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SMITHS' SEA SMITHS' SEA FISHES

---~-- --. ~-- -- @ ------J -~ ------.~­ -- .------.:..-- -~-.------. fDlHD BY MARGARET M. SM ITH AND PHilliP C. HEEMSTRA

Springer-verlag Berlin Heidelberg New York London Paris Tokyo Dedicated to J .L.B. Smith who loved angling and devoted his life to the study of fishes .

Distributio n rights all over the world except for the area of Southern Springer-Verl3g Berlin Heidelberg New York London Paris Tot yo

ISBN 978·3·642·82860·7 ISBN 978-3-642-82858-4 (cBook) 00110.10071978-3-642-82858-4

Copyright 0 1986 by J.L.B. Smith Institute of Ichthyology. GrahamSlown. Softcover reprint of ttle hardcover 6th edition 1986

All righ ts reserved. No part of this publication may be reproduced or transmitted in any form or by any means without prior wriuen permission from the publisher.

Jackel design by Graphitor. after an idea of David Voorvcll.

CTP Book Printers, Cape. Foreword Dr. D. S. Henderson, Chairman of the 1. L. B. Smith Institute of Ichthyology and Vice Chancellor of Rhodes University

This book is a unique, international collaborative effort ranging all of the colour plates for the book. For the past of 76 scientists, representing 15 countries. Several skilled eight years, she has been assisted in the research, writing artists and photographers have also contributed to the and editing of the book by Dr. P. C. Heemstra. numerous and beautiful illustrations. Research done in It is essential for the proper management of the marine South Africa was supported by the Council for Scientific resources of southern Africa that we know what and Industrial Research. Work on the book at the J.L.B. we are dealing with. This book will greatly facilitate the Smith Institute was also funded by the Department of identification of our fishes. It will thus be of vital import• National Education and The Trustees of the Sea Fishes of ance to the understanding and wise use of our tremendously Southern Africa Book Fund. Expenses of visits by ichthy• diverse and valuable fish fauna. ologists from overseas were defrayed by a grant from the We are proud of the affiliation of the J.L.B. Smith Insti• John S. Schlesinger Foundation. tute with Rhodes University and happy to see the successful Although this book is an impressive contribution to the completion of this long-awaited work. zoology of southern Africa, few people will realise the enormous amount of work that has gone into it. Prof. Mar• garet M. Smith has laboured for thirty years: first, to build the foundation of the largest fish collection in Africa and the comprehensive, superbly organised library necessary for the research; second, to lead the project that produced the Dr. D. S. Henderson, book; and third, to participate in the actual research, writ• Chairman of the Institute and ing, editing, production of most of the illustrations, and ar- Vice-Chancellor of Rhodes University

1. P. de Wit, Deputy President, CSIR

When J. L. B. Smith's monumental book The Sea Fishes considered advisable to enlist the assistance of other of Southern Africa was first published in 1949, the first ichthyologists who are experts on the various families of printing was sold out within three weeks. Due in large meas• marine fishes represented in the seas of southern Africa. ure to the excellent and numerous illustrations (mostly by This team effort has produced a uniquely authoritative and Margaret Smith), the book has been extremely popular in comprehensive book on the marine fishes of southern Af• South Africa and also much in demand overseas. It has be• rica. To all of the many collaborators, we owe a debt of come an international ichthyological "classic" and is well gratitude for their unstinting efforts to provide the latest known in museums and marine laboratories all over the and best fruits of their ichthyological research. world. In 1953 and 1961, J. L. B. Smith added a few addi• With this book, on which she laboured for over 15 years, tional species in appendices at the end of the book. Since Margaret Smith has produced not only a worthy tribute to 1961, many new species and new records of fishes have been her late husband's great achievements, but also a lasting discovered in the seas off southern Africa. Also, recent testimonial to her own dedication to the fulfilment of his ichthyological research has considerably altered the classifi• ideals. cation of marine fishes. It thus became apparent that the Sea Fishes book was in need of a thorough revision. Prof. Margaret Smith and Dr P. C. Heemstra began work on the revision in 1978. In view of the specialisation that has taken place in ichthyology (as in science generally), it was Contents

