecilopsettids occur in the western Atlantic, western and central Pacific, and Indian Oceans in tropical and warm temperate seas. They primarily occur in deep water. There are 14 species in 3 genera, and 2 species occur in the Gulf of Mexico. references: Norman 1934; Nelson 1994; Munroe 2002d.

Key to the Species of the Gulf of Mexico (Adapted from Munroe 2002d) 1a. Body depth 32% to 39% of SL; pored lateral line scales more than 80 . . . . . . . . . . . . . . Poecilopsetta beanii p. 862 1b. Body depth 40% to 46% of SL; pored lateral line scales fewer than 79 . . . . . . . . . . . . . Poecilopsetta inermis p. 863

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862 PLEURONECTIFORMES

Poecilopsetta beanii

(Goode, 1881)

Poecilopsetta beanii is elongate and oval in profile, with a relatively short head. Eyes are separated by a very narrow ridge, upper eye extends above margin of head, and lower eye is slightly anterior to upper eye. Mouth is small and rather symmetrical. Maxilla extends to anterior section of lower eye. Jaw teeth are villiform and arranged in two rows. Gill rakers on first arch are moderately short and stout and number 6 on upper limb and 10 or 11 on lower limb. Measurements are expressed as percent of SL: head length 19%–22%, snout length about 3%, eye diameter 7%–9%, upper jaw length 6%– 8%, ocular-side pectoral fin length 9%–10%, body depth 32%–39%. Pectoral fin on ocular side has 10 simple rays. Dorsal fin has 63 to 68 rays. Anal fin has 50 to 56 rays.

Caudal fin is bluntly pointed. Scales are ctenoid on ocular side and cycloid on blind side. Lateral line on ocular side extends onto caudal fin and consists of 80 to 89 scales. Ocular side is brownish, with dark markings that are most pronounced along upper and lower sections of body. Black blotches are located on middle of upper and lower margins of caudal fin. This species occurs between 155 and 1,639 m in the western Atlantic from New England to northern Brazil, including the Gulf of Mexico and the Antilles. Maximum known size is 90 mm SL. references: Breder 1927 (as P. inermis, in part); Norman 1934; Reid 1934 (as P. albomarginata?); Munroe 2002d.

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POECILOPSETTIDAE 863

Poecilopsetta inermis

(Breder, 1927)

Poecilopsetta inermis is elongate and oval in profile, with a relatively large head. Eyes are separated by a very narrow ridge, upper eye extends above margin of head, and lower eye is slightly anterior to upper eye. Mouth is small and rather symmetrical. Maxilla extends to pupil of lower eye. Jaw teeth are small and arranged in two rows, with teeth in blind-side jaws larger than those in ocular-side jaws. Gill rakers on first arch are moderately short and stout and number 4 on upper limb and 10 or 11 on lower limb. Gill opening extends above pectoral fin base. Measurements are expressed as percent of SL: head length 22%–28%, snout length 3%– 4%, eye diameter 7%–9%, upper jaw length 6%–7%, ocular-side pectoral fin length 9%–12%, body depth 40%– 46%. Pectoral fin on ocular side has 9 or 10 simple rays. Dorsal fin has 59 to 64 (usually 63 to 65) rays. Ocular-side pelvic fin inserts near ventral midline, slightly anterior to blind-side fin, and is

longer than blind-side fin. Anal fin has 51 to 53 (usually 53 to 55) rays. Caudal fin is bluntly pointed. Scales are ctenoid on ocular side and cycloid on blind side. Lateral line on ocular side extends onto caudal fin and has 71 to 76 scales. Lateral line on blind side is absent. Ocular side is pale yellowish gray, with margins of scale pockets dark. Pectoral fin on ocular side is dark, and dorsal, anal, and ocular-side pelvic fins are blackish with narrow pale bases and white margins. Black blotches are located on middle of dorsal and ventral margins of caudal fin. This species occurs between 183 and 549 m in the western Atlantic from the eastern Gulf of Mexico and Tobago Island in the Lesser Antilles. Maximum known size is 116 mm SL.

references: Breder 1927; Reid 1934 (as P. albomaculata, in part); Hoshino 2000; Munroe 2002d.

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864

ACHIRIDAE

American soles

Achirids are oblong to ovate in profile and compressed, with both eyes on right side of head, snout hooked, and dorsal and anal fins free of caudal fin. Profile of head is moderately to broadly rounded. Nostrils occur on each side of head and are paired and asymmetrically located, with anterior naris on ocular side ending as a tube and posterior naris opening below lower eye. Eyes are very small, close set, and separated by a bony ridge. Mouth is terminal to subterminal and twisted toward blind side and has fleshy lips that often bear dermal flaps. Jaw teeth are villiform and arranged in bands. Preopercular margin is adinate (concealed beneath scales and skin). Skin of lower jaw and interoperculum is continuous ventrally and covers isthmus and branchiostegal rays. Gill membranes are connected and joined to isthmus, and gill openings are either confluent or separated ventrally. Pectoral fins are minute or absent. Dorsal fin originates near or at tip of snout, with anterior rays concealed in a fleshy dermal envelope. Pelvic fins are developed, or one or both are absent. When present, pelvic fin on ocular side has a long base, is inserted on midline, and is connected to anal fin. Caudal fin rays are unbranched. Lateral line is present on ocular side and is straight. Body, head, and often dorsal and anal fins are either covered with ctenoid scales or are naked. Blind side of head is generally covered with fringes. Maximum known size is about 350 mm TL. Achirids occur in marine and fresh waters along North, Central, and South America. They are benthic usually on inner continental and insular shelves, but some reach depths up to 300 m. There are 28 species in nine genera, 12 species in five genera in the western Central Atlantic, and 5 or 6 species in three genera in the Gulf of Mexico. references: Nelson 1994; Munroe 2002e.

Key to the Species of the Gulf of Mexico (Adapted in part from Dawson 1964; Hoese and Moore 1998) 1a. Body covered with scales; upper eye anterior to lower eye; posterior margin of pelvic fin on blind side not connected to body by membrane . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Body naked; upper eye on vertical line with lower eye; margin of both pelvic fins connected to body by membranes .............................................4 2a. Interbranchial foramen present . . . Achirus lineatus p. 866 2b. Interbranchial foramen absent . . . . . . . . . . . . . . . . . . . . . 3 3a. Pectoral fins present; tufts of cirri absent on body; body pale and patterned with network of dark lines . . . . . . . . . . . . . . . . . . . . . . . . . Trinectes inscriptus p. 870

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ACHIRIDAE 865

3b. Pectoral fins absent; tufts of cirri present on body; body pale with dark bars. . . . . . . . . . Trinectes maculatus p. 871 4a. Number of complete and incomplete stripes on body 25 to 49 (generally 30 or more); number of pores in third accessory lateral line 21 to 48 (generally 27 to 38) . . . . . . . . . . . . . . . . . . . . . . . . . Gymnachirus texae p. 869 4b. Number of complete and incomplete stripes on body 11 to 32 (generally fewer than 31); number of pores in third accessory lateral line 26 to 79 (generally 34 to 62) . . . . . . 5 5a. Number of pores in third accessory lateral line 35 to 79 (usually 47 to 62) . . . . . . . . . . . Gymnachirus melas p. 867 5b. Number of pores in third accessory lateral line 26 to 59 (usually 34 to 44). . . . . . . . . . . Gymnachirus nudus p. 868

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866 PLEURONECTIFORMES

Achirus lineatus Lined sole

(Linnaeus, 1758)

Achirus lineatus is nearly circular in profile, with a matting of dermal folds and fimbriae on blind side of head, an interbranchial foramen, and pectoral fin on ocular side. Mouth is subterminal, and snout slightly overhangs jaws. Jaw teeth are villiform and present in blind side of jaws only. Opercular opening is greatly restricted but confluent ventrally. Measurements are expressed as percent of SL: head length 28.4%–31.1%, interorbital width 0.9%–2.2%, caudal fin length 28.7%–35.7%, body depth 65.2%–70%. Small rudimentary pectoral fin is located above gill slit on ocular side and consists of 3 to 8 rays. Dorsal fin has 49 to 60 rays, and anal fin has 38 to 46 rays. Ocular-side pelvic fin has 5 rays and is continuous with anal fin. Blind-side pelvic fin inserts anterior to ocular-side pelvic fin and has 5 rays. Entire body and most of head are covered with small ctenoid scales, including both sides of dorsal and anal fin rays. Fimbriae run

along margins of mouth on ocular side and are broadly distributed on blind side of head. Tufts of dark-colored cirri are randomly distributed on both sides of body. Ocular side is olive to brown, with darker spots and blotches and very narrow stripes evenly arranged along body. Blind side is white anteriorly, grading to light brown posteriorly. This species occurs in coastal waters in the western Atlantic from the east coast of South Carolina to Argentina, including the Gulf of Mexico and the Antilles. It is found in estuaries but generally at salinities greater than 15‰. Food consists of polychaetes and gammarid amphipods. Maximum known size is 230 mm TL. references: Jordan and Evermann 1898; Hildebrand 1955; Springer and Woodburn 1960; Hoese and Moore 1977, 1998; Robins and Ray 1986; Cervigón 1993a; Munroe 2002e.

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ACHIRIDAE 867

Gymnachirus melas Naked sole

Nichols, 1916

Gymnachirus melas is nearly circular in profile, with a matting of dermal folds and fimbriae on blind side of head and margin of body, fleshy dorsal and anal fins, and a large number of black and white stripes. Mouth is terminal. Jaw teeth are villiform, arranged in patches, and limited to blind side of jaws. Vomer and palatine lack teeth. Gill rakers are absent. Opercular opening is greatly restricted, and opercular cavities are connected by interbranchial septum. Gill membranes are largely free of isthmus. Measurements are expressed as percent of SL: head length 18.9%–31.6%, caudal fin length 20.5%–35.4%, body depth 50.8%– 62.3%. Small rudimentary pectoral fin is located above gill slit on ocular side and has 1 to 3 rays; occasionally it is invaginated or hidden beneath integument. Pectoral fin on blind side is absent. Dorsal and anal fins are very fleshy and are separate from caudal fin. Dorsal fin has 62 to 70 rays, and anal fin has 44 to 52 rays. Pelvic fins are enclosed in common dermal envelope that is continuous with anal fin, and each fin consists of 5 rays. Anus is located on blind side, urinary papilla is located on ocular side, and both are near posterior margin of pelvic fin envelope. Blind side of head and mar-

gin of head on ocular side are covered with matting of dermal folds. Accessory lateral lines on ocular side, running perpendicular to lateral line, number 7 to 10. Third accessory lateral line on ocular side has 35 to 79 (usually 47 to 62) pores. Ocular side is pale tan to brown, with moderate to narrow brown to black stripes. Total stripes number 15 to 32 (usually 20 to 30), and caudal fin stripes number 3 to 5. Light interstripes are as wide as or slightly wider than primary stripes. Juveniles (less than 55 mm SL) are dark and have no stripes. Blind side is pale creamy white. This species occurs between 1 and 183 m in the western Atlantic from Cape Cod and the Bahamas to the Gulf of Mexico. In the Gulf of Mexico it occurs from the Dry Tortugas to Pensacola. Food consists of poriferans, bivalve mollusks, polychaetes, ostracods, amphipods, cumaceans, brachyurans, stomatopods, and lancelets. Maximum known size is 164 mm SL. references: Gunter 1936, 1939 (as Nodogymnus williamsoni); Longley and Hildebrand 1941 (as N. williamsoni); Dawson 1964, 1967c; Böhlke and Chaplin 1968; Robins and Ray 1986; Smith 1997; Munroe 2002e.

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868 PLEURONECTIFORMES

Gymnachirus nudus Kaup, 1858

Gymnachirus nudus is nearly circular in profile, with a matting of dermal folds and fimbriae on blind side of head and margin of body, fleshy dorsal and anal fins, and a large number of black and white stripes. Mouth is terminal. Jaw teeth are villiform, arranged in patches, and limited to blind side of jaws. Vomer and palatine lack teeth. Gill rakers are absent. Opercular opening is greatly restricted, and opercular cavities are connected by interbranchial septum. Gill membranes are largely free of isthmus. Measurements are expressed as percent of SL: head length 22.7%–28.4%, caudal fin length 23.6%–34.4%, body depth 50%– 58.4%. Small rudimentary pectoral fin is located above gill slit on ocular side and has 1 to 3 rays; occasionally it is invaginated or hidden beneath integument. Pectoral fin on blind side is absent. Dorsal and anal fins are very fleshy and are separate from caudal fin. Dorsal fin has 60 to 69 rays, and anal fin has 44 to 50 rays. Pelvic fins are enclosed in common dermal envelope that is continuous with anal fin, and each fin consists of 5 rays. Blind side of head

and margin of head on ocular side are covered with matting of dermal folds. Accessory lateral lines on ocular side, running perpendicular to lateral line, number 7 to 10. Third accessory lateral line on ocular side has 26 to 59 (usually 34 to 44) pores. Ocular side is pale tan to brown, with moderately narrow to narrow brown to black stripes. Total stripes number 11 to 21 (usually 15 to 18), and caudal fin stripes number 2 to 5. Light interstripes are 1.5 times as wide as primary stripes. Juveniles (less than 55 mm SL) are dark and have no stripes. Blind side is pale creamy white to tan. This species occurs between the shoreline and 91 m in the western Atlantic from the southern Gulf of Mexico to northern Brazil, including the Greater Antilles. In the Gulf of Mexico it is known from the northwestern coast of Campeche. Maximum known size is 128 mm SL. references: Hildebrand 1955 (possibly as Gymnachirus williamsoni); Dawson 1964; Matsuura 1983q; Cervigón 1993a; Munroe 2002e.

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ACHIRIDAE 869

Gymnachirus texae Fringed sole

(Gunter, 1936)

Gymnachirus texae is nearly circular in outline, with a matting of dermal folds and fimbriae on blind side of head and margin of body, fleshy dorsal and anal fins, and a large number of black and white stripes. Mouth is terminal. Jaw teeth are villiform, arranged in patches, and limited to blind side of jaws. Vomer and palatine lack teeth. Gill rakers are absent. Opercular opening is greatly restricted, and opercular cavities are connected by interbranchial septum. Branchiostegal membranes are largely free of isthmus. Measurements are expressed as percent of SL: head length 22.8%–34.9%, caudal fin length 23.1%–33.3%, body depth 46.6%– 62%. Small rudimentary pectoral fin is located above gill slit on ocular side and has 1 to 3 rays; occasionally it is invaginated or hidden beneath integument. Pectoral fin on blind side is absent. Dorsal and anal fins are very fleshy and are separate from caudal fin. Dorsal fin has 57 to 66 rays, and anal fin has 41 to 48 rays. Pelvic fins are enclosed in common dermal envelope that is continuous with anal fin, and each fin consists of 5 rays. Blind side of head and margin of head on ocular side

are covered with matting of dermal folds. Accessory lateral lines on ocular side, running perpendicular to lateral line, number five to nine. Third accessory lateral line on ocular side has 21 to 48 (usually 27 to 38) pores. Ocular side is pale tan to brown, with moderate to narrow brown to black stripes. Total stripes number 25 to 49 (usually more than 30), and caudal fin stripes number 3 to 6. Light interstripes are about twice as wide as primary stripes. Juveniles (less than 27 mm SL) may be dark and without stripes. Blind side is pale creamy white to tan. This species occurs between 20 and 187 m on mud bottoms in the western Atlantic from the Gulf of Mexico and Yucatán Peninsula. In the Gulf of Mexico it occurs from Cape San Blas, FL, to Campeche Bank and is absent along the coast of Florida south and east of Cape San Blas. references: Gunter 1936 (as Nodogymnus texae), 1952; Hildebrand 1954, 1955; Springer and Bullis 1956; Dawson 1964, 1967c; Robins and Ray 1986; Smith 1997; Munroe 2002e.

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870 PLEURONECTIFORMES

Trinectes inscriptus Scrawled sole

(Gosse, 1851)

Trinectes inscriptus is nearly circular in profile, with a matting of dermal folds and fimbriae on blind side, ctenoid scales, and a dense reticular pattern of dark lines on a light background on ocular side. Anterior naris on ocular side opens at end of short, stout tube, and posterior naris opens immediately in front of lower eye. Mouth is subterminal, with snout slightly overhanging jaws. Jaw teeth are villiform and present in blind side of jaws only. Opercular opening is greatly restricted, and interbranchial foramen is lacking. Branchiostegal membranes are free of isthmus. Measurements are expressed as percent of SL: head length 28.5%– 30.1%, interorbital width 1.2%–2.6%, caudal fin length 28.4%–30.5%, body depth 58.4%– 65%. Small rudimentary pectoral fin is located above gill slit on ocular side and has 1 to 3 rays. Dorsal fin has 51 to 55 rays, and anal fin has 39 to 41 rays. Ocular-side pelvic fin has 4 rays and is continuous with anal fin. Blind-side pelvic fin inserts anterior to ocular pelvic fin and

has 5 rays. Entire body and most of head are covered with small ctenoid scales, including both sides of dorsal and anal fin rays. Fimbriae run along margins of mouth on ocular side and are broadly distributed on blind side of head and anterior dorsal and anal fin rays. Ocular side is tan, with fine dark brown reticular pattern extending over entire surface of head, body, and fins. Four to six fine and slightly irregular stripes run across body. Blind side is yellowish anteriorly and increasingly darkly pigmented distally. This species occurs in coastal waters, including estuaries, in the western Atlantic from southern Florida and the Bahamas to Venezuela and the Greater Antilles. It has not been reported from the Gulf of Mexico but may occur there. Maximum known size is 150 mm TL. references: Böhlke and Chaplin 1968; Robins and Ray 1986; Cervigón 1993a; Smith 1997; Munroe 2002e.

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ACHIRIDAE 871

Trinectes maculatus Hogchoker

(Bloch and Schneider, 1801)

Trinectes maculatus is elliptical in profile, with a matting of dermal folds and fimbriae on blind side, ctenoid scales, and light and dark spots or lines on a dark background on ocular side. Anterior naris on ocular side opens at end of short, stout tube, and posterior naris opens immediately in front of lower eye. Mouth is subterminal, with snout slightly overhanging jaws. Jaw teeth are villiform and present in blind side of jaws only. Opercular opening is greatly restricted, and interbranchial foramen is lacking. Gill membranes are free of isthmus. Measurements are expressed as percent of SL: head length 26.2%–30.9%, interorbital width 1.6%–2.1%, caudal fin length 28.5%–34.6%, body depth 50.6%– 61.3%. Pectoral fin is absent. Dorsal fin has 50 to 56 rays, and anal fin has 36 to 42 rays. Ocular-side pelvic fin has 3 rays and is continuous with anal fin. Blindside pelvic fin inserts opposite ocular-side pelvic fin and has 4 rays. Entire body and most of

head are covered with small ctenoid scales, including both sides of dorsal and anal fin rays. Fimbriae run along margins of mouth on ocular side and are broadly distributed on blind side of head and on anterior dorsal and anal fin rays. Fimbriae are also scattered over body. Ocular side is dark brown, with darker brown bars. Blind side is white to light tan, and occasionally is blotched or spotted. This species occurs between the shoreline and 75 m in the western Atlantic from the southern Gulf of Maine to Panama, including the entire Gulf of Mexico. It also occurs in estuaries and fresh waters with connections to the sea. Food consists of polychaetes and crustaceans. Maximum known size is 200 mm TL.

references: Springer and Woodburn 1960; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith 1997; Munroe 2002e.

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872

CYNOGLOSSIDAE

Tonguefishes

Cynoglossids are elongate and oval in profile, with eyes on left side of head, pectoral fins absent or represented by a membrane, and dorsal and anal fins confluent with caudal fin. Anterior profile of head is moderately to broadly rounded or pointed. Nostrils are paired and are symmetrically located on either side of head. Mouth is subterminal to inferior, and asymmetrical. Species with subterminal mouths have snout hooked over mouth, and mouth is contorted. Jaw teeth are small. Eyes are usually very small and close set. Pupillary operculum, a triangular or rounded structure, is present in some shallow-water (usually) species of Symphurus. Preopercular margin is concealed beneath scales and skin. Skin of lower jaw and interoperculum is continuous ventrally and covers isthmus and branchiostegal rays. Gill membranes are connected and joined to isthmus. Dorsal fin originates above or anterior to lower eye and is continuous with caudal fin. Blind-side pelvic fin inserts on ventral midline, is often connected with anal fin, and has 4 rays. Ocular-side pelvic fin is present or absent. Anal fin is joined to caudal fin. Caudal fin rays are unbranched. Lateral line is present or absent. Body, head, and often dorsal and anal fins are covered with scales. Cynoglossids occur worldwide in tropical to temperate seas, and occasionally in freshwater with connections to the sea. They are benthic on continental and insular shelves and the upper slopes to a depth of 1,500 m. There are about 125 species in three genera, 20 species in one genus in the western Atlantic, and 11 or 12 species in the Gulf of Mexico.

references: Ochiai 1964; Nelson 1994; Munroe et al. 1995; Munroe 2002f.

Key to the Species of the Gulf of Mexico (Adapted from Munroe 1998; 2002f) 1a. Peritoneum black, usually visible through abdominal wall; caudal fin rays usually 12; pupillary operculum absent; teeth present along entire margin of ocular-side jaws . . . . 2 1b. Peritoneum unpigmented; caudal fin rays 10 to 12; pupillary operculum present or absent; teeth present or absent along entire margin of ocular-side jaws . . . . . . . . . . . . . . 7 2a. Blind side of body peppered with melanophores (usually concentrated along bases of dorsal and anal fins) . . . . . . . . . . . . . . . . . . . . . . . . Symphurus pelicanus p. 881 2b. Blind side of body not peppered with melanophores . . . . 3

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CYNOGLOSSIDAE 873

3a. Dorsal fin rays 93 to 104; anal fin rays 80 to 89; ocular surface uniformly colored and without distinct crossbands but with large dark brown diamond-shaped blotch on caudal region of body; basal margins of dorsal and anal fins with dark brown stripe but without blotches; vertebrae 51 to 56 (usually 52 to 54) . . . . Symphurus marginatus p. 878 3b. Dorsal fin rays usually less than 95; anal fin rays usually 84 or fewer; ocular side of body with or without distinct pattern of crossbands and without dark brown diamondshaped blotch on caudal region of body; dorsal and anal fins with or without pigmented blotches; vertebrae 47 to 53 (usually 52 or fewer) . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4a. Scales in longitudinal series 62 to 75; hypurals five; anal fin rays 68 to 74; inner opercular lining and both sides of isthmus usually lightly pigmented; vertebrae 45 to 49 (usually 47 to 49) . . . . . . . . . . . . . . . . . . . . Symphurus piger p. 882 4b. Scales in longitudinal series usually 77 to 100; hypurals four; anal fin rays 71 to 84; inner opercular lining and isthmus unpigmented; vertebrae 47 to 52 (usually more than 48) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5a. Dorsal fin rays 83 to 88; anal fin rays 71 to 75; vertebrae 47 to 49; scales in longitudinal series 77 to 87; ocular surface usually yellow or light straw colored, with one or two prominent, complete crossbands just posterior to opercular opening; dorsal and anal fins without stripe along basal margins; adults not exceeding 80 mm SL . . . . . . . . . . . . . . . . . . . . . . . . . Symphurus pusillus p. 884 5b. Dorsal fin rays 87 to 95; anal fin rays 74 to 84; vertebrae 50 to 53; scales in longitudinal series 85 to 100; ocular surface usually dark brown, straw colored, or yellowish, with series of mostly incomplete crossbands posterior to opercular opening, or ocular surface uniformly pigmented and without crossbands; dorsal and anal fins frequently with dark brown stripe along basal margins, sometimes in combination with series of large, pigmented blotches alternating with unpigmented areas on dorsal and anal fins .............................................6 6a. Dorsal and anal fins usually with alternating series of prominent, darkly pigmented blotches; blotches usually wider than intervening unpigmented areas; no pigment spot on scaly base of caudal fin; eyeballs round, usually contiguous or nearly contiguous within fleshy orbital sac . . . . . . . . . . . . . . . . . . . . . . . Symphurus stigmosus p. 885 6b. Dorsal and anal fins usually without alternating series of prominent, darkly pigmented blotches; if blotches present, blotches as wide as or slightly narrower than intervening unpigmented areas; pigmented spot present on scaly base

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874 PLEURONECTIFORMES

of caudal fin; eyeballs longer than wide, separated by small space within fleshy orbital sac . . . . . . . . . . . . . . . . . . . . . . . Symphurus billykrietei p. 875 7a. Caudal fin rays usually 12; pupillary operculum absent . . . . . . . . . . . . . . . . . . . . . . . Symphurus civitatium p. 876 7b. Caudal fin rays usually 10 or 11; pupillary operculum present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8a. Caudal fin rays usually 11; large ocellated spot on caudal fin; dorsal and anal fins without spots; pupillary operculum well developed . . . . . . . . . Symphurus urospilus p. 886 8b. Caudal fin rays usually 10; no ocellated spot on caudal fin; if spot present on caudal fin (occasionally in S. diomedeanus), spots also present on posterior aspect of dorsal and anal fins; pupillary operculum present or absent . . . . . . . 9 9a. Dark brown blotch on caudal region of ocular side of body or single ocellated spot on posterior dorsal and anal fins; pupillary operculum present; no fleshy ridge on ocular side of lower jaw; ostia present in membranes at bases of dorsal and anal fins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9b. No dark brown blotch on caudal region of ocular side of body; no ocellated spots on posterior aspects of dorsal and anal fins; pupillary operculum and fleshy ridge on ocular side of lower jaw present or absent; no ostia in membranes at bases of dorsal and anal fins . . . . . . . . . . . . . . . . . . . . 11 10a. Dorsal fin rays 69 to 81 (usually 72 to 77); anal fin rays 55 to 64 (usually 56 to 64); vertebrae 41 to 44 (usually 41 to 43); scales in longitudinal series 55 to 67 . . . . . . . . . . . . . . . . . . . . . . . . . . Symphurus minor p. 879 10b. Dorsal fin rays 75 to 86 (usually 77 to 84); anal fin rays 60 to 70 (usually 62 to 67); vertebrae 43 to 47 (usually 44 to 46); scales in longitudinal series 59 to 78 . . . . . . . . . . . . . . . . . . . . . . . . . . Symphurus parvus p. 880 11a. Posterior dorsal and anal fins usually spotted; pupillary operculum present . . . . . . Symphurus diomedeanus p. 877 11b. Dorsal and anal fins without spots; pupillary operculum absent or only weakly developed . . . . . . . . . . . . . . . . . . . . . . . . Symphurus plagiusa p. 883

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CYNOGLOSSIDAE 875

Symphurus billykrietei Munroe, 1998 Chocolate banded tonguefish Symphurus billykrietei has 12 caudal fin rays, distinct bands on body, and no pupillary operculum. Anterior naris on ocular side terminates as a long tube that usually reaches anterior margin of lower eye. Posterior margin of maxilla extends to anterior margin of lower eye. Lower jaw on ocular side lacks fleshy ridge near posterior margin. Jaw teeth are developed on both sides. Measurements are expressed as percent of SL: head length 18.7%–22.6%, head width 20.7%–27%, body depth 25.1%– 32.8%; and as percent of head length: snout length 15.7%–24%, upper jaw length 17.9%– 25.3%, eye diameter 9.8%–14.3%. Dorsal fin originates above middle of upper eye and has 89 to 95 (usually 91 to 94) rays. Pelvic fin is moderately long, with posteriormost ray connected to body or anal fin by membrane. Anal fin has 76 to 84 (usually 77 to 81) rays. Caudal fin is moderately long. Both sides of body and head are covered with large ctenoid scales. Scales are absent on blind side of dorsal and anal fin rays. Scales in longitudinal series range from 80 to 100. Small scales also occur on

snout and on anterior and median surfaces of eyes. Dermal papillae are well developed on blind side of snout and occasionally on ocular side of snout. Vertebrae number 50 to 53. Ocular side is light to dark brown, with five to eight irregular bands that vary in intensity. Dorsal and anal fins are lightly pigmented anteriorly but increase in pigment intensity posteriorly, and pigment is more concentrated along bases of rays. A small dark spot is present on base of caudal fin. Opercular cavity and isthmus are unpigmented. Blind side is uniformly yellowish. Peritoneum is black. This species occurs between 117 and 650 m in the western Atlantic from southern Nova Scotia to Cape Hatteras, and from the Straits of Florida and the southern Gulf of Mexico off the Yucatán Peninsula. It is rare, however, south of Cape Hatteras. Maximum known size is 112 mm SL.

references: Munroe 1992 (as Symphurus species C, in part), 1998, 2002f.

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876 PLEURONECTIFORMES

Symphurus civitatium Ginsburg, 1951 Offshore tonguefish

Symphurus civitatium has 12 caudal fin rays and lacks a pupillary operculum. Anterior naris on ocular side terminates as a short tube that fails to reach anterior margin of lower eye. Posterior margin of maxilla extends to middle of pupil of lower eye. Lower jaw on ocular side has a fleshy ridge near posterior margin. Jaw teeth are well developed on blind side but are limited to a single row in anterior third of jaws or are totally absent from upper jaw on ocular side. Measurements are expressed as percent of SL: head length 17%–21.9%, head width 21.2%–27.1%, body depth 24.7%–32.8%; and as percent of head length: snout length 16.9%–23.1%, upper jaw length 18.1%– 28.9%, eye diameter 7%–11%. Dorsal fin originates anterior to or over anterior margin of upper eye and has 86 to 93 rays. Pelvic fin is short, and posteriormost pelvic ray is connected to body by membrane. Anal fin has 46 to 50 rays. Caudal fin is short. Both sides of body and head are covered with large ctenoid scales. Scales occur on ocular side of dorsal and anal fins but are largely absent on blind side. Scales in longitudinal series range from 74 to 82. Small scales also occur on snout. Dermal

papillae are well developed on blind side of snout and chin. Vertebrae number 66 to 83. Ocular side is light to dark brown, occasionally with dark bands running across head and body. Inner surface of opercular cavity and isthmus are darkly pigmented. Posterior third of dorsal and anal fins and caudal fin are darkly pigmented. Blind side is unpigmented. This species occurs between 1 and 172 m in the western Atlantic from Cape Hatteras and Bermuda to the Gulf of Mexico. In the Gulf of Mexico it occurs at the Dry Tortugas and from the Florida Panhandle to Yucatán. Adults are most common between 11 and 45 m on sandy to silty substrates, and juveniles occur in estuaries. Maximum known size is 152 mm SL.

references: Ginsburg 1951; Hildebrand 1954, 1955; Springer and Bullis 1956; Reséndez Medina 1979; Topp and Hoff 1972; Hoese and Moore 1977, 1998; Darcy and Gutherz 1984; Robins and Ray 1986; (all previous as S. civitatus); Munroe 1991, 1992 (both as S. civitatius), 1998; Boschung 1992 (as S. civitatus); Munroe et al. 2001; Munroe 2002f.

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CYNOGLOSSIDAE 877

Symphurus diomedeanus Spottedfin tonguefish

(Goode and Bean, 1885)

Symphurus diomedeanus has a pupillary operculum, 10 caudal fin rays, and posterior sections of dorsal and anal fins marked with small dark spots. Anterior naris on ocular side terminates as a moderately short tube that just fails to reach anterior margin of lower eye. Posterior margin of maxilla extends to anterior margin or middle of pupil of lower eye. Lower jaw on ocular side lacks fleshy ridge near posterior margin. Jaw teeth are well developed on blind side but are limited to anterior half of lower jaw on ocular side. Measurements are expressed as percent of SL: head length 16.2%– 20.4%, head width 19.9%–26.3%, body depth 26.8%–32.4%; and as percent of head length: snout length 13%–20.9%, upper jaw length 16.5%–22.3%, eye diameter 8.2%–15.6%. Dorsal fin originates over anterior margin of upper eye and has 86 to 96 rays. Pelvic fin is short, and posteriormost ray is connected to body by a membrane. Anal fin has 69 to 80 rays. Caudal fin is short. Both sides of body and head are covered with large ctenoid scales. Scales occur on ocular side of dorsal and anal fins but are absent on blind side. Scales in longitudinal series range from 79 to 96. Small scales also occur on snout. Dermal papillae are well developed on blind side of snout, chin, and dorsal section of head. Vertebrae number 47 to 50.

Ocular side is dark brown, occasionally with obscure dark bands running across head and body and with mottling or dark blotches. Pupillary operculum is dark brown or black, or silvery in specimens collected over white sand. Inner surfaces of opercular cavity and isthmus are unpigmented. Posterior third of dorsal and anal fins, and caudal fin are dusky. One to five dark brown to black small, round spots are located on posterior third of dorsal and anal fins. Occasionally specimens have a dark spot on caudal fin. Blind side is unpigmented. This species occurs between 6 and 183 m in the western Atlantic from just north of Cape Hatteras to Brazil, including the Gulf of Mexico, Jamaica, and the Virgin Islands. It is most common between 21 and 80 m on hard or shelly substrates. In the Gulf of Mexico it occurs along the west coast of Florida to the south Texas coast, and along the west coast of Yucatán. Maximum known size is 207 mm SL. references: Longley and Hildebrand 1941; Ginsburg 1951; Hildebrand 1954, 1955; Springer and Bullis 1956; Hoese 1958; Cervigón 1966; Topp and Hoff 1972; Chittenden and Moore 1977; Hoese and Moore 1977, 1998; Darcy and Gutherz 1984; Robins and Ray 1986; Boschung 1992 (all previous as S. diomedianus); Munroe 1992, 1998, 2002f; Smith 1997 (as S. diomedianus).

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878 PLEURONECTIFORMES

Symphurus marginatus Margined tonguefish

(Goode and Bean, 1886)

Symphurus marginatus has 12 caudal fin rays and a dark brown diamond-shaped blotch on the caudal fin, but lacks a pupillary operculum. Anterior naris on ocular side terminates as a moderately long tube that just fails to reach anterior margin of lower eye. Posterior margin of maxilla extends to anterior margin of pupil of lower eye. Lower jaw on ocular side lacks a fleshy ridge near posterior margin. Jaw teeth occur on both sides in a single row along anterior four-fifths of upper jaw and along entire lower jaw. Measurements are expressed as percent of SL: head length 12.7%–22.1%, head width 14.7%–22.7%, body depth 20%– 31.5%; and as percent of head length: snout length 16.8%–33.1%, upper jaw length 18%– 33.1%, eye diameter 12.5%–24.8%. Dorsal fin originates over midlength of upper eye and has 93 to 104 rays. Pelvic fin is short, and posteriormost ray is connected to body or anal fin by a membrane. Anal fin has 80 to 89 rays. Caudal fin is short. Both sides of body and head are covered with large ctenoid scales. Scales occur on ocular side of dorsal and anal fins but are absent on blind side. Scales in longitudinal

series range from 86 to 99. Small scales also occur on snout. Dermal papillae are absent or are limited to blind side of snout. Vertebrae number 51 to 56 (usually 52 to 54). Ocular side is uniformly dark brown. Inner surface of opercular cavity and isthmus are unpigmented. Caudal fin blotch may extend onto caudal fin base and posteriormost dorsal and anal fin rays. Posterior third of dorsal and anal fins, and caudal fin are dusky. Dark stripe is present along bases of dorsal and anal fins. Blind side is slightly dusky or yellow. Peritoneum is darkly pigmented. This species occurs between 37 and 752 m in the western Atlantic from New Jersey to southeastern Brazil, including the Bahamas, the Greater Antilles, and the Gulf of Mexico. It is most abundant between 320 and 550 m. In the Gulf of Mexico it occurs from the Dry Tortugas to Louisiana. Maximum known size is 146 mm SL.

references: Ginsburg 1951; Topp and Hoff 1972; Munroe 1992, 1998, 2002f.

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CYNOGLOSSIDAE 879

Symphurus minor Ginsburg, 1951 Largescale tonguefish

Symphurus minor has a well-developed pupillary operculum, 10 caudal fin rays, and a dark brown blotch near caudal peduncle. Anterior naris on ocular side terminates as a long tube that usually reaches anterior margin of lower eye. Posterior margin of maxilla extends to anterior margin of pupil of lower eye. Lower jaw on ocular side lacks fleshy ridge near posterior margin. Jaw teeth are well developed on blind side but are limited to a single row along anterior one-half or three-fourths of upper jaw and entire lower jaw of ocular side. Measurements are expressed as percent of SL: head length 20%–24.7%, head width 19.8%– 25.6%, body depth 20.7%–32.4%; and as percent of head length: snout length 14.6%– 22.2%, upper jaw length 17.2%–24.7%, eye diameter 11.8%–18.2%. Dorsal fin originates between anterior margin and midpoint of upper eye and has 69 to 81 (usually 72 to 77) rays. Pelvic fin is moderately long, and posteriormost ray is connected to body or anal fin by a membrane. Anal fin has 55 to 64 (usually 57 to 60) rays. Both sides of body and head are covered with large ctenoid scales. Scales occur on ocular side of dorsal and anal fins but are absent on blind side. Scales in longitudinal se-

ries range from 55 to 67 (usually 57 to 64). Small scales also occur on snout. Dermal papillae are well developed on blind side of snout. Vertebrae number 41 to 44. Ocular side is light brown or straw colored, with dusky markings and spots. A dark brown oblong to diamond-shaped blotch occurs near caudal fin base. Occasionally faint dark bands occur on body. Inner surfaces of opercular cavity and isthmus are unpigmented. Dorsal and anal fins are progressively darker pigmented posteriorly but lack distinct spots or blotches. Blind side is white or yellowish. Peritoneum is unpigmented. This species occurs between 18 and 170 m in the western Atlantic from North Carolina to the eastern Gulf of Mexico. On rare occasions it has been reported from Nova Scotia. In the Gulf of Mexico it occurs as far west as DeSoto Canyon. This species is apparently associated with live bottoms and seldom occurs on terrigenous substrates. Maximum known size is 78 mm SL. references: Ginsburg 1951; Topp and Hoff 1972; Robins and Ray 1986; Boschung 1992; Munroe 1992, 1998, 2002f; Smith 1997.

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880 PLEURONECTIFORMES

Symphurus parvus Ginsburg 1951 Pygmy tonguefish

Symphurus parvus has a well-developed pupillary operculum, 10 caudal fin rays, and a dark brown blotch near caudal fin base. Anterior naris on ocular side terminates as a long tube that usually reaches anterior margin of lower eye. Posterior margin of maxilla extends to anterior margin of pupil of lower eye. Lower jaw on ocular side lacks a fleshy ridge near posterior margin. Jaw teeth are well developed on blind side but are limited to a single row along one-half to three-fourths of upper jaw and to a single row along three-fourths to entire length of lower jaw on ocular side. Measurements are expressed as percent of SL: head length 18%– 26%, head width 21.1%–27.9%, body depth 24.8%–32.8%; and as percent of head length: snout length 14.6%–22.9%, upper jaw length 18.5%–25.2%, eye diameter 10.5%–19.7%. Dorsal fin originates between anterior margin and midpoint of upper eye and has 75 to 86 (usually 77 to 84) rays. Pelvic fin is long, and posteriormost ray is connected to body or anal fin by a membrane. Anal fin has 60 to 70 (usually 62 to 67) rays. Both sides of body and head are covered with large ctenoid scales. Scales are absent on blind side of dorsal and anal fins. Scales in longitudinal series range from 59 to 78. Small scales also occur on snout, on anterior and medial surfaces of eyes, and on inter-

orbital region. Dermal papillae are well developed on blind side of snout. Vertebrae number 43 to 47 (usually 44 to 46). Ocular side is light brown or yellowish, with irregularly arranged dusky markings and spots. A dark brown oblong or diamond-shaped spot occurs near caudal fin base. Occasionally faint and incomplete dark bands are present on body. Inner surfaces of opercular cavity and isthmus are unpigmented. Dorsal and anal fins are progressively darker pigmented posteriorly but lack distinct spots or blotches. Blind side is white or yellowish. Peritoneum is unpigmented. This species occurs between 20 and 146 m in the western Atlantic from Cape Lookout, NC, to Trinidad, including the Gulf of Mexico. In the Gulf of Mexico it is most common off Florida, Alabama, Mississippi, and Louisiana. It is less common off Texas and in the southern Gulf off Yucatán and is absent in the western Gulf south of Texas. This species is associated with terrigenous substrates. Maximum known size is 87 mm SL. references: Ginsburg 1951; Topp and Hoff 1972; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Munroe 1992, 1998, 2002f; Smith 1997.

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CYNOGLOSSIDAE 881

Symphurus pelicanus Longtail tonguefish

Ginsburg, 1951

Symphurus pelicanus has 12 caudal fin rays, ocular side uniformly pigmented, and no pupillary operculum. Anterior naris on ocular side terminates as a short tube that fails to reach anterior margin of lower eye. Posterior margin of maxilla extends to midpoint of lower eye. Lower jaw on ocular side lacks a fleshy ridge near posterior margin. Jaw teeth are well developed on blind side but are limited to a single row along jaws of ocular side. Measurements are expressed as percent of SL: head length 21.1%–25.4%, head width 21%–25.9%, body depth 23.5%–31.5%; and as percent of head length: snout length 13.6%–22.6%, upper jaw length 19.3%–25%, eye diameter 9.3%–15.8%. Dorsal fin originates between midlength and posterior margin of upper eye and has 77 to 85 (usually 80 to 83) rays. Pelvic fin is long, and posteriormost ray is connected to body or anal fin by a membrane. Anal fin has 64 to 70 rays. Caudal fin is long. Both sides of body and head are covered with large ctenoid scales. Scales are absent on blind side of dorsal and anal fins. Scales in longitudinal series range from 62 to 70. Small scales also oc-

cur on snout. Dermal papillae are well developed on both sides of snout. Vertebrae number 43 to 46. Ocular side is light brown or yellowish, with irregularly arranged dusky markings and spots. Occasionally lightly shaded marks or streaks occur on body. Inner surfaces of opercular cavity and isthmus are unpigmented or lightly pigmented. Dorsal and anal fins are pigmented like body. Blind side is lightly pigmented, with melanophores most concentrated along bases of dorsal and anal fins. Peritoneum is black. This species occurs between 24 and 133 m in the western Atlantic from the Florida Straits to Guyana, including the Gulf of Mexico and the Lesser Antilles. In the Gulf of Mexico it has been captured from Louisiana to southern Texas and the Gulf of Campeche. Maximum known size is 69 mm SL.

references: Ginsburg 1951; Hildebrand 1954; Topp and Hoff 1972; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Munroe 1992, 1998, 2002f; Smith 1997.

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882 PLEURONECTIFORMES

Symphurus piger (Goode and Bean, 1886) Deepwater tonguefish

Symphurus piger has 12 caudal fin rays and distinct bands on body, but lacks a pupillary operculum. Anterior naris on ocular side terminates as a long tube that reaches or nearly reaches anterior margin of lower eye. Posterior margin of maxilla extends to midpoint of lower eye. Lower jaw on ocular side lacks fleshy ridge near posterior margin. Jaw teeth are well developed on blind-side jaws, but are limited to a single row along three-fourths of length of upper jaw and a single row along length of lower jaw on ocular side. Measurements are expressed as percent of SL: head length 18.2%– 25.6%, head width 24.2%–31.3%, body depth 24.4%–35%; and as percent of head length: snout length 17.2%–28.2%, upper jaw length 18.8%–33.8%, eye diameter 10.1%–16.7%. Dorsal fin originates above posterior margin of upper eye and has 80 to 88 rays. Pelvic fin is moderately long, and posteriormost ray is connected to body or anal fin by a membrane. Anal fin has 68 to 74 rays. Both sides of body and head are covered with large ctenoid scales. Scales are absent on blind side of dorsal and anal fins. Scales in longitudinal series range from 62 to 75. Small scales also occur on snout,

on anterior and medial surfaces of eye, and on interorbital region. Dermal papillae are well developed on blind side of snout. Vertebrae number 45 to 49 (usually 47 to 49). Ocular side is dark brown, with 3 to 10 well-developed dark brown and narrow bands on head and body that extend onto fins as blotches. Dorsal and anal fins are lightly pigmented anteriorly but are progressively darker posteriorly. Opercular cavity and isthmus are lightly pigmented. Blind side is uniformly white. Peritoneum is black. This species occurs between 92 and 549 m in the western Atlantic from St. Augustine, FL, and the Bahamas to French Guiana, including the Gulf of Mexico and the Greater and Lesser Antilles. In the Gulf of Mexico it has been captured from the Mississippi Delta, southern Florida, and the west coast of Campeche. It is most common between 141 and 300 m. Maximum known size is 121 mm SL.

references: Ginsburg 1951; Topp and Hoff 1972; Hoese and Moore 1977, 1998; Robins and Ray 1986; Munroe 1992, 1998, 2002f.

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CYNOGLOSSIDAE 883

Symphurus plagiusa (Linnaeus, 1766) Blackcheek tonguefish

Symphurus plagiusa has 10 caudal fin rays and a conspicuous black spot on ocular-side operculum, but lacks a pupillary operculum. Anterior naris on ocular side terminates as a short tube that usually fails to reach anterior margin of lower eye. Posterior margin of maxilla extends to midpoint of lower eye. Lower jaw on ocular side has a fleshy ridge near posterior margin. Jaw teeth on blind side are well developed but are limited to a single row along lower jaw on ocular side. Measurements are expressed as percent of SL: head length 14.5%– 21%, head width 21.2%–26.5%, body depth 28.3%–32.6%; and as percent of head length: snout length 19.4%–24.7%, upper jaw length 18%–24.2%, eye diameter 8.3%–12.6%. Dorsal fin originates above anterior margin of pupil of upper eye and has 81 to 91 rays. Pelvic fin is moderately long, and posteriormost ray is connected to body or anal fin by a membrane. Anal fin has 66 to 75 rays. Both sides of body and head are covered with large ctenoid scales. Scales are present on both sides of posterior dorsal and anal fin rays. Scales in longitudinal series range from 76 to 86. Small scales also occur on snout and on interorbital region. Dermal papillae are well developed on blind side of snout, chin, and dorsal portion of head. Ver-

tebrae number 44 to 49. Ocular side is dull tan to dark brown. Dark bands are variably developed on head and body. Dorsal and anal fins are moderately dusky over their lengths but lack spots and blotches. Opercular cavity and isthmus are heavily pigmented. Black spot on outer surface of upper angle of operculum is most evident in large specimens and may be absent in small specimens. Blind side is uniformly white in adults. Peritoneum is unpigmented. This species occurs between the shoreline and 183 m in the western Atlantic from the New York Bight to the southern Gulf of Mexico, including the Florida Keys, the Bahamas, and Cuba. It also occurs in estuaries. Small specimens are generally found in the shallowest waters. This species is widely distributed in the Gulf of Mexico. Maximum known size is 172 mm SL.

references: Gunter 1945; Ginsburg 1951; Reid 1954; Hildebrand 1954, 1955; Böhlke and Chaplin 1968; Springer and Woodburn 1960; Chittenden and Moore 1977; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Munroe 1992, 1998, 2002f; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Munroe et al. 2001.

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884 PLEURONECTIFORMES

Symphurus pusillus (Goode and Bean, 1885) Northern tonguefish

Symphurus pusillus has 12 caudal fin rays and dark narrow bands on body, but lacks a pupillary operculum. Anterior naris on ocular side terminates as a short tube that usually fails to reach anterior margin of lower eye. Posterior margin of maxilla extends to anterior margin of pupil of lower eye. Lower jaw on ocular side lacks a fleshy ridge near posterior margin. Jaw teeth are well developed on blind side but are limited to a single row along three-fourths of upper jaw and to a single row along lower jaw of ocular side. Measurements are expressed as percent of SL: head length 20.6%– 25.4%, head width 21.3%–26.2%, body depth 25.7%–31.7%; and as percent of head length: snout length 15.6%–23.8%, upper jaw length 18.9%–24.4%, eye diameter 10.3%–15.6%. Dorsal fin originates between anterior margin and midpoint of upper eye and has 83 to 88 rays. Pelvic fin is moderately long, and posteriormost ray is connected to body or anal fin by a membrane. Anal fin has 71 to 75 rays. Both sides of body and head are covered with large ctenoid scales. Scales are absent on blind side

of dorsal and anal fin rays. Scales in longitudinal series range from 77 to 87. Small scales also occur on snout and on interorbital region. Dermal papillae are well developed on blind side of snout, chin, and dorsal portion of head. Vertebrae number 47 to 49. Ocular side is yellowish, with two to six more or less continuous, narrow light brown bands across body. Basal half of dorsal and anal fins is brown pigmented. Blind side is uniformly whitish. Peritoneum is black. This species occurs between 102 and 233 m in the western Atlantic from Long Island to the Florida Straits, including the northern coast of Cuba and the northern Gulf of Mexico. In the Gulf of Mexico it is known from southern Florida to DeSoto Canyon. Specimens reported from Texas (Baughman 1950) are S. plagiusa (Munroe 1998). Maximum known size is 69 mm SL.

references: Ginsburg 1951; Topp and Hoff 1972; Robins and Ray 1986; Monroe 1992, 1998, 2002f.

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CYNOGLOSSIDAE 885

Symphurus stigmosus Blotchfin tonguefish

Munroe, 1998

Symphurus stigmosus has 12 caudal fin rays and distinct blotches on posterior part of dorsal and anal fins, but lacks a pupillary operculum. Anterior naris on ocular side terminates as a long tube that usually reaches anterior margin of lower eye. Posterior margin of maxilla extends to between anterior margin of pupil and anterior margin of lower eye. Lower jaw on ocular side lacks a fleshy ridge near posterior margin. Jaw teeth are well developed on blind side but are limited to a single row along jaws on ocular side. Measurements are expressed as percent of SL: head length 20.6%– 22.8%, head width 23%–27%, body depth 25.2%–31%; and as percent of head length: snout length 16%–22.6%, upper jaw length 17.3%–25%, eye diameter 12.8%–17.1%. Dorsal fin originates between anterior margin of upper eye and anterior margin of pupil of upper eye and has 92 to 95 rays. Pelvic fin is moderately long, and posteriormost ray is connected to body or anal fin by a membrane. Anal fin has 78 to 81 rays. Both sides of body and head are covered with large ctenoid scales. Scales are absent on blind side of dorsal and anal fin rays. Scales in longitudinal series range

from 98 to 100. Small scales also occur on snout and on anterior and medial surfaces of eyes. Dermal papillae are well developed on blind side of snout. Vertebrae number 51 or 52. Ocular surface is yellowish to yellowish brown, without bands or blotches but occasionally with diffuse mottling or small melanophores. Dorsal and anal fins have a continuous brown stripe along their bases and four to six distinct dark brown or black blotches on posterior twothirds of fins. Blotches on dorsal and anal fins are wider than intervening unpigmented areas. Blind side is uniformly yellowish. Peritoneum is black. This species occurs between 192 and 373 m in the western Atlantic from southeastern Florida to southeastern Yucatán Peninsula, including the Lesser Antilles. It has not been collected in the Gulf of Mexico, but records from the Straits of Florida in the Dry Tortugas region and from the Straits of Yucatán suggest that it may occur there. Maximum known size is 126 mm SL. references: Munroe 1992 (in part as Symphurus species C), 1998, 2002f.

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886 PLEURONECTIFORMES

Symphurus urospilus Ginsburg, 1951 Spottail tonguefish

Symphurus urospilus has a well-developed pupillary operculum, 11 caudal fin rays, and 4 to 11 dark brown bands. Anterior naris on ocular side terminates as a short tube that fails to reach anterior margin of lower eye. Posterior margin of maxilla extends to middle of lower eye. Lower jaw on ocular side has a fleshy ridge near posterior margin. Jaw teeth are well developed on blind side but are limited to an incomplete row along lower jaw on ocular side. Measurements are expressed as percent of SL: head length 15.5%–21.3%, head width 22.4%–30.8%, body depth 28.5%–36.6%; and as percent of head length: snout length 13.8%–25.5%, upper jaw length 18.8%– 26.1%, eye diameter 10.2%–17%. Dorsal fin originates above anterior margin of upper eye and has 82 to 90 rays. Pelvic fin is moderately long, and posteriormost ray is connected to body or anal fin by a membrane. Anal fin has 64 to 74 rays. Both sides of body and head are covered with large ctenoid scales. Scales are absent on blind side of dorsal and anal fin rays. Scales in longitudinal series range from 67 to

82. Small scales also occur on snout. Dermal papillae are well developed on blind side of snout. Vertebrae number 44 to 48. Ocular side is dark brown, with 4 to 11 dark brown bands on head and body but not on dorsal and anal fins. Dorsal and anal fins are dark brown. Distal half of caudal fin has a round, dark ocellated spot surrounded by unpigmented area. Blind side is white. Peritoneum is unpigmented. This species occurs between 5 and 324 m, but usually shallower than 50 m., in the western Atlantic from Cape Hatteras to southern Florida and the Florida Keys, including the Gulf of Mexico and Cuba. In the Gulf of Mexico it occurs from the Dry Tortugas, off Florida and Alabama, off Texas, and off the west coast of the Yucatán Peninsula. Maximum known size is 166 mm SL. references: Ginsburg 1951; Hildebrand 1955; Springer and Bullis 1956; Bullis and Thompson 1965; Topp and Hoff 1972; Dennis and Bright 1988a; Boschung 1992; Munroe 1992, 1998, 2002f; Smith 1997; Hoese and Moore 1998.

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887

TETRAODONTIFORMES Tetraodontiforms are a highly corroborated monophyletic group derived from an unknown taxon, presumably within Perciformes. The order consists of nine families, and seven of these occur in the Gulf of Mexico.