List of subscribers ...... ix Acknowledgements ...... xi

Contributors ...... 0 • • • xii Systematic contents ...... xiv Introduction ...... 1 Anatomy of fishes ...... 2 Biology of fishes ...... 6 Dangers of the sea ...... 17 The effect of fishing ...... 19 Common names ...... 20 Scientific nomenclature and classification ...... 21 Oceanography of the Southern African region ...... 22 Ichthyology in South Africa ...... 28 Place names and their localities: map ...... 30 Systematic section Key to superclasses, classes and subclasses ...... 31 Class Pteraspidomorphi ...... 33 Class Chondrichthyes (cartilaginous fishes) ...... 39 Class (bony fishes) ...... 149 Colour plate section ...... 908 Reference section Fin formula key to bony fishes ...... 909 Glossary ...... 939 Bibliography ...... 945 Number conversion table ...... 987 Scientific index ...... 993 English common name index ...... 1029 Afrikaans common name index ...... 1039 THE SEA FISHES TRUST

Chairman: Dr D. S. Henderson Secretary/Treasurer: C. A. Lanham

Professor M. M. Smith Professor M. N. Bruton G. A. Harvey W. M. Smith Dr I. Mackenzie Dr B. E. Hersov A. R. Thorpe

Sponsors, Donors and Subscribers

Corporate Sponsors Anglo American Corporation of South Africa Ltd Gold Fields De Beers Consolidated Mines Ltd Mrs Joan Le May: in memory of the late De Beers Industrial Diamond Division (Pty) Ltd Basil C. Le May

Corporate Donors Barclays National Bank Ltd Lennon Ltd Fisheries Development Corporation of Natal Parks Board South Africa Ltd Professor Margaret M. Smith The Gencor Development Fund William and Jenny Smith Hewlett Packard South Africa (Pty) Ltd Total South Africa (Pty) Ltd

Corporate Subscribers Anglovaal Ltd National Parks Board Atlantic Fishing Enterprises Oceanographic Research Institute, Durban Barlow Rand Limited SAPPI Fine Papers (Pty) Ltd CTP Book Printers (Pty) Ltd Smithsonian Institution Durban City Council The South African Association for Hans Merensky Holdings (Pty) Ltd Marine Biological Research Irvin & Johnson Ltd South African Permanent Building Society Metal Box South Africa Ltd Standard Bank Investment Corporation Monaco Institute of Oceanographic Research A. R. Thorpe E. J. and A. L. Morris Wildlife Society of Southern Africa

Individual Donors Adcan Marine Suppliers Hedley Gerhardt Africana Book Collectors Pat Goss Dr B. R. Allanson Henry J. Gradwell Trevor Ashington Dr D. S. Henderson L. G. Baumann Basil Hersov The Brenthurst Library Dr Brian Hulley Professor J. W. Brommert Isowall SA (Pty) Ltd Craig, Ryan and Tracey Bruton Alf Johnson David Christie Jessica and Len Jones Pearl E. Coppin Errol and Fredia Kreutzer Bert Davids Helen Leibel Claire Davids Robert E. Levitt H. B. Dyer Ian Mackenzie Johan Els Natal Ski-Boat Light Tackle Game Fishing Professor P. K. Faure Association

ix Individual Donors Richards Bay Minerals George Brian Trow Mrs L. M. Russell G. H. Tudor S.A. Shipping News & Fishing Industry Review Umhlali Ski-Boat Club Derek and Linda Souter Pieter W. van Niekerk South African Ski-Boat Angling Association Zoological Society of Southern Africa Nolan Sparg