Key to the Families of the Gulf of Mexico (Adapted from Smith and Heemstra 1986c) 1a. Jaw teeth in each jaw fused to each other and to jaw bones to form sharp-edged dental plates . . . . . . . . . . . . . . . . . . 2 1b. Jaw teeth separate and protruding from jaw bone sockets 4 2a. Body truncate posteriorly; caudal fin absent, replaced by thick rudderlike structure without visible fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Molidae p. 939 2b. Body not truncate posteriorly; caudal fin present and fin rays visible . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3a. Premaxilla and dentary each fused with its antimere, resulting in undivided tooth plate in each jaw; skin bearing large, sharp spines . . . . . . . . . . . . . . . . Diodontidae p. 930 3b. Premaxilla and dentary not fused with their antimeres, resulting in median suture between antimeres; skin naked or with small spines . . . . . . . . . . . . . Tetraodontidae p. 919

4a. Pelvic fins present and consisting of long spine and one or two rays; dorsal fin with six spines, last may be rudimentary . . . . . . . . . . . . . . . . . . Triacanthodidae p. 889 4b. Pelvic fins absent; spinous dorsal fin absent or with two or three spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5a. Body compressed, covered with platelike bony scales or minute spinelike scales; two dorsal fins, first with one or more spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 5b. Body robust, covered with bony armor of hexagonal plates fused into solid box; single dorsal fin consisting of rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ostraciidae p. 913

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888 TETRAODONTIFORMES

6a. Body covered with hard, platelike bony scales; second dorsal fin spine longer than half the length of first; upper and lower jaws each with eight outer teeth . . . Balistidae p. 893

6b. Body covered with minute spinelike scales; second dorsal fin spine shorter than half the length of first; upper and lower jaws each with four or six outer teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monacanthidae p. 901

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889

TRIACANTHODIDAE

Spikefishes

Spikefishes are moderately deep to deep and compressed, with jaw teeth separate from each other and from the jaw bones, and dorsal fin and pelvic fins supported by spines and rays. Snout is short to elongate and tubular. Nostril is paired and located in scaleless area in front of eye. Eye is large and located near anterior margin of head. Mouth is small and usually terminal. Jaw teeth are conical to truncate, with 10 or more in outer row, but may be greatly reduced to lacking in one jaw. Gill opening is short and located in front of pectoral fin base. Pectoral fin inserts below midflank and is relatively small. First dorsal fin has six spines that decrease in size posteriorly. Second dorsal fin has 12 to 18 rays. Pelvic fin has one prominent spine followed by 1 or 2 rays. Anal fin consists of 11 to 16 rays. Most pectoral, dorsal, and anal fin rays are branched. Dorsal and pelvic spines can be locked in place, generally at right angles to body. Caudal fin is rounded to truncate. Caudal peduncle is slender and slightly compressed. Body is covered with small scales that have erect spinulose processes. Lateral line is inconspicuous. Maximum known size is 200 mm TL. Spikefishes occur worldwide in tropical to warm temperate seas. All species occur near the bottom between 46 and 900 m. Food consists of benthic invertebrates. There are 20 species in 11 genera, 5 species in 3 genera in the western Central Atlantic, and 3 species in 2 genera in the Gulf of Mexico.

references: Tyler 1968, 1986; Nelson 1994; Matsuura 2001a, 2002a.

Key to the Species of the Gulf of Mexico (Adapted from Tyler 1968) 1a. Origin of first dorsal fin anterior to (rarely over) upper edge of gill slit; jaw teeth in two rows, with 1 to 10 (usually 2 to 4) medial to complete outer tooth rows . . . . . . . . . . . . . . . . . . . . . . . . Parahollardia lineata p. 892 1b. Origin of first dorsal fin distinctly behind (rarely over) upper edge of gill slit; jaw teeth in a single row . . . . . . . . 2 2a. Dorsal fin rays 16 to 18 (usually 17 or 18); anal fin rays 14 to 16 (usually 15); pectoral fin rays 12 to 15 (usually 14); width of pelvis between pelvic spines 14% to 17% of its length . . . . . . . . . . . . . . . . . . . . . Hollardia hollardi p. 890 2b. Dorsal fin rays 15 or 16; anal fin rays 14 or 15 (usually 14); pectoral fin rays 12 or 13 (usually 12); width of pelvis between pelvic spines 20% to 25% of its length . . . . . . . . . . . . . . . . . . . . . . . . . . . Hollardia meadi p. 891

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890 TETRAODONTIFORMES

Hollardia hollardi Poey, 1861 Reticulate spikefish

Hollardia hollardi has an irregular pattern of dark reticulations on side. Snout is moderately long and acute. Jaw teeth are short, blunt, and conical; number 6 to 11 in a single row in upper jaw; and number 7 to 12 in a single row in lower jaw. Gill opening is slightly arched, slightly oblique, and slightly longer than pectoral fin base. Gill rakers on first arch number 15 to 20. Measurements are expressed as percent of SL: head length 35%– 45%, snout length 10%–14%, eye diameter 14%–19%, interorbital width 5%–9%, pectoral fin length 15%–18%, body depth 49%–73%. Pectoral fin has 12 to 15 (usually 14) rays. Dorsal fin originates posterior to upper edge of gill slit and has six stout spines and 16 to 18 rays. Pelvic fin consists of one spine and 1 or 2 rays;

spine is very long and stout, and rays are very short and unbranched. Anal fin is similar in location, shape, and size to rayed section of dorsal fin and has 14 to 16 rays. Scale spinules become branched with growth. Color in life is pinkish red, with olive to brown reticulations. In preservative, body is light, with irregular pattern of dark reticulations. This species occurs between 230 and 915 m in the western Atlantic from southern Florida and Bermuda to northern South America, including the Dry Tortugas and Campeche Bay in the Gulf of Mexico. Maximum known size is 179 mm SL. references: Kanazawa 1952; Matsuura 1983s, 2001a, 2002a; Robins and Ray 1986.

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TRIACANTHODIDAE 891

Hollardia meadi Tyler 1966 Spotted spikefish

Hollardia meadi has dark spots on upper part of body. Snout is moderately long and acute. Jaw teeth are short, blunt, and conical; number five or six in a single row of upper jaw; and number six to eight in a single row of lower jaw. Gill opening is slightly arched, slightly oblique, and slightly longer than pectoral fin base. Gill rakers on first arch number 13 to 17. Measurements are expressed as percent of SL: head length 38%– 42%, snout length 13%–16%, eye diameter 17%–20%, interorbital width 7%–9%, pectoral fin length 16%–18%, body depth 58%–73%. Pectoral fin has a moderately oblique base and 12 or 13 (usually 12) rays. Dorsal fin originates posterior to upper edge of gill slit and has six stout spines and 15 or 16 (usually 15) rays. Pelvic fin consists of one spine and 1 or 2 rays; spine is very long and

stout, and rays are very short and unbranched. Anal fin is similar in location, shape, and size to rayed section of dorsal fin and has 14 or 15 (usually 14) rays. Scale spinules become branched with growth. Isolated patches of scales are usually present on lips. Color in preservative is pale, and dark spots are arranged more or less in horizontal rows. Peritoneum is pale to silvery white. This species occurs between 54 and 448 m in the western Atlantic from the eastern Gulf of Mexico, the Bahamas, Cuba, Barbados, and northern South America. In the Gulf of Mexico it has been captured at 2624N, 8445W, at 384 m. Maximum known size is 70 mm SL. references: Tyler 1968; Robins and Ray 1986; Matsuura 2002a.

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892 TETRAODONTIFORMES

Parahollardia lineata (Longley, 1935) Jambeau

Parahollardia lineata has 5 to 10 stripes on body. Snout is relatively long and acute. Jaw teeth are short, blunt, and conical; number four to eight in outer row and one to five in inner row of upper jaw; and number five to nine in outer row and zero to two in inner row of lower jaw. Gill opening is straight, slightly oblique, and slightly longer than pectoral fin base. Gill rakers on first arch number 12 to 17. Measurements are expressed as percent of SL: head length 34%– 43%, snout length 14%– 18%, eye diameter 12%–18%, interorbital width 7%–10%, pectoral fin length 12%– 20%, body depth 44%–73%. Pectoral fin has 12 to 14 rays. Dorsal fin originates anterior to upper edge of gill slit and has 15 to 17 rays.

Pelvic fin consists of one spine and 1 ray; spine is very long and stout, and ray is very short and unbranched. Anal fin is similar in location, shape, and size to rayed section of dorsal fin and has 15 to 17 rays. Scale spinules become branched with growth. Color is pale yellow to pinkish, and stripes are reddish brown. This species occurs between 119 and 396 m in the western Atlantic from Virginia to Yucatán, including the northern Gulf of Mexico. Maximum known size is 200 mm TL.

references: Longley and Hildebrand 1940, 1941; Tyler 1968; Robins and Ray 1986; Boschung 1992; Matsuura 2002a.

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893

BALISTIDAE Triggerfishes

Triggerfishes are moderately deep to deep and moderately compressed, with two dorsal fins and branched dorsal and anal fin rays. Eye is large and located near dorsal margin of head. Mouth is terminal or almost terminal and relatively small. Left and right premaxillae and dentaries are not fused medially. Upper jaw is not protractile. Jaw teeth are separate from each other and protrude from sockets in jaw bones. Upper jaw has four outer incisor teeth and three inner and smaller teeth on each side. Lower jaw has four incisor teeth on each side. Gill opening is restricted to lateral slit and is located above or in front of pectoral fin base. Pectoral fin inserts just below midflank. First dorsal fin consists of three spines; first spine is stout and capable of being locked into upright position by second, and second spine is more than half the length of first. Second dorsal fin is separated from first and originates posterior to midlength. Pelvic fin is rudimentary, consisting of four pairs of enlarged scales forming a spine and attached to a long, movable pelvic bone. Anal fin is similar in shape and size to second dorsal fin and originates below second dorsal fin. Most dorsal, anal, and pectoral fin rays are branched. Caudal fin has convex, truncate, or double emarginate posterior margin, and caudal fin rays number 7 to 10. All fin rays are branched. Body and head are covered with strong, rough scales arranged in regular series. Scales above pectoral fin base are usually enlarged and slightly separated from each other to form a flexible tympanum. Lateral line is obscure or absent. Vertebrae number 18: 7 precaudal and 11 caudal. Gas bladder is present. Color ranges from drab to bright and boldly patterned. Maximum known size is 80 cm TL. Triggerfishes occur worldwide in tropical to warm temperate seas and are especially common on coral reefs. Most species are benthic. Food consists of hard-shelled invertebrates, such as mollusks and crabs, and also zooplankton and algae. Development is oviparous. Eggs are benthic. There are 40 species in 11 genera, and 6 species in 4 genera occur in the western Central Atlantic and the Gulf of Mexico. references: Smith and Heemstra 1986a; Tortonese 1986a; Nelson 1994; Matsuura 2001b, 2002b.

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894 TETRAODONTIFORMES

Key to the Species of the Gulf of Mexico (Adapted from Böhlke and Chaplin 1968; Matsuura 2002b) 1a. Teeth unequal, oblique, platelike, and deeply notched . . . 2 1b. Teeth even, perpendicular, and unnotched . . . . . . . . . . . . . . . . . . . . . . . . . . . Melichthys niger p. 899 2a. One or more enlarged bony scutes behind gill opening. . . 3 2b. No enlarged bony scutes behind gill opening . . . . . . . . . . 4 3a. Two distinct curved blue stripes on head; dorsal fin rays 29 to 31 (usually 30); anal fin rays 27 or 28 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Balistes vetula p. 896 3b. No blue stripes on head; dorsal fin rays 26 to 29 (usually 27 or 28); anal fin rays 23 to 26 (usually 24 or 25) . . . . . . . . . . . . . . . . . . . . . . . . . . . Balistes capriscus p. 895 4a. Cheek with three narrow parallel grooves, darker than surrounding areas; chin distinctly projecting . . . . . . . . . . . . . . . . . . . . . . . . Xanthichthys ringens p. 900 4b. Cheek closely scaled, without parallel grooves; chin not projecting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5a. Body covered with pale spots; dorsal fin rays 23 to 25; anal fin rays 20 to 22; pectoral fin rays 13 to 15 . . . . . . . . . . . . . . . . . . . . . . Canthidermis maculata p. 897 5b. Body without pale spots; dorsal fin rays 25 to 28 (usually 26 or 27); anal fin rays 23 to 25; pectoral fin rays 15 or 16 . . . . . . . . . . . . . . . . . . . . . . Canthidermis sufflamen p. 898

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BALISTIDAE 895

Balistes capriscus Gray triggerfish

Gmelin, 1789

Balistes capriscus has notched teeth and bony scutes above pectoral fin base. Deep groove is located in front of eye. Teeth at symphyses of upper and lower jaws are pointed, and those on lateral aspects of jaws have incised biting surfaces. Upper jaw has two rows of teeth, with eight teeth in outer row and six in inner row. Lower jaw has a single row of six teeth. Gill rakers on first arch number 31 to 35. Measurements are expressed as percent of SL: head length 36%–37%, snout length 25%–26%, eye diameter about 8%, pectoral fin length 15%–16%, body depth at origin of anal fin 48%–50%. Pectoral fin has 15 rays. Second dorsal fin is elevated anteriorly and has 26 to 29 (usually 27 or 28) rays. Anal fin is similar in shape to second dorsal fin and has 23 to 26 (usually 24 or 25) rays. Caudal fin is convex in juveniles and double emarginate in adults. Scales in lateral series number 57 to 63. Color is olivaceous gray, with three dark blotches or broad bars and blue to purplish spots on upper

flank, lighter spots on lower flank, and chin light tan. Second dorsal and anal fins are dark brown, with dark spots more or less arranged in rows. This species occurs in the tropical to warm temperate Atlantic Ocean. In the western Atlantic it occurs to a depth of 50 m, from Nova Scotia and Bermuda to Argentina, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is found throughout the Gulf of Mexico over its depth range. Habitats include coral reefs and grassy and sandy bottoms. Food consists of benthic invertebrates. Maximum known size is 300 mm TL.

references: Longley and Hildebrand 1941; Moore 1967; Randall 1968a; Hoese and Moore 1977, 1998; Matsuura 1983t, 2002b; Robins and Ray 1986; Cervigón 1993a; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; SmithVaniz et al. 1999.

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896 TETRAODONTIFORMES

Balistes vetula Linnaeus, 1758 Queen triggerfish

Balistes vetula has notched teeth, bony scutes above pectoral fin base, and two blue bands on cheek. Deep groove is located in front of eye. Jaw teeth at symphyses of upper and lower jaws are pointed, and those on lateral aspects of jaws have incised biting surfaces. Upper jaw has two rows of teeth, with eight teeth in outer row and six in inner row. Lower jaw has a single row of teeth. Gill rakers number 35 to 38. Measurements are expressed as percent of SL: head length 37%–38%, snout length 23%–29%, eye diameter 7%–12%, pectoral fin length 16%–17%, body depth at origin of anal fin 49%– 62%. Pectoral fin has 15 or 16 (usually 15) rays. Second dorsal fin is elevated anteriorly and has 29 to 31 (usually 30) rays. Anterior rays of second dorsal fin are filamentous in adults. Anal fin is similar in shape to second dorsal fin and has 27 or 28 rays. Caudal fin is emarginate in juveniles and lunate, with upper- and lowermost rays filamentous, in adults. Scales in horizontal series number 56 to 63. Color is greenish or bluish gray dorsally and orangish yellow on lower section of head

and belly, with two curved blue bands extending from snout to below pectoral fin base and a third band running below lower lip. Narrow, dark irregular lines radiate from eye. Median fins are dark blue, with median margins light blue. Caudal peduncle has a light blue band. This species occurs in the tropical and warm temperate Atlantic. In the western Atlantic it occurs from near shore to about 100 m, from Nova Scotia and Bermuda to southeastern Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is associated with coral reefs, sandy and rubble areas around reefs, and also sea-grass beds. Food consists of sea urchins and other invertebrates. Maximum known size is 500 mm TL.

references: Longley and Hildebrand 1941; Moore 1967; Böhlke and Chaplin 1968; Randall 1968, 1996; Hoese and Moore 1977, 1998; Matsuura 1983t, 2002b; Robins and Ray 1986; Smith and Heemstra 1986b; Tortonese 1986a; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999.

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BALISTIDAE 897

Canthidermis maculata Rough triggerfish

(Bloch, 1786)

Canthidermis maculata has evenly scaled cheeks and pale spots on body. A deep groove is located in front of eye. Jaw teeth have scalloped biting surfaces, and lateral teeth are diagonally oriented and converge medially. Upper jaw has two rows of teeth, with eight teeth in outer row and six in inner row. Lower jaw has a single row of teeth. Gill opening is slitlike and located largely above pectoral fin base. Gill rakers on first arch number 24 to 32. Measurements are expressed as percent of SL: head length 34%–36%, snout length 19%– 23%, eye diameter 7%–9%, pectoral fin length 13%–15%, body depth 51%– 61%. Pectoral fin has 13 to 15 rays. Second dorsal fin has 23 to 25 rays. Anal fin has 20 to 22 rays. Second dorsal and anal fins are elevated anteriorly and become falcate in large specimens. Caudal fin is slightly truncate in juveniles and double emarginate in adults. Scales in horizontal series

number 36 to 41. Scales above pectoral fin base are not enlarged or fused into tympanum. Vertebrae number 18: 7 precaudal and 11 caudal. Color is brown and slightly lighter ventrally, with small oval spots on body and fins. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs from New Jersey and Bermuda to Argentina, including the Florida Keys, the northern and western Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. It is associated with flotsam. Maximum known size is 500 mm TL. references: Berry and Baldwin 1966; Moore 1967; Randall 1996; Matsuura 1983t, 2001a, 2002b; Robins and Ray 1986; Smith and Heemstra 1986a; Smith 1997; Hoese and Moore 1998; Smith-Vaniz et al. 1999.

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898 TETRAODONTIFORMES

Canthidermis sufflamen (Mitchill, 1815) Ocean triggerfish

Canthidermis sufflamen has very rough scales and lacks a tympanum. A deep groove is located in front of eye. Jaw teeth have scalloped biting surfaces, and lateral teeth are diagonally oriented and converge medially. Upper jaw has two rows of teeth, with eight teeth in outer row and six in inner. Lower jaw has a single row of teeth. Gill opening is slitlike and located largely above pectoral fin base. Gill rakers on first arch number 32 to 36. Measurements are expressed as percent of SL: head length 34%–39%, snout length 23%–26%, eye diameter 7%–11%, pectoral fin length 13%–16%, body depth 50%–55%. Pectoral fin has 15 or 16 rays. First dorsal fin is serrated along leading edge. Second dorsal fin has 25 to 28 (usually 26 or 27) rays. Anal fin has 23 to 25 rays. Second dorsal fin and anal fin are strongly elevated an-

teriorly and become falcate in large specimens. Caudal fin is slightly truncate in juveniles and double emarginate in adults. Scales in horizontal series number 53 to 58. Color is brownish gray, with a large dark brown spot at base of pectoral fun. This species occurs in the western and Central Atlantic. In the western Atlantic it occurs from Massachusetts and Bermuda to northern South America, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is pelagic and is most common over offshore reefs in clear water. Food consists of large zooplankton. Maximum known size is 600 mm TL. references: Randall 1968a, 1996; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Matsuura 2002b.

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BALISTIDAE 899

Melichthys niger Black durgon

(Bloch, 1786)

Melichthys niger has evenly scaled cheeks, and scales on posterior section of body are ridged and form keels. A deep groove is located in front of eye below nares. Jaw teeth have even biting surfaces, and lateral teeth are only slightly diagonally oriented. Upper and lower jaws each have eight teeth in a single row. Gill opening is slitlike and located largely above pectoral fin base. Gill rakers on first arch number 36 or 37. Measurements are expressed as percent of SL: head length 31%–33%, snout length 22%–23%, eye diameter 6%–7%, pectoral fin length 8%–12%, body depth 54%– 58%. Pectoral fin has 14 or 15 rays. Second dorsal fin has 32 to 34 rays. Anal fin has 28 to 31 rays. Second dorsal fin and anal fin are slightly elevated anteriorly and have straight margins. Caudal fin is double emarginate in juveniles and adults. Scales in lateral series number 54 to 60. Scales above pectoral fin and behind gill opening are enlarged and platelike, forming a flexible tympanum. Vertebrae number 18: seven precaudal and 11 caudal. Color is black, with a greenish sheen. In life, blue

lines radiate from eye, run along base of dorsal fin and anal fin, and extend along caudal fin margin. This species occurs worldwide in tropical to warm temperate seas to about 30 m. In the western Atlantic it occurs from southern Florida and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. This species is generally reported to be absent from the Gulf of Mexico, but it is recorded from the West Flower Garden Bank off east Texas (Bright and Cashman 1974). This species most frequently occurs around islands and over outer reefs. Food consists of calcareous algae and zooplankton. Maximum known size is 500 mm TL.

references: Berry and Baldwin 1966; Moore 1967; Böhlke and Chaplin 1968; Randall 1968a, 1996; Randall and Klausewitz 1973; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Robins and Ray 1986; Smith and Heemstra 1986a; Smith 1997; Smith-Vaniz et al. 1999; Matsuura 2001b, 2002b.

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900 TETRAODONTIFORMES

Xanthichthys ringens Sargassum triggerfish

(Linnaeus, 1758)

Xanthichthys ringens has a distinct chin and three slightly diagonal grooves on cheeks. Jaw teeth have scalloped biting surfaces, and lateral teeth are diagonally oriented and converge medially. Gill opening is slitlike and located above pectoral fin base. Gill rakers on first arch number 36 to 40. Measurements are expressed as percent of SL: head length about 38%, snout length about 22%, eye diameter about 9%, pectoral fin length about 16%, and body depth about 56%. Pectoral fin has 13 or 14 rays. First dorsal fin spine is serrated along leading edge. Second dorsal fin has 26 to 29 rays. Anal fin has 23 to 27 rays. Second dorsal fin and anal fin are moderately elevated anteriorly. Caudal fin is slightly convex in juveniles and emarginate in adults. Scales above pectoral fin base are not enlarged or fused to form tympanum. Scales in lateral series number 39 to 44. Color

is light brownish gray, with rows of dark brown spots on body. Grooves on cheek are dark brown. Bases of second dorsal fin and anal fin are dark brown. Caudal fin has orangish red upper, lower, and posterior margins. This species occurs in the western Atlantic from South Carolina and Bermuda to northeastern Brazil, including the Gulf of Mexico, the Bahamas, and the Greater and Lesser Antilles. Juveniles are pelagic and associated with Sargassum, and adults are benthic, generally at depths greater than 31 m. Maximum known size is 243 mm TL. references: Dawson 1962b; Moore 1967; Böhlke and Chaplin 1968; Randall 1968a, 1996; Randall et al. 1978; Dennis and Bright 1988a; Hoese and Moore 1977, 1998; Robins and Ray 1986; Schaldach et al. 1997; Matsuura 2002b.

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901

MONACANTHIDAE

Filefishes

Filefishes are moderately deep to deep and compressed, with two dorsal fins and unbranched dorsal and anal fin rays. Eyes are large and located near dorsal margin of head. Nostril is paired and located in a scaleless depression in front of eye. Mouth is small and usually terminal, and upper jaw is not protractile. Upper jaw has three incisor-like teeth in outer row and two in inner row on each side. Lower jaw has similar-shaped teeth in a single row. Gill opening is short, vertical to oblique, and located in front of or above pectoral fin base. Pectoral fin inserts just below midflank and is fanlike. First dorsal fin consists usually of two spines, but occasionally of one spine, with first rather stout and capable of being locked into upright position by second, when present. Second dorsal fin is separated from first and has 22 to 52 unbranched rays. Pelvic fin consists of three or fewer pairs of scales encasing posterior-end pelvic bone that projects through skin. Pelvic bone is movably attached to pectoral girdle and supports ventral abdominal skin flap that extends from pelvic bone to anus. Abdominal skin flap is expanded or contracted by movement of pelvic bone. Anal fin is similar in shape and size to second dorsal fin and has 20 to 62 unbranched rays. Caudal peduncle bears two pairs of prominent spines or a patch of bristles in some species. Body and head are covered with minute to small scales bearing one or two spines. Gas bladder is present. Maximum known size is 500 mm TL. Filefishes occur worldwide in tropical to temperate seas between shore and 200 m. They are associated with coral or rocky reefs and sea-grass beds. There are 95 species in 31 genera, 10 species in 4 genera in the western Central Atlantic, and 10 species in 4 genera in the Gulf of Mexico. references: Hutchins 1986, 2001b; Tortonese 1986b; Nelson 1994; Smith 1997; Matsuura 2002c.

Key to the Species of the Gulf of Mexico (Adapted from Böhlke and Chaplin 1968) 1a. Pelvic bone with distinct external spine . . . . . . . . . . . . . . 2 1b. Pelvic bone with minute and rudimentary external spine or no external spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 2a. Deep groove on dorsal midline between dorsal fins; external spine on pelvic bone fixed, not flexible; first dorsal fin spine originates above anterior section of eye . . . . . . . . . . 3 2b. No deep groove on dorsal midline between dorsal fins; external spine on pelvic bone flexible; first dorsal fin spine originates above posterior section of eye . . . . . . . . . . . . . 4

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902 TETRAODONTIFORMES

3a. Caudal peduncle with two pairs of stout spines; pectoral fin rays usually 14; gill rakers on first arch 29 to 35 . . . . . . . . . . . . . . . . . . . . Cantherhines macrocerus p. 907 3b. Caudal peduncle with no stout spines; pectoral fin rays usually 13; gill rakers on first arch 34 to 46 . . . . . . . . . . . . . . . . . . . . . . . . . Cantherhines pullus p. 908 4a. Caudal peduncle with two to four pairs of enlarged spines; ventral flap of skin between end of pelvis and anus relatively large; no dorsal fin rays elongated; scales consist of one to eight unbranched spines . . . . . . . . . . . . . . . . . . . . 5 4b. Caudal peduncle without enlarged spines; ventral flap of skin between end of pelvis and anus relatively small; second dorsal fin ray elongated in males; scales consist of one to eight highly branched spines. . . . . . . . . . . . . . . . . . . . . . . 6 5a. Body depth between origins of second dorsal fin and anal fin 39% to 55% of SL; snout length 22% to 26% of SL . . . . . . . . . . . . . . . . . . . . . . . Monacanthus ciliatus p. 909 5b. Body depth between origins of second dorsal fin and anal fin 31% to 39% of SL; snout length 25% to 28% of SL . . . . . . . . . . . . . . . . . . . . . . . . Monacanthus tuckeri p. 910 6a. Dorsal fin rays usually 31 to 34 (rarely 29, 30, or 35); anal fin rays usually 31 to 34 (rarely 30 or 35) . . . . . . . . . . . . . . . . . . . . . . Stephanolepis hispidus p. 911 6b. Dorsal fin rays usually 27 to 29 (rarely 30); anal fin rays usually 27 to 29 (rarely 26 or 30) . . . . . . . . . . . . . . . . . . . . . . . . Stephanolepis setifer p. 912 7a. Dorsal fin rays 43 to 50; anal fin rays 46 to 52; pectoral fin rays usually 14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7b. Dorsal fin rays 32 to 41; anal fin rays 35 to 44; pectoral fin rays usually 12 or 13 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8a. Caudal peduncle depth 65% to 95% of caudal peduncle length; caudal fin length 18 to 26% of SL . . . . . . . . . . . . . . . . . . . . . . . . Aluterus monoceros p. 904 8b. Caudal peduncle depth 124% to 160% of caudal peduncle length; caudal fin length 33% to 61% of SL . . . . . . . . . . . . . . . . . . . . . . . . . . . Aluterus scriptus p. 906 9a. No pelvic spine; sides of body relatively rough . . . . . . . . . . . . . . . . . . . . . . . . . . Aluterus schoepfii p. 905 9b. Rudimentary pelvic spine; sides of body relatively smooth . . . . . . . . . . . . . . . . . . . . . . . . . Aluterus heudelotii p. 903

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MONACANTHIDAE 903

Aluterus heudelotii Dotterel filefish

Hollard, 1855

Aluterus heudelotii is elliptical in profile, with a rudimentary pelvic spine and a slender caudal peduncle. Snout is slightly concave. Jaw teeth are acutely pointed; those in upper jaw are in two rows, with three teeth in outer row and two teeth in inner row, and those in lower jaw are in a single row of three teeth. Head is slightly concave below chin. Gill slit is located in front of and partially above pectoral fin base and is about twice the length of pectoral fin base. Measurements are expressed as percent of SL: head length 29%–35%, snout length 24%– 29%, eye diameter 6%–10%, body depth at origin of anal fin 28%– 47%. Pectoral fin usually has 12 to 14 rays. Second dorsal fin has 36 to 41 rays. Pelvic spine is rudimentary or may

be absent. Anal fin has 39 to 41 rays. Caudal peduncle lacks enlarged scales. Color is brown to olive and scrawled with blue lines and spots. This species occurs in the tropical and warm temperate Atlantic Ocean. In the western Atlantic it occurs from Massachusetts and Bermuda to Brazil, including the northern and southern Gulf of Mexico and the Greater and Lesser Antilles. Maximum known size is 300 mm TL. references: Berry and Vogele 1961; Hoese and Moore 1977, 1998; Matsuura 1983u, 2002c; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999.

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904 TETRAODONTIFORMES

Aluterus monoceros Unicorn filefish

(Linnaeus, 1758)

Aluterus monoceros is elliptical in profile, with no pelvic spine and a slender caudal peduncle. Snout is slightly concave, and head is slightly concave below chin. Jaw teeth are acutely pointed; those in upper jaw are in two rows, with three teeth in outer row and two teeth in inner row, and those in lower jaw are in a single row of three teeth. Gill rakers on first arch number about 31. Gill slit is oblique, located in front of and partially above pectoral fin base, and about twice the length of pectoral fin base. Measurements are expressed as percent of SL: head length 27%–35%, snout length 23%–28%, eye diameter 4%– 8%, body depth at origin of anal fin 34%– 44%. Pectoral fin usually has 14 or 15 rays. Second dorsal fin has 46 to 50 rays. Pelvic spine is rudimentary in juveniles and absent in adults. Anal fin has

47 to 52 rays. Caudal peduncle is longer than it is deep and lacks enlarged scales. Color is gray to brown and mottled with dark blotches and marked with dark spots. Second dorsal fin and anal fin are pale yellowish. This species occurs worldwide in tropical and warm temperate seas. In the western Atlantic it occurs over continental and insular shelves from Massachusetts and Bermuda to Brazil, including the entire Gulf of Mexico. It is rare in the Antilles. Maximum known size is 600 mm TL. references: Berry and Vogele 1961; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Matsuura 1983u, 2002c; Hutchins 1986, 2001b; Robins and Ray 1986; Smith 1997; Smith-Vaniz et al. 1999.

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MONACANTHIDAE 905

Aluterus schoepfii Orange filefish

(Walbaum, 1792)

Aluterus schoepfii is elliptical in profile, with no pelvic spine, a slender caudal peduncle, and spots and stripes on body. Dorsal profile of head is slightly concave to straight. Jaw teeth are acutely pointed; those in upper jaw are in two rows, with three teeth in outer row and two teeth in inner row, and those in lower jaw are in a single row of three teeth. Head is slightly convex below chin. Gill rakers on first arch number about 21 to 27. Gill slit is oblique, located in front of and partially above pectoral fin base, and about twice the length of pectoral fin base. Measurements are expressed as percent of SL: head length 23%–34%, snout length 12%–28%, eye diameter 5%–9%, body depth at origin of anal fin 17%– 47%. Depth increases with growth. Pectoral fin has 11 or 12 rays. Second dorsal fin has 32 to 40 rays. Pelvic spine is absent. Anal fin has 35 to 41

rays. Caudal peduncle is slightly longer than to slightly shorter than it is deep and lacks enlarged scales. Color is pale gray to brown, with numerous small, round orange spots on body. This species occurs in shallow water in the western Atlantic from Nova Scotia and Bermuda to Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is associated with sea grass on sand to mud bottoms. Food consists of algae and sea grasses. Maximum known size is 600 mm TL.

references: Longley and Hildebrand 1941; Berry and Vogele 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Matsuura 1983u, 2002c; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999.

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906 TETRAODONTIFORMES

Aluterus scriptus Scrawled filefish

(Osbeck, 1765)

Aluterus scriptus is elliptical in profile, with no pelvic spine, a deep caudal peduncle, and spots and stripes on body. Snout is slightly upturned, and dorsal profile of head is concave. Jaw teeth are acutely pointed; those in upper jaw are in two rows, with three teeth in outer row and two teeth in inner row, and those in lower jaw are in a single row of three teeth. Head is slightly convex below chin. Gill rakers on first arch number about 21 to 27. Gill slit is very oblique, located in front of and partially above pectoral fin base, and about twice the length of pectoral fin base. Measurements are expressed as percent of SL: head length 29%– 34%, snout length 22%–29%, eye diameter 5%–9%, body depth at origin of anal fin 22%– 35%. Pectoral fin has 13 to 15 (usually 14) rays. Second dorsal fin has 43 to 49 rays. Pelvic spine is rudimentary or absent. Anal fin has 46 to 52 rays. Caudal peduncle is deeper than it is

long and lacks enlarged scales. Color is light bluish to olive or brown, with bluish green spots and lines and small black spots scattered on head and body. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs in shallow water from Nova Scotia and Bermuda to Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. Food consists of algae, sea grasses, hydrozoans, gorgonian corals, sea anemones, and tunicates. Maximum known size is 910 mm TL.

references: Longley and Hildebrand 1941; Berry and Vogele 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Hutchins 1986, 2001b; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Matsuura 2002c.

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MONACANTHIDAE 907

Cantherhines macrocerus (Hollard, 1853) Whitespotted filefish

Cantherhines macrocerus is moderately deep bodied, with a deep groove between dorsal fins and an external spine on pelvic bone. Snout is slightly concave. Jaw teeth are acutely pointed; those in upper jaw are in two rows, with three teeth in outer row and two teeth in inner row, and those in lower jaw are in a single row of three teeth. Gill rakers on first arch number 29 to 35. Gill slit is located in front of pectoral fin base and is relatively long. Pectoral fin usually has 13 or 14 rays. Second dorsal fin has 34 to 36 rays. Pelvic spine is short and stout, is not movable in anteroposterior direction, and has denticles on ventral surface and spinules along both sides. Anal fin has 29 to 32 rays. Scales bear one to several spines, and spines are branched distally. Adults possess two or three pairs of enlarged spines on caudal pe-

duncle. Color is brown, with or without numerous white spots on head and body. Ventral and posterior sections of body are orangish. Caudal fin is black. This species occurs in the western Atlantic from the east coast of Florida and Bermuda to Brazil, including the northern and western Gulf of Mexico, the Bahamas, and the Antilles. It is associated with clear water over reefs. Food consists of sponges. Maximum known size is 460 mm TL. references: Böhlke and Chaplin 1968; Randall 1964, 1968a, 1996; Sonnier et al. 1976; Hoese and Moore 1977, 1998; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Matsuura 2002c.

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908 TETRAODONTIFORMES

Cantherhines pullus (Ranzani, 1842) Orangespotted filefish

Cantherhines pullus is moderately deep bodied, with a deep groove between dorsal fins and an external spine on pelvic bone. Snout is slightly concave. Jaw teeth are acutely pointed; those in upper jaw are in two rows, with three teeth in outer row and two teeth in inner row, and those in lower jaw are in a single row of three teeth. Gill rakers on first arch number 34 to 46. Gill slit is located in front of pectoral fin base and is relatively long. Measurements are expressed as percent of SL: head length 30%– 32%, snout length 26%–27%, eye diameter 6%–7%, pectoral fin length 12%–13%, body depth at origin of anal fin 40%– 43%. Pectoral fin usually has 12 to 14 rays. Second dorsal fin has 33 to 36 rays. Pelvic spine is short and stout, has denticles on ventral surface and spinules along both sides, and is not movable in an anteroposterior direction. Anal fin has 29 to 32 rays. Scales bear one to several spines, and spines are branched distally. Caudal peduncle lacks enlarged scales, but mature males have setose patch (scales bearing long and very

slender spines) just anterior to caudal peduncle. Color is variable but commonly dark olive brown, with two or more light stripes on body and small orange spots on side of head. Caudal peduncle has white spots on dorsal and ventral midlines. This species occurs in the tropical and warm temperate Atlantic. In the western Atlantic it occurs from Massachusetts and Bermuda to southeastern Brazil, including the northern and western Gulf of Mexico and the Greater and Lesser Antilles. It is associated with reefs. Juveniles are pelagic and often are found in oceanic waters. Food consists of algae, sea grasses, sponges, tunicates, and bryozoans. Maximum known size is 200 mm TL.

references: Longley and Hildebrand 1941; Berry and Vogele 1961 (as Amanses pullus); Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; SmithVaniz et al. 1999; Matsuura 2002c.

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MONACANTHIDAE 909

Monacanthus ciliatus Fringed filefish

(Mitchill, 1818)

Monacanthus ciliatus is rectilinear in profile, with a prominent pelvic spine and no groove behind dorsal fin spines. Snout distinctly projects beyond remainder of head and is equally concave dorsally and ventrally. Jaw teeth are acutely pointed; those in upper jaw are in two rows, with three teeth in outer row and two teeth in inner row, and those in lower jaw are in a single row of three teeth. Gill opening is moderately oblique, slightly longer than pectoral fin base, and located above pectoral fin base. Gill rakers on first arch number 15 to 23. Measurements are expressed as percent of SL: head length 29%–39%, snout length 22%–26%, eye diameter 7%–15%, body depth 39%–55%. Pectoral fin usually has 11 rays. First dorsal spine is denticulate along anterior and lateral surfaces and bears retrorse spines along posterior surface. Second dorsal fin has 29 to 37 rays. Pelvic spine is distinct and flexible at base. Anal fin has 28 to 36 rays. Scales bear one or more spines, and the number of spines increases with growth. Adult males

have two pairs of enlarged spines and two pairs of smaller spines associated with each large pair on side of caudal peduncle. Color is tan, olive, or greenish, with broad, dark irregular stripes extending behind eye and behind gill opening. This species occurs in the tropical and warm temperate Atlantic Ocean to about 50 m. In the western Atlantic it occurs in shallow water from Newfoundland and Bermuda to Argentina, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is associated with grass beds, including turtle-grass beds. Juveniles are associated with pelagic Sargassum. Food consists of algae, organic detritus, and zooplankton. Maximum known size is 200 mm TL.

references: Longley and Hildebrand 1941; Berry and Vogele 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999; Matsuura 2002c.

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910 TETRAODONTIFORMES

Monacanthus tuckeri Bean, 1906 Slender filefish

Monacanthus tuckeri is rectilinear in profile, with a prominent pelvic spine and no groove behind dorsal fin spines. Snout distinctly projects beyond remainder of head and is equally concave dorsally and ventrally. Jaw teeth are acutely pointed; those in upper jaw are in two rows, with three teeth in outer row and two teeth in inner row, and those in lower jaw are in a single row of three teeth. Gill opening is moderately oblique, slightly longer than pectoral fin base, and located above pectoral fin base. Gill rakers on first arch number 19 to 24. Measurements are expressed as percent of SL: head length 33%– 42%, snout length 25%–28%, eye diameter 9%–14%, body depth 31%– 39%. Pectoral fin usually has 11 rays. First dorsal spine is denticulate along anterior and lateral surfaces and bears retrorse spines along posterior surface. Second dorsal fin has 32 to 37 rays. Pelvic spine is distinct and flexible at base. Anal fin has 31 to 36 rays. Scales bear

one to eight spines, and the number of spines increases with growth. Adult males have two pairs of enlarged spines on each side of caudal peduncle. Color is tan, olive, or greenish, with small yellowish brown to brown spots and a white reticular pattern on body. Spots behind eye are densely concentrated and form dark band. This species occurs in the western Atlantic from North Carolina and Bermuda to southern Florida and the Antilles, including the eastern Gulf of Mexico and the Bahamas. In the Gulf of Mexico it is known from southwestern Florida and the Dry Tortugas. This species is associated with gorgonian coral on coral reefs. Maximum known size is 90 mm TL. references: Longley and Hildebrand 1941; Berry and Vogele 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986; Smith 1997; Matsuura 2002c.

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MONACANTHIDAE 911

Stephanolepis hispidus (Linnaeus, 1766) Planehead filefish

Stephanolepis hispidus is rectilinear in profile, with a prominent pelvic spine and no groove behind dorsal fin spines. Snout moderately projects beyond remainder of head and is more concave dorsally than ventrally. Jaw teeth are acutely pointed; those in upper jaw are in two rows, with three teeth in outer row and two teeth in inner row, and those in lower jaw are in a single row of three teeth. Gill opening is moderately oblique, slightly longer than pectoral fin base, and located above pectoral fin base. Gill rakers on first arch number 26 to 33. Measurements are expressed as percent of SL: head length 30%– 41%, snout length 14%–28%, eye diameter 7%–17%, body depth 43%– 66%. Pectoral fin usually has 12 to 14 rays. First dorsal spine is denticulate along anterior and lateral surfaces and bears two rows of barbs along posterior surface. Second dorsal fin has 29 to 35 rays, with first ray filamentous in mature males. Pelvic spine is distinct and flexible at base. Anal fin has 30 to

35 rays. Scales bear one or more spines, the number of spines increases with growth, and spines are branched in specimens longer than 30 mm SL. Adult males lack enlarged spines on caudal peduncle. Color is tan, brown, or greenish, with dark blotches, spots, or lines. Bases of second dorsal fin and anal fin are usually dark. This species occurs in the tropical and warm temperate Atlantic. In the western Atlantic it occurs in shallow water from Nova Scotia and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. Maximum known size is 250 mm TL.

references: Longley and Hildebrand 1941 (as Monacanthus hispidus); Berry and Vogele 1961; Hoese and Moore 1977, 1998 (as M. hispidus); Robins and Ray 1986 (as M. hispidus); Boschung 1992 (as M. hispidus); Schaldach et al. 1997 (as M. hispidus); Smith 1997; Vega-Cendejas et al. 1997 (as M. hispidus); Smith-Vaniz et al. 1999; Matsuura 2002c.

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912 TETRAODONTIFORMES

Stephanolepis setifer Pigmy filefish

(Bennett, 1831)

Stephanolepis setifer is rectilinear in profile, with a prominent pelvic spine and no groove behind dorsal fin spines. Snout moderately projects beyond remainder of head and is more concave dorsally than ventrally. Jaw teeth are acutely pointed; those in upper jaw are in two rows, with three teeth in outer row and two teeth in inner row, and those in lower jaw are in a single row of three teeth. Gill opening is moderately oblique, slightly longer than pectoral fin base, and located above pectoral fin base. Gill rakers on first arch number 22 to 31. Measurements are expressed as percent of SL: head length 31%– 40%, snout length 18%–27%, eye diameter 8%–16%, body depth 47%– 60%. Pectoral fin usually has 11 to 13 rays. First dorsal spine is denticulate along anterior and lateral surfaces and bears two rows of barbs along posterior surface. Second dorsal fin has 27 to 29 (rarely 30) rays, with first ray filamentous in mature males. Pelvic spine is distinct and flexible at base. Anal

fin has 27 to 29 (rarely 26 or 30) rays. Scales bear one or more spines, the number of spines increases with growth, and spines are branched in specimens longer than 30 mm SL. Adult males lack enlarged spines on caudal peduncle. Color is tan, brown, or greenish, with short horizontal dark streaks or rows of small dark spots. Caudal fin has two dark bars. This species occurs in the western Atlantic from North Carolina and Bermuda to northern South America, including the Gulf of Mexico, the Bahamas, and the Antilles. It is associated with sea-grass beds. Juveniles are associated with floating Sargassum and flotsam. Maximum known size is 190 mm TL.

references: Berry and Vogele 1961; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998 (as M. setifer); Robins and Ray 1986 (as M. setifer); Schaldach et al. 1997 (as M. setifer); Smith 1997; Smith-Vaniz et al. 1999; Matsuura 2002c.

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913

OSTRACIIDAE

Cowfishes

Cowfishes are ellipsoid to subellipsoid in profile and triangular, square, or pentagonal in cross section, with body encased in bony plates through which eyes, mouth, and fins protrude. Nostril is paired and located in a shallow depression in front of eye. Mouth is terminal, lips are fleshy, and upper jaw is not protractile. Jaw teeth are conical or incisor-like and number three to eight in upper and lower jaws. Opercular opening is small, slitlike, and located in front of pectoral fin base. Branchiostegal rays number five or six. Pectoral fin is fanlike and inserts slightly below midflank. Dorsal and anal fins are similar in shape and size and are located posteriorly near or behind bony carapace. Pelvic fin is absent. Caudal fin is truncate or slightly convex. Carapace consists of hexagonal or polygonal bony scale plates. Scale plates often have granulations forming patterns and are often elongated into spines over eye, on posterolateral angle of disc, or along dorsal angles of body. Maximum known size is 450 mm TL. Cowfishes occur worldwide in tropical and warm temperate seas from near shore to 100 m. They are benthic fishes associated with coral and rocky reefs, sea-grass beds, or sponge-covered bottoms. Food consists of benthic invertebrates. There are 33 species in 14 genera, and 5 species in 3 genera occur in the western Central Atlantic and the Gulf of Mexico. references: Smith 1986; Tortonese 1986c; Nelson 1994; Matsuura 2001c, 2002d.

Key to the Species of the Gulf of Mexico (Adapted from Böhlke and Chaplin 1968) 1a. Spine projecting forward above each eye . . . . . . . . . . . . . 2 1b. No spine above eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2a. Pectoral fin with 12 or (rarely) 13 rays; body patterned with dark spots or blotches and irregular wavy lines; cheek with horizontal parallel lines . . . . . . . . . . . . . . . . . Acanthostracion quadricornis p. 915 2b. Pectoral fin with 13 or (rarely) 12 rays; body patterned with hexagonal or nearly hexagonal markings; cheek with reticular pattern . . . . . . Acanthostracion polygonius p. 914 3a. Carapace incomplete behind dorsal fin base, with an isolated large scaly plate in open area behind dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . Lactophrys trigonus p. 916 3b. Carapace complete behind dorsal fin base, forming a bony bridge over caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . 4 4a. A spine on posterolateral edge of carapace; body with many dark brown or blackish spots . . . . . . . . . . . . . . . . . . . . . . Rhinesomus bicaudalis p. 917 4b. No spine on carapace; body with many white, straw, or golden spots. . . . . . . . . . . . . . Rhinesomus triqueter p. 918

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914 TETRAODONTIFORMES

Acanthostracion polygonius Honeycomb cowfish

Poey, 1876

Acanthostracion polygonius has a spine anterior to eye and a spine on each side of anal fin, and bony armor is complete behind dorsal fin base. Mouth is low on head. Jaw teeth are incisor-like, with sharp-pointed cusps, and arranged in a single row. Anterior naris is at end of a moderately short tube, and posterior naris is at end of a short tube. Eye projects slightly above dorsal margin of head, and preorbital spine is well developed. Gill opening is slightly longer than pectoral fin base. Gill rakers on first arch number 11 to 14. Measurements are expressed as percent of SL: head length 25%– 27%, snout length 20%–23%, eye diameter 7%– 8%, pectoral fin length 17%–19%, maximum body depth 43%– 47%. Pectoral fin has 12 or 13 (usually 13) rays. Dorsal, anal, and caudal fins each have 10 rays. Carapace mostly consists of hexagonal bony plates, and plates are patterned with small, concentrically arranged tubercles and lines of tubercles radiating from geometric centers on plates. Ventral

ridge of carapace bears spine extending beyond level of anal fin origin. Color is olivaceous, with dark reticulate pattern on head and dark polygonal or hexagonal rings on bony plates on body. This species occurs in shallow water in the western Atlantic from New Jersey and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. In the Gulf of Mexico it occurs off Florida, 2516N, 8337W. This species is primarily associated with reefs. Food consists of sponges, tunicates, alcyonarians, and shrimps. Maximum known size is 480 mm TL.

references: Tyler 1965 (as Lactophrys polygonius); Böhlke and Chaplin 1968 (as L. polygonius); Randall 1968a, 1996 (as L. polygonius); Robins and Ray 1986 (as L. polygonia); Smith 1997 (as L. polygonius); Smith-Vaniz et al. 1999 (as L. polygonius); Matsuura 2002d.

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OSTRACIIDAE 915

Acanthostracion quadricornis Scrawled cowfish

(Linnaeus, 1758)

Acanthostracion quadricornis has a spine anterior to eye and a spine on each side of anal fin, and bony armor is complete behind dorsal fin base. Mouth is low on head. Jaw teeth are incisor-like, with sharp-pointed cusps, and arranged in a single row. Anterior naris is at end of a moderately short tube, and posterior naris is at end of a short tube. Eye projects slightly above dorsal margin of head, and preorbital spine is well developed. Gill opening is slightly longer than pectoral fin base. Gill rakers on first arch number 13 to 17. Measurements are expressed as percent of SL: head length 24%– 27%, snout length 20%–24%, eye diameter 8%–9%, pectoral fin length 18%–20%, maximum body depth 43%– 47%. Pectoral fin usually has 12 rays. Dorsal, anal, and caudal fins each have 10 rays. Carapace consists mostly of hexagonal bony plates, and plates are densely covered with small tubercles. Ventral ridge of carapace bears spine extending to anus. Carapace often bears posteriorly directed spines be-

hind dorsal and anal fin insertions. Color is grayish brown to yellow, with blue spots, bars, and reticulations on head and body. This species occurs in shallow water in the western Atlantic from Massachusetts and Bermuda to southeastern Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. It is associated with sea-grass beds. Food consists of sponges, colonial sea anemones, hermit crabs and other crustaceans, and tunicates. Maximum known size is 480 mm TL.

references: Longley and Hildebrand 1941 (as Lactophrys tricornis); Tyler 1965; Böhlke and Chaplin 1968; Randall 1968a, 1996; Hoese and Moore 1977, 1998; Castro-Aguirre and MárquezEspinoza 1981; Matsuura 1983v, 2002d; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999 (all above except Hildebrand 1941 as L. quadricornis).

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916 TETRAODONTIFORMES

Lactophrys trigonus Trunkfish

(Linnaeus, 1758)

Lactophrys trigonus has a spine on each side of anal fin, and bony armor is incomplete behind dorsal fin base. Mouth is low on head. Jaw teeth are incisor-like, with sharp-pointed cusps, and arranged in a single row. Anterior naris has a thick flap on posterior margin, and posterior naris is at end of a short tube. Eye projects slightly above dorsal margin of head, and preorbital spine is lacking. Gill opening is slightly longer than pectoral fin base. Gill rakers on first arch number 17 or 18. Measurements are expressed as percent of SL: head length 27%–33%, snout length 23%–25%, eye diameter 8%–13%, pectoral fin length 17%–21%, maximum body depth 48%–58%. Pectoral fin has 11 to 13 rays. Dorsal, anal, and caudal fins each have 10 rays. Carapace is patterned with small, concentrically arranged tubercles. Ventral ridge of carapace bears a spine extending to level of anus. Carapace is

usually incomplete behind dorsal fin but is followed by one or two separate bony plates. Color is greenish to tannish, with small, diffuse white spots. Hexagonal plates behind base of pectoral fin, at mid-distance between gill opening and end of carapace, are dark edged, forming chainlike markings. This species occurs in the western Atlantic from Massachusetts and Bermuda to Brazil, including the northern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with sea-grass beds. Food consists of polychaetes, sea urchins, bivalve mollusks, crabs, and sessile tunicates. Maximum known size is 450 mm TL references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986; Boschung 1992; Smith 1997; Smith-Vaniz et al. 1999; Matsuura 2002d.

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OSTRACIIDAE 917

Rhinesomus bicaudalis Spotted trunkfish

(Linnaeus, 1758)

Rhinesomus bicaudalis has a spine present on each side of anal fin, and bony armor is complete behind dorsal fin base. Anterior naris has a thick flap on posterior margin, and posterior naris is at end of a short tube. Eye projects slightly above dorsal margin of head, and preorbital spine is lacking. Mouth is low on head. Jaw teeth are incisor-like, with sharp-pointed cusps, and arranged in a single row. Gill opening is slightly longer than pectoral fin base. Gill rakers on first arch number 12 to 14. Measurements are expressed as percent of SL: head length 33%–34%, snout length 25%–28%, eye diameter 10%–11%, pectoral fin length 19%–20%, maximum body depth 49%–53%. Pectoral fin has 12 rays. Dorsal, anal, and caudal fins each have 10 rays. Carapace is patterned with small, concentrically arranged tu-

bercles. Ventral ridge of carapace bears spine extending to level of anal fin origin. Color is whitish, with small dark spots covering head, body, and caudal fin. Lips are whitish, and bases of pectoral, dorsal, and anal fins are dark. This species occurs in shallow water in the western Atlantic from Florida to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. It is found in the northern and southern Gulf of Mexico. Food consists of tunicates, sea cucumbers, brittle stars, and sea urchins. Maximum known size is 480 mm TL. references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997 (all above as Lactophrys bicaudalis); Matsuura 2002d.

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918 TETRAODONTIFORMES

Rhinesomus triqueter Smooth trunkfish

(Linnaeus, 1758)

Rhinesomus triqueter lacks spines on body, and bony armor is complete behind dorsal fin base. Mouth is low on head. Jaw teeth are incisor-like, with sharp-pointed cusps, and arranged in a single row. Anterior naris has a thick flap on posterior margin, and posterior naris is at end of a short tube. Eye projects slightly above dorsal margin of head. Gill opening is slightly longer than pectoral fin base. Gill rakers on first arch number eight or nine. Measurements are expressed as percent of SL: head length 30%–36%, snout length 23%–27%, eye diameter 8%–13%, pectoral fin length 17%–22%, maximum body depth 49%– 64%. Body becomes proportionally more slender with growth. Pectoral fin has 12 rays. Dorsal, anal, and caudal fins each have 10 rays. Carapace is patterned with small, concentrically ar-

ranged tubercles. Color is pale yellow to grayish, with dense covering of yellow spots on side of head and white spots on remainder of body. Mouth, dorsal fin base, and proximal and distal margins of caudal peduncle are blackish. This species occurs in shallow water in the western Atlantic from Massachusetts and Bermuda to Brazil, including the northern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with coral reefs. Food consists of algae, sponges, polychaetes, shrimps, and crabs. Maximum known size is 300 mm TL. references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986 (all above as Lactophrys triqueter); Matsuura 2002d.

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919

TETRAODONTIDAE

Puffers

Puffers are moderately slender and robust when uninflated, with jaw teeth forming dental plates and body either naked or covered by fine spinules. Head is large and blunt. Nostril is either paired or single. Eye is moderately large and located near dorsal margin of head. Mouth is terminal to slightly superior. Upper and lower dental plates have median suture and are fused to jaw bones. Gill opening is a short slit in front of pectoral fin. Pectoral fin is fanlike, inserts on midflank or above midflank, and has nearly vertical base. Dorsal and anal fins are similar in shape and size, possess 7 to 18 rays, and are located posteriorly. Pelvic fin and girdle are absent. Caudal fin is moderately forked to convex and has 10 principal rays. Skin is very tough and elastic. Spinules are present on back, belly, and flank in some species. Lateral line is present or absent, and when present, it often forms interconnected patterns on head and body. Stomach possesses ventral diverticulum that enables fish to swallow water or air and inflate to large size. Ribs and epineural bones are absent. Vertebrae number 17 to 29. Maximum known size is 80 cm TL. Puffers occur worldwide in tropical to warm temperate seas, generally at shallow depths. Most species are marine, but some enter fresh waters with access to the sea, and a few live permanently in freshwater. The skin, liver, and ovaries of most species contain alkaloid poisons (tetrodotoxin) that can be lethal to animals, including humans. Some species excrete poison through the skin. There are 121 species in 19 genera, 16 species in 4 genera in the western Central Atlantic, and 9 species in 3 genera in the Gulf of Mexico. references: Tortonese 1986d; Nelson 1994; Smith and Heemstra 1986b; Smith 1997; Matsuura 2001d; Shipp 2002.

Key to the Species of the Gulf of Mexico (Adapted from Böhlke and Chaplin 1968; Shipp and Yerger 1969a; Shipp 2002) 1a. Dorsal fin and anal fins each with 12 to 15 rays . . . . . . . . . . . . . . . . . . . . . Lagocephalus laevigatus p. 923 1b. Dorsal and anal fins each with fewer than 11 rays . . . . . . 2 2a. Dorsal fin with 9 or 10 rays . . . . . . . . . . . . . . . . . . . . . . . 3 2b. Dorsal fin with 6 to 9 rays . . . . . . . . . . . . . . . . . . . . . . . . 4 3a. Caudal fin with distinct dark dorsal and ventral margins; dorsal fin base without black spot . . . . . . . . . . . . . . . . . . . . . . . . Canthigaster rostrata p. 922 3b. Caudal fin base without distinct dark dorsal and ventral margins, dorsal fin base with black spot . . . . . . . . . . . . . . . . . . . . . Canthigaster jamestyleri p. 921

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920 TETRAODONTIFORMES

4a. Dorsal surface of body with small, fleshy tabs or lappets, either one pair behind eye or many on posterior part of body. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4b. Dorsal surface of body without small, fleshy lappets. . . . . 6 5a. Single black lappet behind eye . . . . . . . . . . . . . . . . . . . . . . . . Sphoeroides dorsalis p. 924 5b. Many tan lappets on posterior dorsal part of body . . . . . . . . . . . . . . . . . . . . . . . Sphoeroides spengleri p. 928 6a. Body uniformly darkly pigmented except for a few spots; body smooth . . . . . . . . . . . Sphoeroides pachygaster p. 926 6b. Body mottled; body with prickles, at least dorsally. . . . . . 7 7a. One or two distinct interorbital bars; often with white circular markings behind eye . . . . . . . . . . . . . . . . . . . . . Sphoeroides testudineus p. 929 7b. One indistinct interorbital bar; no white circular markings behind eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8a. Axil of pectoral fin with distinct dark spot . . . . . . . . . . . . . . . . . . . . . . . Sphoeroides nephelus p. 925 8b. Axil of pectoral fin without distinct dark spot . . . . . . . . . . . . . . . . . . . . . . . . . Sphoeroides parvus p. 927

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TETRAODONTIDAE 921

Canthigaster jamestyleri Goldface toby

Moura and Castro, 2002

Canthigaster jamestyleri has minute nostrils, an indistinct lateral line, and a dark spot on dorsal fin base. Snout is relatively long and very acute. Nostril is single and surrounded by a fleshy rim. Interorbital region is concave. Eye is located slightly above dorsal margin of head. Gill opening is arch shaped, located just anterior to pectoral fin base, and about half the length of pectoral fin base. Measurements are expressed as percent of SL: head length 36%– 39%, uninflated body depth 32%– 40%; and as percent of head length: snout length 56%– 63%, eye diameter 21%–28%, interorbital width 24%–27%, pectoral fin length 33%– 40%. Pectoral fin has 15 or 16 rays. Dorsal fin is anterior to and larger than anal fin and has

9 rays. Anal fin has 9 rays. Caudal fin is truncate to slightly convex. Small spinules are located on lower part of head and abdomen and have two lateral roots that fold into small sockets. Color is tan, darker dorsally than ventrally, with roundish spots and irregular diagonal lines ventral to pectoral fin base. Side of head and snout are marked with horizontal lines, and two stripes run along body. This species occurs in the western Atlantic from South Carolina to the northern Gulf of Mexico. It is apparently restricted to depths greater than 90 m and is likely associated with hard substrates. reference: Moura and Castro 2002.

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922 TETRAODONTIFORMES

Canthigaster rostrata Sharpnose puffer

(Bloch, 1786)

Canthigaster rostrata has minute nostrils, an indistinct lateral line, and a fleshy keel on back. Snout is relatively long and very acute. Nostril is single and surrounded by a fleshy rim. Interorbital region is concave. Eye is located slightly above dorsal margin of head. Gill opening is arch shaped, located just anterior to pectoral fin base, and about half the length of pectoral fin base. Gill rakers on first arch number six or seven. Measurements are expressed as percent of SL: head length 36%– 41%, snout length 23%–27%, eye diameter 9%– 11%, interorbital width 10%–12%, pectoral fin length 14%–16%, uninflated body depth 32%– 40%. Pectoral fin has 16 to 18 rays. Dorsal fin is anterior to and larger than anal fin and has 10 rays. Anal fin has 9 rays. Caudal fin is truncate to slightly convex. Small spinules are located on lower part of head and abdomen and have two lateral roots that fold into small sockets. Color is brown dorsally, with faint blue spots, and white with a bluish tint ventrally. Upper and lower margins of caudal pe-

duncle and caudal fin are black, and lower margin of caudal peduncle is marked with wavy or diagonal blue lines. This species occurs in the tropical and warm temperate Atlantic. In the western Atlantic it occurs in shallow water from North Carolina and Bermuda to northern South America, including the northern and southern Gulf of Mexico, the Bahamas, and the Antilles. It is associated with coral reefs. Food consists of algae, sea grasses, sponges, hydroids, polychaetes, sea urchins, starfishes, and crabs and other crustaceans. Maximum known size is 110 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Castro-Aguirre and MárquezEspinoza 1981; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; SmithVaniz et al. 1999; Shipp 2002.

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TETRAODONTIDAE 923

Lagocephalus laevigatus Smooth puffer

(Linnaeus, 1766)

Lagocephalus laevigatus has falcate dorsal and anal fins and an emarginate caudal fin. Snout is blunt. Interorbital space is broad and flat. Nostril is paired, and nares are located in an oval depression separated from surrounding integument by a shallow groove. Gill slit is arch shaped. Gill rakers are very short and number 10 to 12. Measurements are expressed as percent of SL: head length 31%–35%, snout length 14%–16%, eye diameter 8%–9%, pectoral fin length 18%–20%, uninflated body depth 22%–23%. Pectoral fin has 17 or 18 rays (rarely 15, 16, or 19). Dorsal fin has 13 or 14 (rarely 15) rays. Anal fin is similar in shape to dorsal fin and has 12 or 13 rays. Dorsal lobe of caudal fin is longer than ventral lobe. Most of body is covered with spinules that possess three well-developed basal prongs. Spines on

chin and belly have a single basal prong. A fleshy fold extends from pectoral fin base to caudal fin base. Color is dark green to brownish gray dorsally and white gray to silvery ventrally. Juveniles have three or four dark saddles. This species occurs in the tropical to warm temperate Atlantic Ocean. In the western Atlantic it occurs from Massachusetts to Brazil, including the Gulf of Mexico and the Bahamas. Juveniles occur near shore and on offshore banks, and adults are pelagic over continental shelves. Maximum known size is 600 mm TL. references: Shipp 1974, 2002; Hoese and Moore 1977, 1998; Matsuura 1983w; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999.

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924 TETRAODONTIFORMES

Sphoeroides dorsalis Marbled puffer

Longley, 1934

Sphoeroides dorsalis has a pair of lappets on dorsal surface behind eyes. Snout is long and acute. Nostril is paired, and nares are at ends of short tubes. Interorbital space is narrow and slightly concave. Gill slit is arch shaped and is slightly shorter than pectoral fin base. Gill rakers are very short and number 9 to 12. Measurements are expressed as percent of SL: head length 32%– 40%, snout length 19%–25%, eye diameter 6%–10%, pectoral fin length 13%–17%, uninflated body depth 21%–24%. Pectoral fin has 15 to 17 rays. Dorsal fin is narrow based and has 8 rays. Anal fin is similar in shape to dorsal fin and has 7 rays. Body is covered with small spinules except on dorsum posterior to lappets, on cheeks, and on belly anterior to pectoral fin base. Color is uniformly gray to brown dorsally, with a diffuse bar over

interorbital region and one or two diffuse blotches on head behind orbit, and white ventrally, with one to five diffuse spots on lower flank behind pectoral fin base. Lappets on back are dark in adults but may be tan in juveniles. This species occurs between 18 and 90 m in the western Atlantic from North Carolina and the Bahamas to northern Suriname, including the northern and southern Gulf of Mexico. Juveniles apparently occur near shore. Maximum known size is 200 mm TL. references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Shipp 1974, 2002; Hoese and Moore 1977, 1998; Boschung 1992; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999.

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TETRAODONTIDAE 925

Sphoeroides nephelus Southern puffer

(Goode and Bean, 1882)

Sphoeroides nephelus has a light reticular pattern on upper half of body and lacks lappets. Snout is long and acute. Nostril is paired, and nares are located at ends of short tubes. Interorbital space is narrow and slightly concave. Gill slit is arch shaped and slightly shorter than pectoral fin base. Gill rakers are very short and number eight or nine. Measurements are expressed as percent of SL: head length 34%– 39%, snout length 18%–25%, eye diameter 5%–9%, pectoral fin length 13%–18%, uninflated body depth 22%–30%. Pectoral fin has 13 to 15 rays. Dorsal fin is narrow based and has 7 to 9 rays. Anal fin is similar in shape to dorsal fin and has 6 or 7 rays. Body varies from naked to nearly covered with spinules anterior to dorsal fin base. Color is dark olive gray dorsally, often with pale gray rings, and

white ventrally. Axil of pectoral fin has a black blotch, and sides behind pectoral fin are often patterned with dark gray spots. Upper section of cheek is often patterned with dark slashes. This species occurs in shallow water, including bays and estuaries, in the western Atlantic from northeastern Florida and the Bahamas to Yucatán and the Lesser Antilles, including the eastern Gulf of Mexico off Florida and the west coast of Yucatán. Maximum known size is 250 mm TL.

references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Shipp and Yerger 1969b; Shipp 1974, 2002; Hoese and Moore 1977, 1998; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999.

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926 TETRAODONTIFORMES

Sphoeroides pachygaster Blunthead puffer

(Müller and Troschel, 1848)

Sphoeroides pachygaster lacks spinules and is uniformly colored. Snout is long and blunt. Nostril is paired, and nares are located at ends of short tubes. Interorbital space is narrow and slightly concave. Gill slit is arch shaped and slightly shorter than pectoral fin base. Gill rakers are very short. Measurements are expressed as percent of SL: head length 36%– 46%, snout length 17%–22%, eye diameter 8%–13%, interorbital width 6%–12%, pectoral fin length 13%–16%. Pectoral fin has 14 to 17 (usually 14 to 16) rays. Dorsal fin is narrow based and has 8 to 10 (usually 9) rays. Anal fin is similar in shape to dorsal fin and has 7 to 9 (usually 8

or 9) rays. Color is dark olive gray dorsally and white ventrally. Adults may have dark brown spots and blotches. This species occurs in the western Atlantic and possibly in the eastern Atlantic and Indian Oceans. In the western Atlantic it occurs between 55 and 180 m, from South Carolina to Argentina, including the Gulf of Mexico. It is apparently a benthic species. Maximum known size is 250 mm TL. references: Shipp and Yerger 1969a; Shipp 1974, 2002; Hoese and Moore 1977; Matsuura 1983w; Robins and Ray 1986; Boschung 1992; Smith 1997.

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TETRAODONTIDAE 927

Sphoeroides parvus Least puffer

Shipp and Yerger, 1969

Sphoeroides parvus has irregularly placed spots on lower flank and lacks lappets. Snout is short and acute. Nostril is paired, and nares are located at ends of short tubes. Interorbital space is relatively broad and slightly flat. Gill slit is arch shaped and slightly shorter than pectoral fin base. Gill rakers are very short and number seven or eight. Measurements are expressed as percent of SL: head length 33%– 39%, snout length 17%–19%, eye diameter 6%–11%, interorbital width 4%– 6%, pectoral fin length 15%–21%, uninflated body depth 15%–24%. Pectoral fin has 13 to 16 (usually 14 or 15) rays. Dorsal fin is narrow based and has 8 or 9 (usually 8) rays. Anal fin is similar in shape to dorsal fin and has 6 to 8 rays. Spinules occur dorsally from nasal tube

to dorsal fin origin, laterally from cheek to dorsal fin origin, and ventrally from chin to anus. Color is dark olive gray dorsally, with pale green spots and usually dark brown spots and blotches, and white ventrally. If pectoral fin axil is darkly pigmented, it is no darker than spots or blotches on sides. This species occurs in the western Atlantic from the Gulf of Mexico in coastal waters, bays, and estuaries. It is known from Pensacola, FL, to Yucatán. Maximum known size is 150 mm TL. references: Shipp and Yerger 1969a; Shipp 1974, 2002; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Smith 1997; CastroAguirre et al. 1999.

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928 TETRAODONTIFORMES

Sphoeroides spengleri Bandtail puffer

(Bloch, 1785)

Sphoeroides spengleri has a regular series of round dark spots on lower flank and two dark bands on caudal fin. Snout is long and acute. Nostril is paired, and nares are located at ends of short tubes. Interorbital space is narrow and moderately concave. Gill slit is arch shaped and slightly shorter than pectoral fin base. Gill rakers are very short and number 8 to 10. Measurements are expressed as percent of SL: head length 33%–39%, snout length 17%–22%, eye diameter 5%–9%, interorbital width 4%– 6%, pectoral fin length 10%–14%, uninflated body depth 19%–22%. Pectoral fin has 13 or 14 (usually 13) rays. Dorsal fin has 7 or 8 (usually 8) rays. Anal fin is similar in shape to dorsal fin and has 6 to 8 rays. Spinules occur ventrally anterior to anus, are present or absent dorsally from nape to dorsal fin base, and if present laterally, are limited to cheek. Color is dark brown dorsally, light tan laterally, and white ventrally, with a row of large, round dark

brown spots along lower side from chin to caudal fin base. Numerous small light brown lappets are present on posterior section of body. This species occurs in shallow water in the western Atlantic from Massachusetts and Bermuda to Brazil, including the Gulf of Mexico, the Bahamas, and the Antilles. It is associated with sea-grass beds and reef flats. Food consists of algae, sea grass, mollusks, polychaetes, sea urchins, brittle stars, and crabs and other crustaceans. Maximum known size is 180 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Shipp 1974, 2002; Hoese and Moore 1977, 1998; Shipp and Hopkins 1978; Castro-Aguirre and Márquez-Espinoza 1981; Boschung 1992; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; Castro-Aguirre et al. 1999; Smith-Vaniz et al. 1999.

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TETRAODONTIDAE 929

Sphoeroides testudineus Checkered puffer

(Linnaeus, 1758)

Sphoeroides testudineus has a light honeycomb pattern on dorsal surface and one or two light bars in interorbital region. Snout is long and acute. Nostril is paired, and nares are located at ends of short tubes. Interorbital space is moderately narrow and flat to slightly moderately concave. Gill slit is arch shaped and slightly shorter than pectoral fin base. Gill rakers are very short and number 8 to 10. Measurements are expressed as percent of SL: head length 33%–38%, snout length 16%–19%, eye diameter 5%–10%, interorbital width 6%– 8%, pectoral fin length 13%–17%, uninflated body depth 24%–27%. Pectoral fin has 13 to 16 (usually 14 or 15) rays. Dorsal fin has 7 to 9 rays. Anal fin is similar in shape to dorsal fin and has 6 to 8 rays. Spinules occur dorsally from interorbital region to dorsal fin origin, ventrally from chin to anus, and laterally in

broad band behind pectoral fin. Color is dark gray to olive dorsally, with a network of circular tan to yellowish lines forming a honeycomb pattern; tan laterally, with pupil-sized dark round spots; and white ventrally. This species occurs in coastal waters, bays, and estuaries in the western Atlantic from Rhode Island to southeastern Brazil, including the Florida Keys, the southern Gulf of Mexico, the Bahamas, and the Antilles. It is rare to absent in the northern Gulf of Mexico. This species is associated with sea-grass beds and is rare on coral reefs. Food consists of shellfishes. Maximum known size is 300 mm TL. references: Böhlke and Chaplin 1968; Shipp 1974, 2002; Hoese and Moore 1977, 1998; Matsuura 1983w; Schaldach et al. 1997; Smith 1997; Castro-Aguirre et al. 1999.

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930

DIODONTIDAE

Porcupinefishes

Porcupinefishes are depressed anteriorly, globular in shape, and greatly inflatable, with jaw teeth fused into beaks and large spinelike scales. Head is broad and blunt. Nostril is usually located in tentacle. Mouth is terminal and very small. Left and right premaxillae and dentaries are fused. Teeth of upper and lower jaws are fused together and to jaw bones, and sutures between teeth on each side of jaws are absent. Gill opening is slitlike and located immediately in front of pectoral fin base. Pectoral fin inserts on midflank and is fanlike. Dorsal fin is single and short based and inserts just anterior to caudal peduncle. Pelvic fin and pelvic girdle are absent. Anal fin is similar in size and shape to dorsal fin and is located beneath dorsal fin. Fin spines are absent. Caudal peduncle is rather deep. Caudal fin has truncate to convex posterior margin. Stomach possesses ventral diverticulum that enables fish to inflate to large size. Gas bladder is present. Lateral line is inconspicuous. Body and head are covered with large spinelike scales, with double or triple subdermal bases. Long spines have double bases and are erectile when body is inflated, and short spines have triple bases and are permanently erect. Vertebrae number 18 to 22. Maximum size is 100 cm TL. Porcupinefishes occur worldwide in tropical to warm temperate seas in shallow water. Species are benthic or associated with floating seaweed. Food consists of sea urchins, mollusks, and crustaceans. There are 19 species in six genera, 8 species in two genera in the western Central Atlantic, and 6 or 7 species in two genera in the Gulf of Mexico. references: Leis 1984, 2001, 2002; Tortonese 1986e; Nelson 1994.

Key to the Species of the Gulf of Mexico (Adapted from Böhlke and Chaplin 1968; Leis 2002) 1a. Spinelike scales long and with double subdermal bases; spines fold back when fish not inflated . . . . . . . . . . . . . . . 2 1b. Spinelike scales short and with triple subdermal bases; spines fixed in erect position. . . . . . . . . . . . . . . . . . . . . . . 4 2a. One or more spinelike scales exclusively located on dorsal surface of caudal peduncle; body without large dark dorsal blotches; fins (with possible exception of anal fin) spotted .............................................3 2b. No spinelike scales exclusively located on caudal peduncle; body with several large dark dorsal blotches; fins without dark spots . . . . . . . . . . . . . . . . Diodon holocanthus p. 937 3a. Pectoral fin rays 19 to 22; anal fin rays 16 to 18; dorsal and anal fins somewhat pointed in adults; head width of

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DIODONTIDAE 931

3b.

4a.

4b.

5a.

5b. 6a.

6b.

adults 25% to 30% of SL; spinelike scales between lower jaw and anus 10 to 14 . . . . . . . . . Diodon eydouxii p. 936 Pectoral fin rays 22 to 25 (rarely 21); anal fin rays 14 to 16; dorsal and anal fins rounded in adults; head width of adults 30% to 42% of SL; spinelike scales between lower jaw and anus 14 to 19 . . . . . . . . . . Dyodon hystrix p. 938 One or two spinelike scales exclusively on caudal peduncle; caudal fin rays normally 10; nostril consists of open ridged cup in adults; fins spotted in adults; some spinelike scales on top of head with four roots . . . . . . . . . . . . . . . . . . . . Chilomycterus reticulatus p. 934 No spines exclusively on caudal peduncle; caudal fin rays normally 9; nostril consists of tentacle with two openings; fins without spots in adults; all spinelike scales with three roots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Large tentacle above eye; color pattern on back and sides consisting of large blotches and small spots; fin bases with small spots . . . . . . . . . . . Chilomycterus antennatus p. 932 Tentacles above eye small or absent; no small spots on body or fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Color pattern on back and sides consists of network of hexagonal or circular black lines . . . . . . . . . . . . . . . . . . . . Chilomycterus antillarum p. 933 Color pattern on back and sides consists of wavy to nearly parallel nonintersecting lines . . . . . . . . . . . . . . . . . . . . . Chilomycterus schoepfii p. 935

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932 TETRAODONTIFORMES

Chilomycterus antennatus Bridled burrfish

(Cuvier, 1816)

Chilomycterus antennatus has moderately short, spinelike scales covering body, small black spots, and large dark blotches. Interorbital region is slightly concave near midline. Snout is strongly oblique, rather straight, and relatively short. Measurements are expressed as percent of SL: head length 38%– 44%, snout length 11%–14%, eye diameter about 9%, pectoral fin length 17%–18%, uninflated body depth about 39%. Pectoral fin has a truncate posterior margin and 23 to 26 rays. Dorsal fin has 11 to 13 rays, and anal fin has 11 or 12 rays. Spinelike scales are short and triangular, with triple subdermal bases. Two spines occur over orbit. Spine height is about half of eye diameter. Fleshy tentacles are located over eye and on sides and below mouth. Color is tan to light brown, with dorsal and lateral sides

of head and body covered with pupil-sized dark brown spots. Large dark blotches occur on dorsum over pectoral fin base and on base of dorsal fin. Fins are lightly pigmented. This species occurs in the tropical to warm temperate Atlantic. In the western Atlantic it occurs from near shore to 25 m, from southeastern Florida and the Bahamas to northern South America. It has not been recorded from the Gulf of Mexico, but it may occur there. This species is associated with coral reefs and grass beds. Food consists of hard-shelled invertebrates. Maximum known size is 230 mm TL.

references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Robins and Ray 1986; Smith 1997; Leis 2002.

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DIODONTIDAE 933

Chilomycterus antillarum Web burrfish

Jordan and Rutter, 1897

Chilomycterus antillarum has moderately short spinelike scales covering the body, large dark blotches, and dark lines demarcating celllike or hexagonal areas. Interorbital region is strongly concave. Snout is strongly oblique, rather straight, and relatively short. Measurements are expressed as percent of SL: head length 40%– 42%, snout length 15%–17%, eye diameter 13%–14%, pectoral fin length 19%–22%, uninflated body depth 31%–35%. Pectoral fin has a truncate posterior margin and 21 or 22 rays. Dorsal fin has 12 or 13 rays, and anal fin has 11 rays. Spinelike scales are moderately short and moderately slender, with triple subdermal bases. Spine height is about half of eye diameter. Fleshy tentacles are located over eye and below mouth. Tentacles accompany spines along sides of belly. Color is tan to light brown, with dorsal and lateral sides of head and body covered with dark lines form-

ing cells or hexagonal areas about half the size of eye. Large dark brown to black blotches occur above pectoral fin, behind pectoral fin base, and at base of dorsal fin. Light-colored rings surround all blotches. Ventral surface is tan anteriorly, grading to dark brown posteriorly. This species occurs from near shore to 44 m in the western Atlantic from North Carolina and Bermuda to Brazil, including the northeastern Gulf of Mexico, the Bahamas, and the Antilles. It has been recorded only once from Bermuda (Smith-Vaniz et al. 1999). Food consists of hard-shelled invertebrates. Maximum known size is about 250 mm SL. references: Böhlke and Chaplin 1968; Randall 1968a, 1996; Matsuura 1983x; Robins and Ray 1986; Boschung 1992; Smith 1997; Smith-Vaniz et al. 1999; Leis 2002.

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934 TETRAODONTIFORMES

Chilomycterus reticulatus Spotted burrfish

(Linnaeus, 1758)

Chilomycterus reticulatus has very short, spinelike scales covering the body and small black spots. Interorbital region is nearly horizontal. Snout is strongly oblique, rather straight, and relatively long. Nostril consists of an open ridged cup. Measurements are expressed as percent of SL: head length about 37%, snout length about 16%, eye diameter about 9%, pectoral fin length about 19%, uninflated body depth about 27%. Pectoral fin has a truncate posterior margin and 19 to 22 rays. Dorsal fin has 12 to 14 rays, and anal fin has 11 to 14 rays. Spinelike scales are short and triangular, with triple subdermal bases, but some spines on top of head have four bases. Spine height is about one-third of eye diameter. Three spines are located over orbit. Fleshy tentacles on head are lacking. Color is tan, with

dorsal and lateral sides of head and body covered with pupil-sized dark brown spots. Light brown gular band extends from ventral margin of eye and is continuous under chin. Fins are covered with smaller, regularly arranged dark brown spots. Pelagic-stage juveniles are blue, with dark spots on dorsal and lateral surfaces. This species occurs worldwide in tropical waters. In the western Atlantic it occurs from New Jersey to Brazil, including the western Gulf of Mexico. Larval and early juvenile stages (up to 200 mm TL) are pelagic, and large juveniles and adults are benthic in coastal waters to 100 m. Maximum known size is 750 mm TL. references: Robins and Ray 1986 (as Chilomycterus atinga); Tortonese 1986e (as C. atinga); Smith 1997(as C. atinga); Leis 2002.

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DIODONTIDAE 935

Chilomycterus schoepfii Striped burrfish

(Walbaum, 1792)

Chilomycterus schoepfii has short, erect, spinelike scales and a pattern of curved lines and dark blotches covering body. Interorbital region is strongly concave. Nares are located on either side of tip of tentacle in front of eye. Snout is strongly oblique, moderately concave, and relatively short. Measurements are expressed as percent of SL: head length 39%– 44%, snout length 15%–19%, eye diameter 9%–12%, pectoral fin length 20%–25%, uninflated body depth 30%–37%. Pectoral fin has a truncate posterior margin and 19 to 22 rays. Dorsal fin has 10 rays, and anal fin has 10 rays. Spinelike scales are moderately short and moderately slender, with triple subdermal bases. Spines running from front of head to origin of first dorsal fin number six. Maximum spine height is about half of eye diameter. Four to six fleshy tentacles are located on chin, and one fleshy tentacle is located above eye. Tentacles accompany spines along sides of belly. Color is yellowish brown dorsally and white to

yellowish ventrally, with dark blotches above and behind pectoral fin base, on base of dorsal fin, and ventral to dorsal fin base. Dorsal and lateral sides of body are patterned with narrow to moderately broad, wavy, and approximately parallel lines. This species occurs in shallow water in the western Atlantic from Nova Scotia and Bermuda to Brazil, including the entire Gulf of Mexico and the Bahamas. It is rare in the Bahamas and the Antilles. Larvae and juveniles to about 20 mm TL are pelagic, and large juveniles and adults are demersal and associated with sea-grass beds in protected waters, including bays and estuaries. Maximum known size is 280 mm TL. references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Vega-Cendejas et al. 1997; Smith-Vaniz et al. 1999; Leis 2002.

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936 TETRAODONTIFORMES

Diodon eydouxii Brissout de Barneville, 1846 Pelagic porcupinefish Diodon eydouxii is relatively slender and has relatively pointed dorsal and anal fins and relatively long, erectile, spinelike scales. Interorbital region is slightly concave. Nares are located on either side of tip of tentacle in front of eye. Snout is strongly oblique, moderately concave, and relatively short. Pectoral fin has a truncate posterior margin and 19 to 22 rays. Dorsal fin has 16 to 18 rays, and anal fin has 16 to 18 rays. Spinelike scales are long and slender, with double subdermal bases. Spines located mid-dorsally in front of head are shorter than those below or posterior to pectoral fin base. A small spine is located on dorsal aspect of caudal peduncle. Spines between lower

jaw and anus number 10 to 14. Tentacles are absent on body. Color is blue dorsally and white ventrally, with numerous small dark blue blotches on head and body and dark blue spots on bases of fins. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs in oceanic waters from North Carolina to northern South America, including the southern and western Gulf of Mexico. Food consists of zooplankton. Maximum known size is 250 cm TL.

reference: Leis 2002.

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DIODONTIDAE 937

Diodon holocanthus Balloonfish

Linnaeus, 1758

Diodon holocanthus has relatively long, erectile, spinelike scales and body patterned with round black spots and large black blotches. Interorbital region is moderately concave. Nares are located on either side of tip of tentacle in front of eye. Snout is strongly oblique, moderately concave, and relatively short. Measurements are expressed as percent of SL: head length 39%– 44%, snout length 14%–17%, eye diameter 10%–11%, pectoral fin length 17%–21%, uninflated body depth 34%–35%. Pectoral fin has a truncate posterior margin and 22 or 23 rays. Dorsal fin has 13 or 14 rays, and anal fin has 13 or 14 rays. Spinelike scales are long and slender, with double subdermal bases. Spines mid-dorsally in front of head are longer than those below or posterior to pectoral fin base. Spines along mid-dorsal line from snout to dorsal fin origin number 14 to 16. Tentacles are absent on body. Color is light brown dorsally and white ventrally, with a large dark bar above and below eye and on occipital region of head, a large dark blotch above pectoral fin

base, and a large dark blotch on base of dorsal fin. Dorsal section of body and posterior part of head are covered with small, round, dark spots. Fins are light yellow and lack spots. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs from near shore to 100 m, from Massachusetts to Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. Larvae and juveniles to 90 mm TL are pelagic, and large juveniles and adults are demersal and more common in the vicinity of mangrove areas than on reefs. Food consists of gastropods, crabs, other crustaceans, and sea urchins. Maximum known size is 300 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1996; Bright and Cashman 1974; Hoese and Moore 1977, 1998; Robins and Ray 1986; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Matsuura 2001e; Leis 2002.

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938 TETRAODONTIFORMES

Diodon hystrix Porcupinefish

Linnaeus, 1758

Diodon hystrix has relatively long, erectile, spinelike scales and body patterned with round black spots. Interorbital region is slightly concave. Nares are located on either side of tip of tentacle in front of eye. Snout is strongly oblique, moderately concave, and relatively short. Measurements are expressed as percent of SL: head length about 47%, snout length about 16%, eye diameter about 13%, pectoral fin length about 24%, body depth about 33%. Pectoral fin has a truncate posterior margin and 22 to 25 rays. Dorsal fin has 15 to 17 rays, and anal fin has 14 to 16 rays. Spinelike scales are long and slender, with double subdermal bases. One or two small spines are located on dorsal aspect of caudal peduncle. Spines located mid-dorsally in front of head are shorter than those below or posterior to pectoral fin base. Spines along mid-dorsal line from snout to dorsal fin origin number about 15 to 20, and from lower jaw to anus number 14 to 19. Ten-

tacles are absent on body. Color is olivaceous dorsally and white ventrally, with numerous small dark spots on head, body, and fins. This species occurs worldwide in tropical to warm temperate seas. In the western Atlantic it occurs from Massachusetts and Bermuda to southern Brazil, including the entire Gulf of Mexico, the Bahamas, and the Antilles. Larvae and juveniles to 200 mm TL are pelagic, and large juveniles and adults are demersal and associated with reefs. Food consists of sea urchins, mollusks, hermit crabs, and other crabs. Maximum known size is 750 mm TL.

references: Longley and Hildebrand 1941; Böhlke and Chaplin 1968; Randall 1968a, 1998; Hoese and Moore 1977, 1998; Robins and Ray 1986; Boschung 1992; Schaldach et al. 1997; Smith 1997; Smith-Vaniz et al. 1999; Matsuura 2001e; Leis 2002.

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939

MOLIDAE

Ocean sunfishes

Ocean sunfish are orbicular to elongate-oval shaped, compressed, and truncate posteriorly, with caudal fin absent and dorsal and anal fins high and short based. Eye is small. Mouth is very small and terminal. Teeth are united in each jaw to form beak but without median suture. Palatine teeth are absent. Gill opening is small and porelike. Pectoral fin is small to moderate, inserts slightly above midflank, and fits into concavity in side of body. Dorsal fin is high and has a short base and 15 to 21 rays. Pelvic fin is absent, and pelvic bone is poorly developed. Anal fin is similar in size and shape to dorsal fin and has 15 to 21 rays. Caudal fin is replaced by a leathery, rudderlike structure (clavus) supported by fin rays derived from dorsal and anal fins. Skin is rough, naked, spinous, or tessellated. Small juveniles are scattered with large spiny scales. Gas bladder is absent. Maximum size is 350 cm TL. Ocean sunfish occur worldwide in tropical and warm temperate seas. Species are pelagic and are often observed on the water surface partially on their sides. Food consists of jellyfishes, salps, ctenophores, eel larvae, and occasionally other ray-finned fishes. There are four or five species in three genera, and three species in three genera occur in the Gulf of Mexico. references: Gregory and Raven 1934; Raven 1939a,b; Fraser-Brunner 1951; Heemstra 1986r; Tortonese 1986f.

Key to the Species of the Gulf of Mexico (Adapted from Fraser-Brunner 1951; Heemstra 1986r) 1a. Body elongate-oval shaped; body depth 25% to 48% of SL; body surface smooth, thin, and composed of fused hexagonal plates; body obliquely truncate posteriorly; gill rakers free of skin . . . . . . . . . . . . . . . . . . . Ranzania laevis p. 942 1b. Body orbicular; body depth 55.6% to 125% of SL; body surface thick and tough, with small, rough denticles; body vertically truncate posteriorly; gill rakers concealed in thick skin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Body depth about 67% of body length; median rays of clavus (rudderlike structure forming modified caudal fin) forming pronounced lobe . . . . Masturus lanceolatus p. 940 2b. Body depth usually equal to SL; median rays of clavus not forming pronounced lobe . . . . . . . . . . . . Mola mola p. 941

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940 TETRAODONTIFORMES

Masturus lanceolatus Sharptail mola

(Liénard, 1840)

Masturus lanceolatus is orbicular, compressed, and truncate posteriorly, with clavus supported by vertebral and dorsal and anal fin elements and possessing a pronounced lobe. Head lacks ridges above and below eye. Mouth is small and terminal. Nostril is very small. Eye possesses a nictitating membrane. Gill opening is small, semicircular, and directed posteriorly. Gill rakers are covered with skin. Pectoral fin is short and rounded and has 7 to 10 rays. Dorsal fin has 18 to 20 rays. Median lobe of clavus is about equal to head length, is supported by three to nine rays, and lacks terminal ossifications. Body is covered with thick collagenous skin and bears small scutelike scales. Color is silvery blue, often with silvery spots on sides.

This species occurs worldwide in tropical and warm temperate seas. In the western Atlantic it occurs from Massachusetts Bay to the northern Gulf of Mexico. In the Gulf of Mexico it has been recorded from Mississippi and Louisiana. Like other members of the family, this species is oceanic but a weak swimmer. Most records are based on strandings along the shore, resulting from unfavorable winds and currents that transport specimens into shallow water, where low salinities, temperatures, or other factors lead to death and beaching. Maximum known size is 200 cm TL. references: Raven 1939b; Fraser-Brunner 1951; Tortonese 1986f; Boschung 1992; Smith 1997; Klein-MacPhee 2002c; Matsuura 2002e.

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MOLIDAE 941

Mola mola (Linnaeus 1758) Ocean sunfish

Mola mola is orbicular, compressed, and truncate posteriorly, with clavus supported by dorsal and anal fin elements and lacking a pronounced lobe. Snout projects slightly beyond mouth. Head possesses a supraoccipital ridge extending from snout to posterior margin of eye and a less distinct ridge running from snout to pectoral fin base. Mouth is small and terminal. Eye possesses a nictitating membrane. Gill opening is small. Gill rakers on first arch are concealed beneath thick skin. Pectoral fin is located behind gill slit and has 11 to 13 rays. Dorsal fin is short, high, and falcate and originates above posterior third of body. Dorsal fin has 15 to 20 rays, and anal fin has 14 to 18 rays. Clavus is gently curved and has about 12 rays, and 8 or 9 of these have terminal ossifications. Body is covered with thick collagenous skin, and trunk and fins are covered with small, bony tubercles. Color is dark gray dorsally, with side grayish brown and belly dusky. Dark bar runs along base of dorsal fin, anal fin, and clavus. Snout projection is capped with a bony tubercle.

This species occurs worldwide in tropical and warm temperate seas. In the western Atlantic it occurs from Newfoundland to northern South America, including the Gulf of Mexico and the Caribbean Sea. In the Gulf of Mexico it has been reported from along the Texas and Louisiana coasts. Most reports are based on strandings of specimens along the shore. It is a pelagic oceanic species with very poor means of locomotion. Onshore winds and currents can cause it to be stranded in shallow low-salinity water, leading to its death and beaching. Food consists of jellyfishes and siphonophores. Maximum known size is 350 cm TL.

references: Jordan and Evermann 1898; Gunter 1941; Baughman 1950; Fraser-Brunner 1951; Springer and Bullis 1956; Kemp 1957; Dawson 1965; Anderson and Cupka 1973; Robins and Ray 1986; Tortonese 1986f; Klein and MacPhee 2002c; Matsuura 2002e.

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942 TETRAODONTIFORMES

Ranzania laevis (Pennant, 1776) Slender mola

Ranzania laevis is elongate and oval shaped in profile, compressed, and truncate posteriorly, with clavus supported by vertebral and dorsal and anal fin elements and body covering consisting of smooth hexagonal plates. Mouth is terminal. Lips protrude beyond teeth, forming funnel, and close as a vertical slit. Gill rakers are free of integument. Pectoral fin is falcate and has an oblique base and 12 or 13 rays. Dorsal fin originates at end of body, is high and short, and has 17 or 18 rays. Anal fin is similar in shape and size to dorsal fin, is located beneath dorsal fin, and has 20 rays. Clavus has a nearly straight posterior margin and 22 rays. Body is covered with smooth hexagonal plates.

Color is blackish blue dorsally and gray on side and ventrally. Head and belly are patterned with silvery lines. This species occurs worldwide in tropical and warm temperate seas. In the western Atlantic it occurs from Florida, off Palm Beach on the east coast of Florida and Little Marco Pass (2602N, 8146W) in the eastern Gulf of Mexico, to Brazil. Maximum known size is 810 mm TL. references: Jordan and Evermann 1898; Raven 1939a; Fraser-Brunner 1951; Robins 1968; Topp and Girardin 1972; Robins and Ray 1986; Heemstra 1986r; Tortonese 1986f; Matsuura 2002e.

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GLOSSARY

(Irregular plural form of noun is in parentheses following singular term.) Adinate: condition in which bony margin is

concealed beneath scales and skin; often refers to opercular bones. Adipose eyelid: transparent to translucent tissue covering anterior and posterior sections of eye and leaving an elongate, elliptical depression over pupil of some fishes (e.g., Mugilidae, Carangidae). Air bladder: See gas bladder. Alveolar process: elongate, generally toothbearing projection of premaxilla that forms part or all of upper gape of mouth. Anteroventral: refers to location on anterior lower, abdominal plane or axis of a bilaterally symmetrical animal body. Anthiine: the vernacular name for a tribe of Serranidae. Antimere: other member of a paired structure; generally antimeres are mirror images of each other and located on each side of the longitudinal axis of the body (e.g., right and left premaxillae, tooth-bearing bones of upper jaw). Antrorse: directed forward or upward, in reference to spines, spinelike structures, or teeth; opposite of retrorse. Appress: function of moving an appendage (fin) to lie flat against the surface (body) to which it is attached. Aricular: needlelike, in reference to scales. Auxillary scales: small scales superimposed on the surface of larger scales. Axil: inner or posterior margin of a fin base,

or the angle formed by inner edge of fin and body. Axillary process: enlarged and usually elongated scale at base of pectoral or pelvic fin. Band: vertical color marking on side of fishes that has more or less straight margins and meets along dorsal and ventral midlines. Barbel: fleshy, tentacle-like protuberance located on head, near mouth, chin, or snout of some fishes (e.g., Gadidae, Mullidae, Sciaenidae), and is sensitive to tactile and/or chemical stimuli. Basibranch: median skeletal element of gill arch (branchial) skeleton, located behind tongue. Basicaudal: related to or located in area at base of caudal fin. Bilobed/bilobate: having two lobes. Biramous: having two arms or limblike structures. Bony ridge scale: thin, transparent scale lacking dense enameloid and dentinal layers. Scale consists of a surface “bony” layer of ridges and valleys and an inner layer of fibrous connective tissue. Bony stay: See suborbital stay. Branchiostegal membrane: membrane below operculum connecting branchiostegals and enclosing gill chamber ventrally, often connected to isthmus and/or to antimere; also called “gill membrane.” Branchiostegal ray: one of a series of long, curved, generally slender bones that connect the hyoid arch and ventral margin of operculum and support gill, or branchiostegal, membrane.

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944 GLOSSARY

Bulla (bullae): a swelling or elevation in a

bony skeletal element produced by a capsule housing a sensory organ. Canine: a prominent, conical, and sharppointed tooth (usually larger than other jaw teeth). Caninoid: refers to teeth that are caninelike. Cardiform: refers to short, fine, pointed teeth arranged in a patch and resembling a wool card or comb. Catadromous: refers to fishes that migrate from freshwaters to marine waters to spawn. Caudal peduncle: section of fish body between the end of anal fin and base of caudal fin (end of hypural plate). Cephalic disc: an oval-shape adhesive disc located on dorsal midline behind the head and derived from the first dorsal fin elements; distinctive of Echeneidae (remoras). Cheek: surface of head between the eye and the anterior margin of the preoperculum. Circumpeduncular: refers to the circumference of the caudal peduncle; generally applied to the number of scales in a more or less vertical line around the caudal peduncle. Circumtropical: term used to describe taxa that occur worldwide at tropical latitudes. Cirrus (cirri): small, slender, fleshy protuberance found on head of some fishes. Clavus: caudal fin in which the fin rays are derived from the dorsal and anal fins; distinctive of Molidae. Cleithrum (cleithra): major dermal bone of the pectoral fin girdle that extends dorsally from pectoral fin base and forms posterior margin of opercular cavity. It articulates dorsally with the supracleithrum and ventrally with the scapula and coracoid. Commensal: describes a symbiotic relationship between two species in which one species is benefited and the other species is neither helped nor harmed. Commissural: refers to the site (commissure) where two corresponding parts meet (e.g., symphysis of the jaws); also, to the band of fibers joining two symmetrical structures or antimeres.

Compressed: laterally flattened, from side

to side. Coranoid process: elevated region of dentary

bone of lower jaw that serves as a site for muscle insertions. Corselet: densely scaled area behind the pectoral fins; typical of a number of species of Scombridae and thought to reduce drag during rapid swimming. Countercolored: refers to color pattern that is dark above and light below. Crenate: refers to an irregular bony margin with small rounded projections (crenulations). Ctenii: small, slender spinelike structures found on the posterior field or margin of ctenoid scales. Ctenoid scale: thin and flexible to slightly thickened and inflexible scale bearing small spines or projections (ctenii) on posterior exposed surface. Cycloid scale: thin, flexible scale with a smooth posterior exposed surface. Deciduous: refers to scales that are loosely attached to body and thus easily lost. Dentary: large tooth-bearing bone of lower jaw of fishes with right and left antimeres meeting at symphysis. Depressed: dorsoventrally flattened. Dewlap: fleshy fold or ridge of skin along midline of chin, or large loose flap of skin between pelvic bone and belly in Monacanthidae. Dichromism/dichromatic: refers to having two color patterns; differences may be between growth stages or between the sexes. Dorsoventral: refers to dorsal ventral plane or axis of an organism. Ectopterygoid: bone or cartilage of the upper jaw, part of hyoid arch that articulates with palatine anteriorly and quadrate posteriorly. Ellipsoid: elliptical in shape. Emarginate: refers to a surface, such as a fin, with a concave margin. Entire: whole; refers to a bony margin that lacks rugosities, serrations, or spines and thus is smooth.

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GLOSSARY 945

Epibranch: bone or cartilaginous element of

branchial or gill arch that bears gill filaments and gill rakers. It articulates with pharyngobranch dorsally and ceratobranch ventrally. Ethmoid: unpaired bone or cartilage of anterior section of neurocranium above vomer that forms part of nasal capsule. Evagination: everting of an embryonic structure. Filamentous/filiform: having a round, very slender structure that is cordlike or filament-like. Fimbria (fimbriae): threadlike or fingerlike filaments producing a fringed margin. FL/Fork length: distance from tip of snout to end of middle of caudal fin when fin is maximally spread. Flange: refers to a projecting ridge for strength or for support. Foramen (foramina): small opening or perforation, generally in a bone or cartilage, through which nerves or blood vessels run. Frenum: a connecting or restraining membrane (e.g., membrane connecting pelvic fin spines in Gobiidae, or membrane connecting upper lip to skin of snout, thus inhibiting jaw protrusion). Gas bladder: a diverticulum of the anterior section of the gastrointestinal tract in rayfinned fishes that is filled with gases or fats and enables the fishes that possess them to be suspended in the water column. Gill chamber: space on each side of head containing gills. Gill raker: bony projection on anterior section of epibranchial and ceratobranch bones; cartilage of gill arch. Gular region: throat; area behind chin and between dentary bones of lower jaw, or area in front of pectoral fins. Hyoid: second branchial arch that supports tongue, forms floor of mouth, and suspends jaws from cranium. Imbricate: overlapping evenly; refers to arrangement of cycloid and ctenoid scales in which the anterior scales overlap anterior sections of adjacent posterior scales.

Incisiform/incisor-like: shaped like an incisor;

refers to teeth. Incisor: laterally expanded tooth with blade-

like cusp used for cropping prey (sessile animal or plant organisms). Included: refers to jaws in which either the upper or the lower jaw extends beyond the other jaw. Infracaudal gland: luminous gland below caudal peduncle. [Correction of definition in volume 1.] Inframedian: refers to position of series of bony scutes running along body ventral to supramedian series of scutes. Infraorbital: refers to area below orbit. [Correction of definition in volume 1.] Infraorbital bones: six dermal bones that form anterior, ventral, and posterior margins of eye. Inquiline: inhabitant; refers to the symbiotic relationship in which one organism lives within the body cavity of another organism. Insertion: site of attachment, as in attachment of fin to body of fish. Interbranchial foramen (foramina): a small opening in the interbranchial septum that connects the right and left gill chamber (e.g., Achiridae). Interbranchial septum: membrane lining mesial aspect of gill chamber. Intercalate: to insert; refers to scales between lateral line scales. Interpelvic process: fleshy projection between inner edges of pelvic fins; may be single or bifid, small or large (e.g., Scombridae). Invagination: enclosing of an embryonic membrane. Isthmus: fleshy throat region of a fish that separates the gill chambers and joins ventral section of body to shoulder girdle. Jugular: refers to the ventral part of the body between the end of the lower jaw and the pectoral fin insertion; used to describe pelvic fin that inserts anterior to pectoral fin insertion. Junior synonym: a taxonomic name of a species that is not recognized as valid because

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946 GLOSSARY

it was described after the taxonomic name that is accepted. Labial fold: fold of skin above and/or below lips; separates the lips from skin covering the head. Lachrymal bone: anteriormost of the dermal bones that surround or partially surround orbits; also called “first infraorbital” or “preorbital bone.” Lamella (lamellae): a minute structure on gill filaments that serves as the site of the gaseous exchange between water and gills; it consists of epithelial cells, pillar cells, and basement membrane. Lamina (laminae): a thin layer of tissue. Lanceolate: shaped like the head of a lance; narrow and tapering to a point. Lappet: small fleshy tab or flap on body surface. Lateral: refers to the side of a bilaterally symmetrical body, versus the medial or middle of the body along the longitudinal axis. Lateral ethmoid: dermal bone forming the anterior margin of the orbit and the side of the nasal cavity. Lateral line: system of tubes, canals, or pores containing sensory cells (neuromasts) that respond to low-frequency sounds and water movements. On head, the system usually consists of a network of branching tubes, canals, and pores, and on trunk, of a single series of pores or a perforated canal extending from dorsal margin of operculum to caudal fin base. Lateral line scales: scales on trunk over lateral line that are perforated by lateral line pores or canals. Longitudinal axis: line in the median sagittal plane extending from head to tail. Lunate: shaped like a half moon; refers to a fin (often pectoral) that is long, gently curved, and narrow throughout length (e.g., Scombridae). Maxilla: dermal bone articulating with palatine and often with articular process of premaxilla, and either forming posterior section of upper gape of mouth or located above alveolar process of premaxilla.

Medial/median: refers to the vertical longitu-

dinal plane passing from head to tail through the center of the body. Melanophore: pigment cell that contains melanin (black, brown, or red pigment). Membranous: membranelike; refers to bony margin that is covered by integument. Mental: refers to chin region of head. Mesial: toward or along the middle, or median, plane. Mesopelagic: refers to the region of the water column between 200 and 1,000 m and the organisms that live in this region. Molar: stout tooth with broad, flattened, or rounded cusp used for crushing prey. Molariform: molar shaped; refers to teeth that are stout, blunt or rounded, and designed for crushing hard-bodied invertebrates. Monophyletic: belonging to a natural group of organisms; refers to a group of species or higher taxa that share a single unique common ancestor. Multifid: divided into many lobes or divisions. Myomere: trunk musculature or muscle block. Nape: area on dorsal surface of body between the end of the cranium and the origin of the dorsal fin; region of head above and behind eyes. Nasal: referring to nostrils or olfactory organs. Negatively sloped: refers to a threedimensional structure that extends forward more on the dorsal side than on the ventral side. Neoteleostei: a derived taxonomic group (clade) of fishes that includes the Stomiiformes, Ateleopodiformes, Aulopiformes, and the remainder of derived teleost fishes. Neritic: refers to organisms that occur in the water column over continental and insular shelves (i.e., in less than 200 m of water). Neuromast: mechanoreceptor that responds to movement of liquid (water or endolymph fluid) and is composed of a gelatinous cap, cilia, and a cell body. Nonimbricate: nonoverlapping; refers to scales that do not overlap.

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GLOSSARY 947

Nuchal: related to or associated with the nape. Occipital pit: mid-dorsal depression in

head at posterior region of cranium (e.g., Scorpaenidae). Occipital region: upper region at posterior end of cranium. Occiput: back of the head or cranium; region of cranium posterior to eyes; posterior border of cranium. Ocellated: having a color pattern consisting of eyespots: pigment spots surrounded by one or more different-colored rings. Operculum (opercula): dermal bone covering much of the gill cavity. Orbicular: spherical, ringlike, circular shaped (e.g., Molidae). Origin: fixed attachment of a muscle; refers to anterior end of a fin base, especially dorsal or anal fin. Ostia: pores on basal membranes of dorsal and anal fins (e.g., Cynoglossidae). Outcurved: curved forward; refers to teeth, spines, or spinelike structures; opposite of recurved. Oviparous: reproductive mode in which fertilized eggs develop and hatch outside of the female’s body. Palatine: bone or cartilage of upper jaw, part of hypoid arch that often bears teeth and posteriorly articulates with ectopterygoid. Palmate: hand-shaped; refers to shape of tentacles of blennioid fishes. Papilla (papillae): small nipplelike projection or process. Parapophysis (parapophyses): lateral process on precaudal vertebrae. Parietal bone: cranial bone forming part of roof of cranium, located posterior to frontal bone and lateral to supraoccipital bone. Pectinate: having toothlike projections resembling the teeth of a comb. Pedicel: lateral, tooth-bearing extension of the premaxilla that forms part or all of the upper gap of the mouth in neoteleostean fishes. Also called “alveolar process.” Pelvic frenum: membrane connecting two pelvic fin spines (e.g., Gobudae).

Penultimate: next-to-the-last position in a

series of discrete variables such as vertebrae or fin rays. Perifacial rim: bony ridge extending from infraorbital bones to posterior margin of preoperculum (e.g., Peristediidae). Pharyngeal sac: toothed saccular outgrowth of the esophagus (e.g., Stromateidae). Physoclistous gas bladder: gas bladder that lacks a connection (pneumatic duct) with the esophagus. Polytomy: a phylogenetic relationship of more than two taxa that is unresolved. Posttemporal region: anterior section of body behind neurocranium, usually above anterodorsal aspect of operculum. Precaudal vertebrae: vertebrae that precede caudal vertebrae; they bear ribs and lack hemal spines. Premaxilla (premaxillae): dermal bone forming anterior section or entire upper gap of mouth in ray-finned fishes. Preoperculum (preopercula): dermal bone attached to hyomandibula and located anterior to operculum. Preorbital bone: See lachrymal bone. Procumbent: lying flat in a forward direction, in reference to orientation of spines, spinelike structures, and teeth. Procurrent: inclined forward; refers to small unsegmented, unbranched rays located above and below posterior section of caudal peduncle. Produced: elongated, projecting, or extended. Protandry: type of sequential hermaphroditism in which individuals first develop and function as males and subsequently switch to and function as females. Protogyny: type of sequential hermaphroditism in which individuals first develop and function as females and subsequently switch to and function as males. Protractile: able to be lengthened; refers to forward movement of premaxilla over the ethmovomer block that leads to protrusion of the upper jaw. Premaxillae that are capable of protraction are located anterior and ventral to the max-

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948 GLOSSARY

illa and have high ascending and articular processes. Protrusible: able to be thrust forward; refers to forward movement of upper jaw as a result of protraction of the premaxilla. Pseudobranch(s): small gill-like structure located on inner surface of operculum. Pterygiophore: splintlike bone or cartilage, located in the flesh medial to the unpaired fins, that articulates and supports the fin rays. Pyloric caeca (caecae): one to many blind sacs located on the anterior end of the intestine, near its junction with the pyloric end of the stomach, that function in digestion and absorption. Ray-finned fishes: a putative monophyletic group of fishes comprising the taxon Actinopterygii. Recess: small cavity, hollow, or indentation. Recurved: curved backward; refers to teeth, spines, or spinelike structures; opposite of outcurved. Rete mirabile (retia mirabilia): capillary bed in which closely spaced arterial capillaries run parallel and in opposite direction to venous capillaries, thus facilitating the exchange of gases or heat between the oppositely flowing blood vessels; also called “wonder net.” Retrorse: bent or turned backward or downward; refers to teeth or spines; opposite of antrorse. Rictus: corner of mouth. Rostral exsertion: anterior projection of lachrymal bones (e.g., Triglidae and Peristediidae). Rostral frenum: integumentary bridge uniting rostrum and lip fold. Rostrum: projecting snout or beak. Rugose: ridged; generally refers to surface structure of dermal bones of cranium. Saddle: vertical color marking that meets along dorsal midline and extends variable distances along side but does not meet along ventral midline. Sagittal plane: vertical longitudinal plane through body, either in median plane or parallel to median plane.

Scute: thickened scale that may possess spines

and/or ridges. Secondary squamation: scales partially or

totally covering primary scales. Semilunate: shaped like a half moon or

crescent. Senior synonym: accepted name; oldest avail-

able name of a taxon. Setiform: bristlelike; refers to sandpaper-like teeth. Setose: bristlelike. Simple rays: unbranched fin rays. SL/Standard length: distance from tip of snout to end of hypural plate. End of plate is at seam produced when tail is bent laterally toward body. Sphenotic: cranial bone or cartilage articulating with frontal bone anteriorly. Spinule: small spine. Squamation: body covering, such as scales or scale derivatives. Stripes: horizontal color markings with more or less straight margins. Subabdominal: located posterior to pectoral fin base under anterior section of abdomen; refers to pelvic fin location. Subequal: slightly less than something else; generally used to quantitatively describe an object by comparing it to a similar object. Suborbital stay: bony ridge on third infraorbital bone extending from cheek to preopercular margin; distinctive of the scorpaeniform fishes. Subtended: included in, as in lower jaw is subtended or included in upper jaw when mouth is closed. Supracleithrum: dorsalmost bone or cartilage of pectoral girdle; articulates ventrally with cleithrum. Supramaxilla: small dermal bone located along dorsal margin of maxilla. Supramedian: refers to location of series of bony scutes on body running ventral to dorsal series of scutes (e.g., Peristediidae). Symphysis: junction between two bones or between two bony or cartilaginous antimeres. Synapomorphy: a derived character that is shared by two or more taxa.

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GLOSSARY 949

Tab: small membranous fold or flap, generally

located on head. Tapetum lucidum: iridescent choroid membrane in the eye. Terete: cylindrical and tapered in shape. Test: exoskeleton of crustaceans. Thermocline: interface between mixed surface waters and cold deep waters where the temperature declines by about one degree or more per meter. Tholichthys: larvae of chaetodontids that are covered with bony plates. Thoracic: in chest region; refers to location of pelvic fin below pectoral fin base. TL/Total length: distance from tip of snout to tip of caudal fin. Tooth plate: several to many teeth embedded in bony plate (e.g., basibranchial tooth plate).

Trifid: divided into three lobes or parts. Tubercle: temporary epidermal growth

on head or a knoblike bony structure on body. Unicuspid: refers to teeth with a simple, or single, point or cusp. Urohyal: unpaired bone or cartilage of hyoid arch that lines inside rami of lower jaw. Vent/Venter: ventral region of body in area of anus and urigenital openings. Vermiculation: wormlike or undulating color pattern. Villiform: shaped like a small bristle; refers to very fine, relatively long brushlike teeth. Vomer: median, unpaired dermal cranial bone or cartilage located below ethmoid and often bearing teeth.

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REFERENCES

NOTE: Multiple entries by the same author in the same year from the same edited volume or journal issue are listed by the page order in which they appeared in the original work. Entries by the same author from different works in the same year are ordered alphabetically by title. Title abbreviations and full publication information for frequently cited works are listed below.

FAO-PC-O  Guía FAO para la identificación de especies para los fines de la pesca, Pacífico Centro-oriental, ed. W. Fischer, K. Krupp, W. Schneider, C. Sommer, K. E. Carpenter, and V. H. Niem. 1995. Rome: FAO. 3 vols., 1813 pp. FAO-WCA  FAO species identification guide for fishery purposes, Western Central Atlantic (Fishing area 31), ed. K. Carpenter. 2002. Rome: FAO. 3 vols., 2127 pp. FAO-WCP  FAO species identification guide for fishery purposes, Western Central Pacific (Fishing area 71), ed. K. Carpenter and V. H. Niem. 2001. Rome: FAO. 6 vols., 4218 pp. FJA  The fishes of the Japanese Archipelago, ed. H. Masuda, K. Amaoka, C. Araga, T. Uyeno, and T. Yoshino. 1984. Tokyo: Tokai University Press. 437 pp., 370 pls. FN-EAM  Fishes of the North-eastern Atlantic and the Mediterranean, ed. P. J. P. Whitehead, M. L. Bauchot, J. C.

Hureau, J. Nielsen, and E. Tortonese. 3 vols.: Vol. 1, 1984: 1–510; Vol. 2, 1986: 511–1008; Vol. 3, 1986: 1009 – 1473. Paris: UNESCO. FTSFG  Fishes trawled off Suriname and French Guiana, ed. T. Uyeno, K. Matsuura, and E. Fujii. 1983. Tokyo: Japan Marine Fishery Resource Research Center. 519 pp. OSF  Ontogeny and systematics of fishes, ed. H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall, Jr., and S. L. Richardson. Special Publ. No. 1. Lawrence (KS): Amer. Soc. Ichthyol. Herpet. 759 pp. SSF  Smith’s sea fishes, ed. M. M. Smith and P. C. Heemstra. 1986. New York: Springer-Verlag. 1047 pp. Agafonova, T. B. 1994. Systematics and distribution of Cubiceps (Nomeidae) of the world ocean. J. Ichthyol. 34: 116 –143. Agafonova, T. B., and E. I. Kukuyev. 1990. New data on the distribution of Cubiceps gracilis. Voprosy Ikhtiologii 30: 1028 – 1031 (1991 Scripta Technica, Inc. ISSN0032 – 452/91/0002 – 0132). Agafonova, T. B., and V. F. Poluyaktov. 1992. Age and growth rate of two species of cigarfishes, Cubiceps caeruleus and C. pauciradiatus (Nomeidae). Voprosy Ikhtiologii 32: 111–119. Ahlstrom, E. L., J. L. Butler, and B. Y. Sumida. 1976. Pelagic stomateoid fishes

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(Pisces: Perciformes) of the eastern Pacific: kinds, distributions, and early life histories and observations on five of these from the northwestern Atlantic. Bull. Mar. Sci. 26: 285 – 402. Allen, G. R. 1980. Butterfly and angelfishes of the world. Vol. 2. New York: John Wiley & Sons. 352 pp. Allen, G. R. 1985. FAO species catalogue. Vol. 6, Snappers of the world: an annotated and illustrated catalogue of lujanid species known to date. FAO Fish. Synop. 125(6): 1–208. Allen, G. R. 1986. Family No. 219: Pomacentridae. In: SSF: 670 – 682. Allen, G. R. 1991. Damselfishes of the world. Melle, Germany: Mergus. 271 pp. Allen, G. R. 2001a. Apogonidae. In: FAOWCP: 2602 –2610. Allen, G. R. 2001b. Pomacentridae. In: FAOWCP: 3337–3345. Amaoka, K. 2001. Paralichthyidae. In: FAOWCP: 3842 –3862. Amaya, F., A. Acero P., and M. M. Criales. 2002. Taxonomy and zoogeography of Microdesmus carri Gilbert (Microdesmidae). Bull. Mar. Sci. 70: 33 –39. Anderson, M. E. 1984. Zoarcidae: development and relationships. In: OSF: 578 –582. Anderson, M. E. 1989. Review of the eelpout genus Pachycara Zugmayer, 1911 (Teleostei: Zoarcidae), with descriptions of six new species. Proc. Calif. Acad. Sci. 46: 221–242. Anderson, M. E. 1990. Zoarcidae. In: Fishes of the southern ocean, ed. O. Gon and H. Heemstra, pp. 256 –276. Grahamstown, South Africa: J. L. B. Smith Institute of Ichthyology. Anderson, M. E. 1994. Systematics and osteology of the Zoarcidae (Teleostei: Perciformes). Ichthyol. Bull. J. L. B. Smith Inst. Ichthyol. 60: 1–120. Anderson, M. E. 2001. Zoarcidae. In: FAO WCP: 3493 –3494. Anderson, M. E. 2002. Zoarcidae. In: FAOWCA: 1740 –1741.

Anderson, W. D., Jr. 1966. A new species of Pristipomoides (Pisces: Lutjanidae) from the tropical western Atlantic. Bull. Mar. Sci. 16: 814 – 826. Anderson, W. D., Jr. 1967. Field guide to the snappers (Lutjanidae) of the western Atlantic. NOAA Tech. Rep. NMSF Circ. 252: 1–14. Anderson, W. D., Jr. 1970. Revision of the genus Symphysanodon (Pisces: Lutjanidae), with descriptions of four new species. Fish. Bull. Fish. Wildl. Serv. 68: 325 –346. Anderson, W. D., Jr. 1972. Notes on western Atlantic lutjanid fishes of the genera Pristipomoides and Etelis. Copeia: 359 –362. Anderson, W. D., Jr. 1981. A new species of Indo-West Pacific Etelis (Pisces: Lutjanidae), with comments on other species of the genus. Copeia: 820 – 825. Anderson, W. D., Jr. 1986. Family No. 181: Lutjanidae. In: SSF: 572 –579. Anderson, W. D., Jr. 2001. Symphysanodontidae. In: FAO-WCP: 2438 –2441. Anderson, W. D., Jr. 2002a. Symphysanodontidae. In: FAO-WCA: 1304 –1307. Anderson, W. D., Jr. 2002b. Serranidae: Anthiinae. In: FAO-WCA: 1330 –1333, 1350 –1352, 1364. Anderson, W. D., Jr. 2002c. Lutjanidae. In: FAO-WCA: 1479 –1505. Anderson, W. D., Jr., and D. M. Cupka. 1973. Records of the ocean sunfish, Mola mola, from the beaches of South Carolina and adjacent waters. Ches. Sci. 14: 295 –298. Anderson, W. D., Jr., and E. J. Gutherz. 1967. Revision of the flatfish genus Trichopsetta (Bothidae), with descriptions of three new species. Bull. Mar. Sci. 17: 892 – 913. Anderson, W. D., Jr., and P. C. Heemstra. 1980. Two new species of western Atlantic Anthias (Pisces: Serranidae), redescription of A. asperilinguis and review of Holanthias martinicensis. Copeia: 72 – 87. Anderson, W. D., Jr., and W. F. Smith-Vaniz.

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INDEX OF SCIENTIFIC NAMES

Abudefduf saxatilis, 490 Abudefduf taurus, 491 Acanthemblemaria aspera, 612 Acanthocybium solandri, 774 Acanthostracion polygonius, 914 Acanthostracion quadricornis, 915 Acanthuridae, 740 Acanthurus bahianus, 742 Acanthurus chirurgus, 743 Acanthurus coeruleus, 744 Achiridae, 864 Achirus lineatus, 866 Acromycter perturbator, 15 Acropomatidae, 111 Acyrtops beryllinus, 643 Alectis ciliaris, 274 Alepocephalidae, 17 Alepocephalus australie, 17 Alphestes afer, 128 Aluterus heudelotii, 903 Aluterus monoceros, 904 Aluterus schoepfii, 905 Aluterus scriptus, 906 Amblycirrhitus pinos, 486 Ancylopsetta dilecta, 837 Ancylopsetta quadrocellata, 838 Anisotremus surinamensis, 364 Anisotremus virginicus, 365 Anthias nicholsi, 129 Anthias tenuis, 130 Anthias woodsi, 131 Aphanopus intermedius, 766 Apogon affinis, 220 Apogon aurolineatus, 221 Apogon binotatus, 222 Apogon lachneri, 223 Apogon maculatus, 224

Apogon planifrons, 225 Apogon pseudomaculatus, 226 Apogon quadrisquamatus, 227 Apogon townsendi, 228 Apogonidae, 216 Apsilus dentatus, 321 Archosargus probatocephalus, 389 Archosargus rhomboidalis, 390 Ariidae, 16 Ariomma bondi, 812 Ariomma melanum, 813 Ariomma regulus, 814 Ariommatidae, 810 Arius assimilis, 16 Astrapogon alutus, 229 Astrapogon puncticulatus, 230 Astrapogon stellatus, 231 Astroscopus y-graecum, 569 Aulopidae, 17 Aulopus filamentosus, 17 Auxis rochei, 775 Auxis thazard, 776 Awaous banana, 668 Bairdiella chrysoura, 415 Bairdiella ronchus, 416 Balistes capriscus, 895 Balistes vetula, 896 Balistidae, 893 Barathrites parri, 17 Barbulifer ceuthoecus, 669 Bassozetus normalis, 17 Bathyanthias cubensis, 132 Bathyclupea argentea, 456 Bathyclupea schroederi, 457 Bathyclupeidae, 455 Bathygobius curacao, 670 98

16-T3324-IX1 9/13/05 11:22 AM Page 988

988 INDEX OF SCIENTIFIC NAMES

Bathygobius mystacium, 671 Bathygobius soporator, 672 Bathysphyraenops simplex, 113 Batrachoides surinamensis, 18 Batrachoididae, 18 Bellator brachychir, 55 Bellator egretta, 56 Bellator militaris, 57 Bellottia apoda, 18 Bembrops anatirostris, 564 Bembrops gobioides, 565 Bembrops macromma, 566 Benthobatis marcida, 15 Benthodesmus tenuis, 767 Blenniidae, 624 Bodianus pulchellus, 508 Bodianus rufus, 509 Bollmannia boqueronensis, 673 Bollmannia communis, 674 Bollmannia eigenmanni, 675 Bollmannia litura, 676 Bothidae, 821 Bothus lunatus, 823 Bothus ocellatus, 824 Bothus robinsi, 825 Brama caribbea, 306 Brama dussumieri, 307 Bramidae, 304 Bythitidae, 18 Caelorinchus ventrilux, 18 Calamus arctifrons, 391 Calamus bajonado, 392 Calamus calamus, 393 Calamus campechanus, 394 Calamus leucosteus, 395 Calamus nodosus, 396 Calamus penna, 397 Calamus pennatula, 398 Calamus proridens, 399 Callionymidae, 646 Cantherhines macrocerus, 907 Cantherhines pullus, 908 Canthidermis maculata, 897 Canthidermis sufflamen, 898 Canthigaster jamestyleri, 921 Canthigaster rostrata, 922 Carangidae, 269

Caranx bartholomaei, 275 Caranx crysos, 276 Caranx hippos, 277 Caranx latus, 278 Caranx lugubris, 279 Caranx ruber, 280 Caristiidae, 311 Caristius maderensis, 312 Cathorops aguadulce, 16 Cathorops melanopus, 16 Cathorops spixii, 16 Caulolatilus chrysops, 245 Caulolatilus cyanops, 246 Caulolatilus intermedius, 247 Caulolatilus microps, 248 Centrodraco acanthopoma, 653 Centrolophidae, 796 Centrolophus niger, 798 Centropomidae, 97 Centropomus ensiferus, 99 Centropomus mexicanus, 100 Centropomus parallelus, 101 Centropomus pectinatus, 102 Centropomus poeyi, 103 Centropomus undecimalis, 104 Centropristis ocyurus, 133 Centropristis philadelphica, 134 Centropristis striata, 135 Centropyge argi, 473 Cephalopholis cruentata, 137 Cephalopholis fulva, 138 Cerdale floridana, 731 Chaenopsidae, 610 Chaenopsis limbaughi, 613 Chaenopsis ocellata, 614 Chaenopsis roseola, 615 Chaetodipterus faber, 736 Chaetodon capistratus, 460 Chaetodon ocellatus, 462 Chaetodon sedentarius, 464 Chaetodon striatus, 466 Chaetodontidae, 458 Chascanopsetta lugubris, 826 Chasmodes longimaxilla, 627 Chasmodes saburrae, 628 Chiasmodon niger, 553 Chiasmodon subniger, 554 Chiasmodontidae, 550

16-T3324-IX1 9/13/05 11:22 AM Page 989

INDEX OF SCIENTIFIC NAMES 989

Chilomycterus antennatus, 932 Chilomycterus antillarum, 933 Chilomycterus reticulatus, 934 Chilomycterus schoepfii, 935 Chloroscombrus chrysurus, 281 Chriolepis benthonis, 677 Chriolepis vespa, 678 Chromis cyanea, 492 Chromis enchrysura, 493 Chromis insolata, 494 Chromis multilineata, 495 Chromis scotti, 496 Cirrhitidae, 485 Citharichthys abbotti, 839 Citharichthys arctifrons, 840 Citharichthys cornutus, 841 Citharichthys dinoceros, 842 Citharichthys gymnorhinus, 843 Citharichthys marcops, 845 Citharichthys spilopterus, 846 Citharichthys uhleri, 847 Clepticus parrae, 510 Congridae, 15 Conodon nobilis, 366 Corvula batabana, 417 Corvula sanctaeluciae, 418 Coryphaena equiselis, 267 Coryphaena hippurus, 268 Coryphaenidae, 265 Coryphopterus dicrus, 679 Coryphopterus eidolon, 680 Coryphopterus glaucofraenum, 681 Coryphopterus hyalinus, 682 Coryphopterus lipernes, 683 Coryphopterus personatus, 684 Coryphopterus punctipectophorus, 685 Coryphopterus thrix, 686 Cosmocampus brachycephalus, 19 Cryptotomus roseus, 531 Ctenogobius boleosoma, 687 Ctenogobius claytonii, 688 Ctenogobius fasciatus, 689 Ctenogobius saepepallens, 690 Ctenogobius shufeldti, 691 Ctenogobius smaragdus, 692 Ctenogobius stigmaticus, 693 Ctenogobius stigmaturus, 694 Cubiceps capensis, 803

Cubiceps gracilis, 804 Cubiceps pauciradiatus, 805 Cyclopsetta chittendeni, 848 Cyclopsetta fimbriata, 849 Cynoglossidae, 872 Cynoscion arenarius, 419 Cynoscion jamaicensis, 420 Cynoscion nebulosus, 421 Cynoscion nothus, 422 Cynoscion regalis, 423 Dactylopteridae, 24 Dactylopterus volitans, 25 Dactyloscopidae, 578 Dactyloscopus moorei, 580 Dactyloscopus tridigitatus, 581 Decapterus macarellus, 282 Decapterus punctatus, 283 Decapterus tabl, 284 Decodon puellaris, 511 Dermatolepis inermis, 139 Diapterus auratus, 347 Diapterus rhombeus, 348 Diodon eydouxii, 936 Diodon holocanthus, 937 Diodon hystrix, 938 Diodontidae, 930 Diplectrum bivittatum, 140 Diplectrum formosum, 141 Diplodus argenteus, 400 Diplodus holbrookii, 401 Diplogrammus pauciradiatus, 648 Diplospinus multistriatus, 755 Doratonotus megalepis, 512 Dormitator maculatus, 656 Draconettidae, 652 Dysalotus alcocki, 555 Dysalotus oligoscolus, 556 Echeneidae, 253 Echeneis naucrates, 255 Echeneis neucratoides, 256 Ectreposebastes imus, 30 Elacatinus horsti, 695 Elacatinus macrodon, 696 Elacatinus oceanops, 697 Elacatinus prochilos, 698 Elacatinus saucrus, 699

16-T3324-IX1 9/13/05 11:22 AM Page 990

990 INDEX OF SCIENTIFIC NAMES

Elacatinus xanthiprora, 700 Elagatis bipinnulata, 285 Eleotridae, 654 Eleotris amblyopsis, 657 Emblemaria atlantica, 616 Emblemaria pandionis, 617 Emblemaria piratula, 618 Emblemariopsis bahamensis, 619 Emblemariopsis diaphana, 620 Emmelichthyidae, 313 Emmelichthyops atlanticus, 383 Emmelichthys ruber, 315 Engyophrys senta, 827 Enneanectes altivelis, 574 Enneanectes boehlkei, 575 Enneanectes jordani, 576 Enneanectes pectoralis, 577 Entomacrodus nigricans, 629 Ephippidae, 735 Epigonidae, 235 Epigonus denticulatus, 237 Epigonus macrops, 238 Epigonus occidentalis, 239 Epigonus oligolepis, 240 Epigonus pandionis, 241 Epigonus pectinifer, 242 Epinephelus adscensionis, 142 Epinephelus drummondhayi, 143 Epinephelus flavolimbatus, 144 Epinephelus guttatus, 145 Epinephelus itajara, 146 Epinephelus morio, 147 Epinephelus mystacinus, 148 Epinephelus nigritus, 149 Epinephelus niveatus, 150 Epinephelus striatus, 151 Epinnula magistralis, 756 Equetus lanceolatus, 424 Equetus punctatus, 426 Erotelis smaragdus, 658 Erythrocles monodi, 316 Etelis oculatus, 322 Etmopterus robinsi, 15 Etropus crossotus, 850 Etropus cyclosquamus, 851 Etropus rimosus, 852 Eucinostomus argenteus, 349 Eucinostomus gula, 350

Eucinostomus harengulus, 351 Eucinostomus havana, 352 Eucinostomus jonesii, 353 Eucinostomus lefroyi, 354 Eucinostomus melanopterus, 355 Eugerres brasilianus, 356 Eugerres plumieri, 357 Eurypharyngidae, 16 Eurypharynx pelecanoides, 16 Eustomias achirus, 17 Eustomias monoclonus, 17 Euthynnus alletteratus, 777 Evermannichthys spongicola, 701 Evorthodus lyricus, 702 Evoxymetopon taeniatus, 768 Exechodontes daidaleus, 547 Foetorepus agassizii, 649 Foetorepus goodenbeani, 650 Gastropsetta frontalis, 853 Gempylidae, 753 Gempylus serpens, 757 Gerreidae, 344 Gerres cinereus, 358 Gillellus greyae, 582 Gillellus healae, 583 Gillellus uranidae, 584 Gnathagnus egregius, 570 Gnatholepis thompsoni, 703 Gobiesocidae, 641 Gobiesox punctulatus, 644 Gobiesox strumosus, 645 Gobiidae, 661 Gobioides broussonetii, 704 Gobiomorus dormitor, 659 Gobionellus oceanicus, 705 Gobiosoma bosc, 706 Gobiosoma ginsburgi, 707 Gobiosoma grosvenori, 708 Gobiosoma hemigymnum, 709 Gobiosoma longipala, 710 Gobiosoma robustum, 711 Gobulus myersi, 712 Gonioplectrus hispanus, 152 Gramma loreto, 196 Grammatidae, 194 Guavina guavina, 660

16-T3324-IX1 9/13/05 11:22 AM Page 991

INDEX OF SCIENTIFIC NAMES 991

Gymnachirus melas, 867 Gymnachirus nudus, 868 Gymnachirus texae, 869 Haemulidae, 359 Haemulon album, 367 Haemulon aurolineatum, 368 Haemulon bonariense, 369 Haemulon boschmae, 370 Haemulon carbonarium, 371 Haemulon chrysargyreum, 372 Haemulon flavolineatum, 373 Haemulon macrostomum, 374 Haemulon melanurum, 375 Haemulon parra, 376 Haemulon plumierii, 377 Haemulon sciurus, 378 Haemulon striatum, 379 Halichoeres bathyphilus, 513 Halichoeres bivittatus, 514 Halichoeres caudalis, 515 Halichoeres cyanocephalus, 516 Halichoeres garnoti, 517 Halichoeres maculipinna, 518 Halichoeres pictus, 519 Halichoeres poeyi, 520 Halichoeres radiatus, 521 Harriotta raleighana, 14 Helicolenus dactylopterus, 31 Hemanthias aureorubens, 153 Hemanthias leptus, 154 Hemanthias vivanus, 155 Hemicaranx amblyrhynchus, 286 Hemiemblemaria simulus, 621 Heteropriacanthus cruentatus, 213 Hippoglossina oblonga, 854 Holacanthus bermudensis, 474 Holacanthus ciliaris, 476 Holacanthus tricolor, 478 Hollardia hollardia, 890 Hollardia meadi, 891 Howella brodiei, 118 Howellidae, 117 Hyperoglyphe bythites, 799 Hyperoglyphe perciformis, 800 Hypleurochilus bermudensis, 630 Hypleurochilus caudovittatus, 631 Hypleurochilus multifilis, 632

Hypoplectrus chlorurus, 157 Hypoplectrus gemma, 158 Hypoplectrus nigricans, 159 Hypoplectrus puella, 160 Hypoplectrus unicolor, 162 Hypsoblennius hentz, 633 Hypsoblennius invemar, 634 Hypsoblennius ionthas, 635 Inermia vittata, 384 Inermiidae, 382 Istiophorus albicans, 791 Joturus pichardi, 19 Kali indica, 557 Kali macrodon, 558 Kali normani, 559 Kathetostoma albigutta, 571 Katsuwonus pelamis, 778 Kyphosidae, 482 Kyphosus incisor, 483 Kyphosus sectator, 484 Labridae, 504 Labrisomidae, 586 Labrisomus albigenys, 590 Labrisomus bucciferus, 591 Labrisomus gobio, 592 Labrisomus guppyi, 593 Labrisomus haitiensis, 594 Labrisomus kalisherae, 595 Labrisomus nigricinctus, 596 Labrisomus nuchipinnis, 597 Lachnolaimus maximus, 522 Lactophrys trigonus, 916 Laemonema goodebeanorum, 18 Lagocephalus laevigatus, 923 Lagodon rhomboides, 402 Larimus breviceps, 428 Larimus fasciatus, 429 Leiostomus xanthurus, 430 Lepidocybium flavobrunneum, 758 Lepidopus altifrons, 769 Liopropoma eukrines, 163 Liopropoma rubre, 164 Lipogramma anabantoides, 197 Lipogramma trilineatum, 198

16-T3324-IX1 9/13/05 11:22 AM Page 992

992 INDEX OF SCIENTIFIC NAMES

Lobotes surinamensis, 343 Lobotidae, 342 Lonchopisthus mircognathus, 202 Lophogobius cyprinoides, 713 Lopholatilus chamaeleonticeps, 249 Lupinoblennius nicholsi, 636 Lupinoblennius vinctus, 637 Lutjanidae, 317 Lutjanus analis, 323 Lutjanus apodus, 324 Lutjanus buccanella, 325 Lutjanus campechanus, 326 Lutjanus cyanopterus, 327 Lutjanus griseus, 328 Lutjanus jocu, 329 Lutjanus mahogoni, 330 Lutjanus purpureus, 331 Lutjanus synagris, 332 Lutjanus vivanus, 333 Luvaridae, 738 Luvarus imperialis, 739 Lycenchelys bullisi, 548 Lythrypnus elasson, 714 Lythrypnus nesiotes, 715 Lythrypnus phorellus, 716 Lythrypnus spilus, 717 Macrouridae, 18 Makaira nigricans, 792 Malacanthidae, 243 Malacanthus plumieri, 250 Malacoctenus aurolineatus, 598 Malacoctenus gilli, 599 Malacoctenus macropus, 600 Malacoctenus triangulatus, 601 Masturus lanceolatus, 940 Melanostomiidae, 17 Melichthys niger, 899 Menticirrhus americanus, 431 Menticirrhus littoralis, 432 Menticirrhus saxatilis, 433 Microdesmidae, 730 Microdesmus carri, 732 Microdesmus lanceolatus, 733 Microdesmus longipinnis, 734 Microgobius carri, 718 Microgobius gulosus, 719 Microgobius microlepis, 720

Microgobius thalassinus, 721 Micropogonias furnieri, 434 Micropogonias undulatus, 435 Microspathodon chrysurus, 497 Mitsukurina owstoni, 15 Mitsukurinidae, 15 Mola mola, 941 Molidae, 939 Monacanthidae, 901 Monacanthus ciliatus, 909 Monacanthus tuckeri, 910 Monolene sessilicauda, 828 Moridae, 18 Morone chrysops, 107 Morone mississippiensis, 108 Morone saxatilis, 109 Moronidae, 105 Mugil gaimardianus, 19 Mugilidae, 19 Mullidae, 447 Mulloidichthys martinicus, 449 Mullus auratus, 450 Mustelus higmani, 14 Mustelus sinusmexicanus, 14 Mycteroperca acutirostris, 165 Mycteroperca bonaci, 166 Mycteroperca interstitialis, 167 Mycteroperca microlepis, 168 Mycteroperca phenax, 169 Mycteroperca tigris, 170 Mycteroperca venenosa, 171 Narcine bancroftii, 15 Narcinidae, 15 Naucrates ductor, 287 Nealotus tripes, 759 Nemaclinus atelestos, 602 Neoepinnula americana, 760 Neomerinthe beanorum, 32 Neomerinthe hemingwayi, 33 Nes longus, 722 Nesiarchus nasutus, 761 Nicholsina usta, 532 Nomeidae, 801 Nomeus gronovii, 806 Ocyurus chrysurus, 334 Odontoscion dentex, 436

16-T3324-IX1 9/13/05 11:22 AM Page 993

INDEX OF SCIENTIFIC NAMES 993

Oligoplites saurus, 288 Ophidiidae, 17 Ophidion holbrookii, 18 Ophidion josephi, 18 Ophidion selenops, 18 Ophioblennius macclurei, 638 Opistognathidae, 199 Opistognathus aurifrons, 203 Opistognathus lonchurus, 204 Opistognathus macrognathus, 205 Opistognathus maxillosus, 206 Opistognathus melachasme, 207 Opistognathus nothus, 208 Opistognathus robinsi, 209 Opistognathus whitehursti, 210 Opsanus dichrostomus, 18 Orthopristis chrysoptera, 380 Ostraciidae, 913 Oxynotidae, 15 Oxynotus caribbaeus, 15 Oxyurichthys stigmalophius, 723 Pachycara sulaki, 549 Pagrus pagrus, 403 Palatogobius paradoxus, 724 Parablennius marmoreus, 639 Paraclinus cingulatus, 603 Paraclinus fasciatus, 604 Paraclinus marmoratus, 605 Paraclinus nigripinnis, 606 Paradiplogrammus bairdi, 651 Parahollardia lineata, 892 Paralichthyidae, 832 Paralichthys albigutta, 855 Paralichthys lethostigma, 856 Paralichthys squamilentus, 857 Paralonchurus brasiliensis, 437 Paranthias furcifer, 172 Pareques acuminatus, 438 Pareques iwamotoi, 440 Pareques umbrosus, 441 Pempheridae, 453 Pempheris schomburgkii, 454 Peprilus burti, 818 Peprilus paru, 819 Percophidae, 563 Peristediidae, 70 Peristedion antillarum, 73

Peristedion brevirostre, 74 Peristedion ecuadorense, 75 Peristedion gracile, 76 Peristedion greyae, 77 Peristedion imberbe, 78 Peristedion longispatha, 79 Peristedion miniatum, 80 Peristedion sp., 84 Peristedion thompsoni, 81 Peristedion truncatum, 82 Peristedion unicuspis, 83 Phaenacoscorpius nebris, 34 Phaeoptyx conklini, 232 Phaeoptyx pigmentaria, 233 Phaeoptyx xenus, 234 Phtheirichthys lineatus, 257 Platygillellus rubrocinctus, 585 Plectranthias garrupellus, 173 Poecilopsetta beanii, 862 Poecilopsetta inermis, 863 Poecilopsettidae, 861 Pogonias cromis, 442 Polydactylus octonemus, 407 Polydactylus oligodon, 408 Polydactylus virginicus, 409 Polynemidae, 405 Pomacanthidae, 471 Pomacanthus arcuatus, 479 Pomacanthus paru, 481 Pomacentridae, 487 Pomadasys crocro, 381 Pomatomidae, 251 Pomatomus saltatrix, 252 Pontinus longispinis, 35 Pontinus rathbuni, 36 Priacanthidae, 211 Priacanthus arenatus, 214 Priolepis hipoliti, 725 Prionotus alatus, 58 Prionotus longispinosus, 59 Prionotus martis, 60 Prionotus murielae, 61 Prionotus ophryas, 62 Prionotus paralatus, 63 Prionotus punctatus, 64 Prionotus roseus, 65 Prionotus rubio, 66 Prionotus scitulus, 67

16-T3324-IX1 9/13/05 11:22 AM Page 994

994 INDEX OF SCIENTIFIC NAMES

Prionotus streansi, 68 Prionotus tribulus, 69 Pristigenys alta, 215 Pristipomoides aquilonaris, 335 Pristipomoides macrophthalmus, 336 Prognathodes aculeatus, 468 Prognathodes aya, 470 Promethichthys prometheus, 762 Pronotogrammus martinicensis, 174 Psenes arafurensis, 807 Psenes cyanophrys, 808 Psenes pellucidus, 809 Pseudocaranx dentex, 289 Pseudoscopelus altipinnis, 560 Pseudoscopelus obtusifrons, 561 Pseudoscopelus scriptus, 562 Pseudupeneus maculatus, 451 Ptereleotris calliura, 726 Pterycombus brama, 308 Rachycentridae, 263 Rachycentron canadum, 264 Ranzania laevis, 942 Remora australis, 258 Remora brachyptera, 259 Remora osteochir, 260 Remora remora, 261 Remorina albescens, 262 Rhinesomus bicaudalis, 917 Rhinesomus triqueter, 918 Rhinochimaeridae, 14 Rhinoptera brasiliensis, 15 Rhinopteridae, 15 Rhomboplites aurorubens, 337 Risor rubber, 727 Ruvettus pretiosus, 763 Rypticus bistrispinus, 176 Rypticus maculatus, 177 Rypticus randalli, 178 Rypticus saponaceus, 179 Rypticus subbifrenatus, 180 Sarda sarda, 779 Scaridae, 527 Scartella cristata, 640 Scarus coelestinus, 533 Scarus coeruleus, 534 Scarus guacamaia, 535

Scarus iseri, 536 Scarus taeniopterus, 537 Scarus vetula, 538 Schultzea beta, 181 Sciaenidae, 410 Sciaenops ocellata, 443 Scomber colias, 780 Scomberomorus cavalla, 781 Scomberomorus maculatus, 782 Scomberomorus regalis, 783 Scombridae, 771 Scombrolabracidae, 745 Scombrolabrax heterolepis, 746 Scophthalmidae, 830 Scophthalmus aquosus, 831 Scorpaena agassizii, 37 Scorpaena albifimbria, 38 Scorpaena bergii, 39 Scorpaena brasiliensis, 40 Scorpaena calcarata, 41 Scorpaena dispar, 42 Scorpaena elachys, 43 Scorpaena grandicornis, 44 Scorpaena inermis, 45 Scorpaena plumieri, 46 Scorpaenidae, 26 Scorpaenodes caribbaeus, 47 Scorpaenodes tredecimspinosus, 48 Scyliorhinidae, 14 Scyliorhinus meadi, 14 Selar crumenophthalmus, 290 Selene brownii, 291 Selene setapinnis, 292 Selene vomer, 293 Seriola dumerili, 295 Seriola fasciata, 296 Seriola rivoliana, 297 Seriola zonata, 298 Serraniculus pumilio, 182 Serranidae, 119 Serranus annularis, 183 Serranus atrobranchus, 184 Serranus baldwini, 185 Serranus chionaraia, 186 Serranus luciopercanus, 187 Serranus notospilus, 188 Serranus phoebe, 189 Serranus subligarius, 190

16-T3324-IX1 9/13/05 11:22 AM Page 995

INDEX OF SCIENTIFIC NAMES 995

Serranus tabacarius, 191 Serranus tigrinus, 192 Serranus tortugarum, 193 Setarches guentheri, 49 Sicydium gymnogaster, 728 Sparidae, 385 Sparisoma atomarium, 539 Sparisoma aurofrenatum, 540 Sparisoma chrysopterum, 541 Sparisoma radians, 542 Sparisoma rubripinne, 543 Sparisoma viride, 544 Sphoeroides dorsalis, 924 Sphoeroides nephelus, 925 Sphoeroides pachygaster, 926 Sphoeroides parvus, 927 Sphoeroides spengleri, 928 Sphoeroides testudineus, 929 Sphyraena barracuda, 749 Sphyraena borealis, 750 Sphyraena guachancho, 751 Sphyraena picudilla, 752 Sphyraenidae, 747 Squalidae, 15 Starksia lepicoelia, 607 Starksia ocellata, 608 Starksia starcki, 609 Stathmonotus hemphillii, 622 Stathmonotus stahli, 623 Stegastes adustus, 498 Stegastes diencaeus, 499 Stegastes leucostictus, 500 Stegastes partitus, 501 Stegastes planifrons, 502 Stegastes variabilis, 503 Stellifer colonesis, 444 Stellifer lanceolatus, 445 Stenotomus caprinus, 404 Stephanolepis hispidus, 911 Stephanolepis setifer, 912 Stromateidae, 817 Syacium gunteri, 858 Syacium micrurum, 859 Syacium papillosum, 860 Symphurus billykrietei, 875 Symphurus civitatium, 876 Symphurus diomedeanus, 877 Symphurus marginatus, 878

Symphurus minor, 879 Symphurus parvus, 880 Symphurus pelicanus, 881 Symphurus piger, 882 Symphurus plagiusa, 883 Symphurus pusillus, 884 Symphurus stigmosus, 885 Symphurus urospilus, 886 Symphysanodon berryi, 340 Symphysanodon octoactinus, 341 Symphysanodontidae, 338 Synagrops bellus, 114 Synagrops spinosus, 115 Synagrops trispinosus, 116 Syngnathidae, 19 Taractes rubescens, 309 Taractichthys longipinnis, 310 Tetragonuridae, 815 Tetragonurus atlanticus, 816 Tetraodontidae, 919 Tetrapturus albidus, 793 Tetrapturus pfluegeri, 794 Thalassoma bifasciatum, 523 Thunnus alalunga, 784 Thunnus albacares, 785 Thunnus atlanticus, 786 Thunnus obesus, 787 Thunnus thynnus, 788 Trachinotus carolinus, 299 Trachinotus falcatus, 300 Trachinotus goodei, 301 Trachurus lathami, 302 Trachyscorpia cristulata, 50 Triacanthodidae, 889 Triakidae, 14 Trichiuridae, 764 Trichiurus lepturus, 770 Trichopsetta ventralis, 829 Triglidae, 51 Trinectes inscriptus, 870 Trinectes maculatus, 871 Tripterygiidae, 572 Umbrina coroides, 446 Upeneus parvus, 452 Uranoscopidae, 567 Uraspis secunda, 303

16-T3324-IX1 9/13/05 11:22 AM Page 996

996 INDEX OF SCIENTIFIC NAMES

Varicus marilynae, 729 Xanthichthys ringens, 900 Xiphias gladius, 795 Xiphiidae, 789

Xyrichtys martinicensis, 524 Xyrichtys novacula, 525 Xyrichtys splendens, 526 Zoarcidae, 545

17-T3324-IX2 9/13/05 11:22 AM Page 997

INDEX OF COMMON NAMES

African pompano, 274 Albacore, 784 Almaco jack, 297 American sole, 864 Anchor tilefish, 247 Angelfish, 471 Anglefin whiff, 843 Apricot bass, 173 Armored searobin, 80 Armored searobins, 70 Arrow stargazer, 582 Atlantic bonito, 779 Atlantic bumper, 281 Atlantic croaker, 435 Atlantic cutlassfish, 770 Atlantic moonfish, 292 Atlantic sailfish, 791 Atlantic spadefish, 736 Atlantic thornyhead, 50 Atlantic threadfin, 407 Balloonfish, 937 Banded blenny, 604 Banded butterflyfish, 466 Banded drum, 429 Banded jawfish, 205 Banded rudderfish, 298 Bandtail puffer, 928 Bandtail searobin, 62 Bank butterflyfish, 470 Bank seabass, 133 Banner blenny, 616 Banner goby, 720 Bar jack, 280 Barbfish, 40 Barbu, 409 Barracudas, 747

Barred blenny, 630 Barred cardinalfish, 222 Barred grunt, 366 Barred hamlet, 160 Barred searobin, 60 Barrelfish, 800 Bartail goby, 686 Basslets, 194 Bay whiff, 846 Bearded goby, 669 Beau gregory, 500 Belted cardinalfish, 228 Belted sandfish, 190 Bermuda chub, 484 Bicolor damselfish, 501 Bigeye, 214 Bigeye jack, 290 Bigeye mojarra, 352 Bigeye searobin, 59 Bigeye squaretail, 816 Bigeye tuna, 787 Bigeyes, 211 Bighead searobin, 69 Bigmouth sleeper, 659 Bigscale pomfret, 310 Bigtooth cardinalfish, 220 Billfishes, 789 Black driftfish, 799 Black drum, 442 Black durgon, 899 Black gemfish, 761 Black grouper, 166 Black grunt, 369 Black jack, 279 Black margate, 364 Black ruff, 798 Black scabbardfish, 766

17-T3324-IX2 9/13/05 11:22 AM Page 998

998 INDEX OF COMMON NAMES

Black seabass, 135 Black snake mackerel, 759 Black snapper, 321 Blackbar drum, 440 Blackbelly blenny, 622 Blackbelly rosefish, 31 Blackcheek blenny, 607 Blackcheek tonguefish, 883 Blackear bass, 184 Blackear wrasse, 520 Blackfin blenny, 606 Blackfin cardinalfish, 230 Blackfin snapper, 325 Blackfin tuna, 786 Blackhead blenny, 619 Blackline tilefish, 246 Blackmouth bass, 114 Blackwing searobin, 66 Blotchfin tonguefish, 885 Blue angelfish, 474 Blue chromis, 492 Blue croaker, 417 Blue goby, 726 Blue marlin, 792 Blue parrotfish, 534 Blue runner, 276 Blue tang, 744 Bluefin driftfish, 809 Bluefin tuna, 788 Bluefish, 252 Bluefishes, 251 Bluegold goby, 717 Bluehead, 523 Blueline tilefish, 248 Bluelip parrotfish, 531 Bluenose goby, 683 Bluespotted searobin, 65 Bluestriped grunt, 378 Bluethroat pikeblenny, 614 Blunthead puffer, 926 Bluntnose jack, 286 Boga, 384 Bonnetmouth, 383 Bonnetmouths, 382 Bridle cardinalfish, 221 Bridled burrfish, 932 Bridled goby, 681 Broad flounder, 857 Bronze cardinalfish, 229

Brown driftfish, 813 Bucktooth parrotfish, 542 Bullet mackerel, 775 Burro grunt, 381 Butter hamlet, 162 Butterfishes, 817 Butterflyfishes, 458 Caesar grunt, 371 Cardinalfishes, 216 Cero, 783 Chalk bass, 193 Channel flounder, 859 Checkered blenny, 608 Checkered puffer, 929 Cherubfish, 473 Chocolate banded tonguefish, 875 Chub mackerel, 780 Clingfishes, 641 Clown goby, 719 Clown wrasse, 518 Cobia, 264 Cobias, 263 Cocoa damselfish, 503 Code goby, 711 Colon goby, 679 Colon stardrum, 444 Comb grouper, 165 Combtooth blennies, 624 Conchfish, 231 Coney, 138 Convict goby, 716 Coral blenny, 603 Coral scorpionfish, 38 Cottonmouth jack, 303 Cottonwick, 375 Cowfishes, 913 Creole wrasse, 510 Creolefish, 172 Crested goby, 713 Crevalle jack, 277 Crimson rover, 316 Cubbyu, 441 Cubera snapper, 327 Cutlassfishes, 764 Damselfishes, 487 Darter goby, 687 Dash goby, 690

17-T3324-IX2 9/13/05 11:22 AM Page 999

INDEX OF COMMON NAMES 999

Deepreef scorpionfish, 48 Deepwater cardinalfishes, 235 Deepwater flounder, 828 Deepwater tonguefish, 882 Doctorfish, 743 Dog snapper, 329 Dolphin, 268 Dolphinfishes, 265 Dotterel filefish, 903 Downy blenny, 595 Draconetts, 652 Dragonets, 646 Driftfishes, 801 Drums, 410 Duckbill flathead, 564 Dusky basslet, 197 Dusky blenny, 599 Dusky cardinalfish, 233 Dusky damselfish, 498 Dusky flounder, 860 Dusky jawfish, 210 Dwarf goatfish, 452 Dwarf scorpionfish, 43 Dwarf seabass, 140 Dwarf wrasse, 512 Eelgrass blenny, 623 Eelpouts, 545 Emerald clingfish, 643 Emerald goby, 692 Emerald parrotfish, 532 Emerald sleeper, 658 Fat sleeper, 656 Fat snook, 101 Feather blenny, 633 Filefishes, 901 Flagfin mojarra, 355 Flamefish, 224 Flatheads, 563 Flecked pikeblenny, 615 Florida blenny, 628 Florida pompano, 299 Flying gurnard, 25 Flying gurnards, 24 Foureye butterflyfish, 460 Fourspot flounder, 854 Freckled blenny, 635 Freckled cardinalfish, 232

Freckled driftfish, 808 Freckled soapfish, 176 Freckled stargazer, 570 French angelfish, 481 French grunt, 373 Freshwater goby, 691 Frigate mackerel, 776 Frillfin goby, 672 Fringed filefish, 909 Fringed flounder, 850 Fringed sole, 869 Gag, 168 Glass blenny, 620 Glass goby, 682 Glasseye snapper, 213 Glassy sweeper, 454 Goatfishes, 447 Gobies, 661 Goby flathead, 565 Goldface tilefish, 245 Goldface toby, 921 Goldline blenny, 598 Goldspot goby, 703 Goosehead scorpionfish, 39 Grass porgy, 391 Gray angelfish, 479 Gray flounder, 852 Gray snapper, 328 Gray triggerfish, 895 Graysby, 137 Great barracuda, 749 Greater amberjack, 295 Greater soapfish, 179 Green goby, 721 Green razorfish, 526 Greenband wrasse, 513 Greenblotch parrotfish, 539 Grunts, 359 Guaguanche, 751 Gulf butterfish, 818 Gulf flounder, 855 Gulf kingfish, 432 Gulf stream flounder, 840 Hairy blenny, 597 Harlequin bass, 192 Harvestfish, 819 Hawkfishes, 485

17-T3324-IX2 9/13/05 11:22 AM Page 1000

1000 INDEX OF COMMON NAMES

Highfin blenny, 636 Highfin goby, 705 Highfin scorpionfish, 36 High-hat, 438 Hogchoker, 871 Hogfish, 522 Honeycomb cowfish, 914 Horned searobin, 57 Horned whiff, 841 Horse-eye jack, 278 Hunchback scorpionfish, 42 Irish pompano, 347 Island frillfin, 671 Island goby, 715 Jackknife-fish, 424 Jacks, 269 Jambeau, 892 Jawfishes, 199 Jewfish, 146 Jolthead porgy, 392 Keelcheek bass, 115 Key blenny, 609 King mackerel, 781 Knobbed porgy, 396 Lancer dragonet, 651 Lancer stargazer, 571 Lancetail wormfish, 733 Lane snapper, 332 Lantern bass, 185 Largescale tonguefish, 879 Least puffer, 927 Leatherjack, 288 Lefteye flounders, 821 Leopard searobin, 67 Lesser amberjack, 296 Lined sole, 866 Little tunny, 777 Littlehead porgy, 399 Littlescale threadfin, 408 Lofty triplefin, 574 Longbill spearfish, 794 Longfin blenny, 594 Longfin damselfish, 499 Longfin scorpionfish, 37

Longsnout butterflyfish, 468 Longspine porgy, 404 Longspine scorpionfish, 35 Longtail bass, 154 Longtail tonguefish, 881 Lookdown, 293 Louvar, 739 Louvars, 738 Lyre goby, 702 Mackerel scad, 282 Mackerels and tunas, 771 Mahogany snapper, 330 Manefishes, 311 Mangrove blenny, 637 Man-of-war fish, 806 Marbled blenny, 605 Marbled grouper, 139 Marbled puffer, 924 Margate, 367 Margined tonguefish, 878 Marked goby, 693 Marlinsucker, 260 Masked goby, 684 Masked stargazer, 583 Mauve goby, 724 Medusafishes, 796 Mexican flounder, 848 Mexican searobin, 63 Midnight parrotfish, 533 Mimic blenny, 593 Mimic triplefin, 576 Misty grouper, 148 Mojarras, 344 Molly miller, 640 Mottled jawfish, 206 Mottled mojarra, 354 Moustache jawfish, 204 Mushroom scorpionfish, 45 Mutton hamlet, 128 Mutton snapper, 323 Naked goby, 706 Naked sloe, 867 Nassau grouper, 151 Neon goby, 697 Night sergeant, 491 Northern kingfish, 433

17-T3324-IX2 9/13/05 11:22 AM Page 1001

INDEX OF COMMON NAMES 1001

Northern sennet, 750 Northern tonguefish, 884 Notchtongue goby, 670

Purple chromis, 496 Pygmy seabass, 182 Pygmy tonguefish, 880

Ocean sunfish, 941 Ocean sunfishes, 939 Ocean surgeon, 742 Ocean triggerfish, 898 Ocellated flounder, 838 Ocular flounder, 824 Offshore tonguefish, 876 Oilfish, 763 Orange filefish, 905 Orangeback bass, 183 Orangespotted filefish, 908 Orangespotted goby, 722

Queen angelfish, 476 Queen parrotfish, 538 Queen snapper, 322 Queen triggerfish, 896

Painted wrasse, 515 Pale cardinalfish, 225 Paleback goby, 712 Palehead blenny, 592 Pallid goby, 680 Palometa, 301 Parrotfishes, 527 Pearl blenny, 629 Pearly razorfish, 525 Pelagic porcupinefish, 936 Pelican flounder, 826 Peppermint bass, 164 Permit, 300 Pigfish, 380 Pigmy filefish, 912 Pilotfish, 287 Pinfish, 402 Pink wormfish, 734 Pirate blenny, 618 Planehead filefish, 911 Plumed scorpionfish, 44 Pomfrets, 304 Pompano dolphin, 267 Porcupinefish, 938 Porcupinefishes, 930 Porgies, 385 Porkfish, 365 Princess parrotfish, 537 Puddingwife, 521 Puffcheek blenny, 591 Puffers, 919

Ragged goby, 674 Rainbow parrotfish, 535 Rainbow runner, 285 Rainbow wrasse, 519 Red barbier, 155 Red goatfish, 450 Red grouper, 147 Red hind, 145 Red hogfish, 511 Red porgy, 403 Red snapper, 326 Redband parrotfish, 540 Reddrum, 443 Redeye triplefin, 577 Redfin parrotfish, 543 Redlip blenny, 638 Redspotted hawkfish, 486 Redtail parrotfish, 541 Redtail scad, 284 Reef butterflyfish, 464 Reef croaker, 436 Reef scorpionfish, 47 Remora, 261 Remoras, 253 Reticulate spikefish, 890 Rimspine searobin, 81 River goby, 668 Rock beauty, 478 Rock hind, 142 Rock seabass, 134 Rockcut goby, 708 Rosy blenny, 600 Rosy razorfish, 524 Rough scad, 302 Rough triggerfish, 897 Roughhead blenny, 612 Roughhead triplefin, 575 Roughtongue bass, 174 Round scad, 283

17-T3324-IX2 9/13/05 11:22 AM Page 1002

1002 INDEX OF COMMON NAMES

Rovers, 313 Royal gramma, 196 Rugjaw wormfish, 731 Rusty goby, 725 Sackfish, 756 Saddle bass, 188 Saddle stargazer, 585 Saddled blenny, 601 Sailfin blenny, 617 Sailors choice, 376 Sand drum, 446 Sand perch, 141 Sand seatrout, 419 Sand stargazer, 581 Sand stargazers, 578 Sand tilefish, 250 Sargassum triggerfish, 900 Sash flounder, 829 Saucereye porgy, 393 Sawcheek cardinalfish, 227 Scaly blennies, 586 Scamp, 169 School bass, 181 Schoolmaster, 324 Scorpionfishes, 26 Scrawled cowfish, 915 Scrawled filefish, 906 Scrawled sole, 870 Sea bream, 390 Sea chubs, 482 Seabasses, 119 Seaboard goby, 707 Searobins, 51 Seaweed blenny, 639 Seminole goby, 718 Sergeant major, 490 Sharksucker, 255 Sharpnose puffer, 922 Sharptail mola, 940 Sheepshead, 389 Sheepshead porgy, 397 Shelf flounder, 851 Shoal flounder, 858 Short bigeye, 215 Short-tube scorpionfish, 34 Shortwing searobin, 68 Shrimp flounder, 853

Silk snapper, 333 Silver driftfish, 807 Silver jenny, 350 Silver perch, 415 Silver pinfish, 400 Silver rag, 812 Silver seatrout, 422 Skilletfish, 645 Skipjack, 778 Sleepers, 654 Slender filefish, 910 Slender mojarra, 353 Slender mola, 942 Slender suckerfish, 257 Slippery dick, 514 Smallmouth grunt, 372 Smooth puffer, 923 Smooth trunkfish, 918 Smoothhead scorpionfish, 41 Snake mackerel, 757 Snake mackerels, 753 Snappers, 317 Snook, 104 Snooks, 97 Snow bass, 186 Snowy grouper, 150 Southern flounder, 856 Southern kingfish, 431 Southern puffer, 925 Southern sennet, 752 Southern stargazer, 569 Spadefishes, 735 Spanish flag, 152 Spanish grunt, 374 Spanish hogfish, 509 Spanish mackerel, 782 Spearfish remora, 259 Speckled hind, 143 Speckled stargazer, 580 Spikefishes, 889 Spiny flounder, 827 Spiny searobin, 58 Spinycheek scorpionfish, 33 Spinycheek sleeper, 657 Sponge cardinalfish, 234 Sponge goby, 701 Spot, 430 Spotcheek blenny, 596

17-T3324-IX2 9/13/05 11:22 AM Page 1003

INDEX OF COMMON NAMES 1003

Spotfin butterflyfish, 462 Spotfin dragonet, 649 Spotfin flounder, 849 Spotfin goby, 723 Spotfin hogfish, 508 Spotfin jawfish, 209 Spotfin mojarra, 349 Spotlight parrotfish, 544 Spottail goby, 694 Spottail pinfish, 401 Spottail tonguefish, 886 Spotted burrfish, 934 Spotted dragonet, 648 Spotted driftfish, 814 Spotted drum, 426 Spotted goatfish, 451 Spotted goby, 685 Spotted scorpionfish, 46 Spotted seatrout, 421 Spotted soapfish, 180 Spotted spikefish, 891 Spotted trunkfish, 917 Spotted whiff, 845 Spottedfin tonguefish, 877 Spotwing scorpionfish, 32 Star drum, 445 Stargazers, 567 Stippled clingfish, 644 Streamer bass, 153 Streamer searobin, 56 Striped bass, 109 Striped burrfish, 935 Striped croaker, 418 Striped grunt, 379 Striped mojarra, 357 Striped parrotfish, 536 Sunshinefish, 494 Surgeonfishes, 740 Sweepers, 453 Swordfish, 795 Swordspine snook, 99 Swordtail jawfish, 202 Tarpon snook, 102 Tattler, 189 Temperate basses, 105 Temperate ocean-basses, 111 Tessellated blenny, 634

Threadfin blenny, 602 Threadfins, 405 Threadnose bass, 130 Three-eye flounder, 837 Threeline basslet, 198 Threespot damselfish, 502 Tidewater mojarra, 351 Tiger goby, 696 Tiger grouper, 170 Tilefish, 249 Tilefishes, 243 Tobaccofish, 191 Tomtate, 368 Tonguefishes, 872 Triggerfishes, 893 Tripletail, 343 Tripletails, 342 Trunkfish, 916 Tube blennies, 610 Tusked goby, 727 Twoscale goby, 710 Twospot cardinalfish, 226 Two-spot flounder, 825 Unicorn filefish, 904 Vermilion snapper, 337 Violet goby, 704 Wahoo, 774 Warsaw grouper, 149 Warteye stargazer, 584 Weakfish, 423 Web burrfish, 933 Wenchman, 335 Whalesucker, 258 White bass, 107 White grunt, 377 White marlin, 793 White suckerfish, 262 Whitebone porgy, 395 Whitecheek blenny, 590 White-eye goby, 673 Whitefin sharksucker, 256 Whitespotted filefish, 907 Whitespotted soapfish, 177 Whitestar cardinalfish, 223 Windowpane, 831

17-T3324-IX2 9/13/05 11:22 AM Page 1004

1004 INDEX OF COMMON NAMES

Wormfishes, 730 Wrasse bass, 163 Wrasse blenny, 621 Wrasses, 504 Yellow bass, 108 Yellow chub, 483 Yellow goatfish, 449 Yellow jack, 275 Yellowcheek wrasse, 516 Yellowedge cromis, 495 Yellowedge grouper, 144 Yellowface pikeblenny, 613 Yellowfin bass, 129

Yellowfin grouper, 171 Yellowfin mojarra, 358 Yellowfin tuna, 785 Yellowhead jawfish, 203 Yellowhead wrasse, 517 Yellowline goby, 695 Yellowmouth grouper, 167 Yellowmouth jawfish, 207 Yellowprow goby, 700 Yellowtail bass, 132 Yellowtail damselfish, 497 Yellowtail hamlet, 157 Yellowtail reeffish, 493 Yellowtail snapper, 334