Individual Subscribers Christopher C. Appleton City of Johannesburg Peter J. Ashton Marthinus J. S. Jordaan Eugene K. Balon Professor D. M. Joubert E. J. B. Bishop· Hillel M. Kahn Border Ski-Boat Angling Association Sidney H. Kahn Alan and Jiff Bowmaker Dr P. G. Landsberg The Braine family Dr B. E. la Trobe, Mayor Grahamstown 1985/86 George and Margo Branch R. H. Lefebvre Alastair and Patricia Buchan In memory of the late Dr C. Mauritz Lombaard Ian D. J. Cochrane-Murray R. Loots Dr Allan Connell W. McLevland Douglas T. Cooke E. K. Moorcroft Mrs H. J. Creswell Natal Ski-Boat Association Brendan Davids National Zoological Gardens of South Africa John-Paul Davids Dan Niven Thomas Davids Patrick Niven Durban Ski-Boat Club Oceana Fishing Group Ltd The Mayor of East London Mike and Mary-Lou Penrith East London Museum Robin G. Piper East London Ski-Boat Club Port Alfred Municipality D. H. Eccles Gavin, Howard, Charles, Mark and Jamie Pote Karl Zachary Edwards R. McC Pott Lloyd Edwards Ray Preen John Dudley Engelbrecht Requin Angling Club S.A. Director, Fishing Industry Research Institute Dr G. J. B. Ross Professor Vernon S. Forbes and W. H. Wood SAICCOR (Pty) Ltd Mayor, Councillors and Citizens of Fort Beaufort R. Allan Scott Eugene and Lalie Fourie Sea Fisheries Research Institute L. P. Dormehl Gertenbach C. P. Skinner M. C. V. Ginn Rodney P. Smith Gonubie Marine Club W. H. Stoops Ken Gough Sandy Stretton Dr C. E. and Mrs D. Gow Oscar B. Susman Grahamstown Municipality Swaziland Underwater Group J. T. Hart Professor John Tew Glen Harvey H. P. Thesen Alan S. Henderson Umzimkulu Ski-Boat Club Ian M. Henderson Rudy van der Elst Ethnee Joy Hepburn (nee Amm) Dr John H. Wallace Viktor E. Hesse E. G. Wanliss Dr P. Alexander Hulley Mr and Mrs T. N. Webb Humansdorp Municipality City of Welkom Indian Export Company (Pty) Ltd Mr and Mrs K. C. Whyte P. B. N. Jackson Mr and Mrs J. O. M. Wilkinson J. W. Jagger Library, University of Cape Town Alastair and Gerry Williams x Acknowledgements

It is impossible to mention all of the many people who to collect fishes for us. In some cases these specimens have assisted in the completion of this book. We are in• have represented new species or new records for southern debted to our numerous collaborators for their contribu• Africa. We are especially grateful to A. R. Thorpe, tions and their patience in the long period that it has Dave Bickell, Rudy van der Elst, Malcolm Smale, Alan taken to complete this work. We are grateful to our as• Connell, Colin Buxton, Dennis King, Len Jones, Alf sistant, Mrs Joan Wright, whose careful work, industry Johnson, Elizabeth Tarr, Rob Tarr, Brian Trow, Hennie and skill with the word processor have been of tremend• Kok, Wouter Holleman, Anton Bok, Tom Hecht, ous help to us. We thanks Mrs Jean Pote for her unique Robin Stobbs, Mike Bruton and Lynneth Beckley. We ability to decipher the illegible handwriting and editorial also thank the East London and Port Elizabeth ciphers presented to her; Mrs Pote has also done an Museums and the Oceanographic Research Institute in enormous job on the word processor. Durban for their efforts in sending specimens. We are Special thanks and credit are due to the artists who indebted to P. A. Hulley and S. X. Kannemeyerfor their have contributed the paintings and drawings that illus• prompt loans of material from the South African trate this work. We thank Elizabeth Tarr for her enorm• Museum and for providing data on various fishes in their ous contribution; Ms Tarr's accuracy and attention to collection. detail are exemplary. We are much indebted to David Most of the funds for the work done in Grahamstown Voorvelt, whose remarkable industry and skill as an ar• were provided by the South African Council for Scien• tist have been essential to the completion of this work. tific and Industrial Research (CSIR) through the South We are also grateful to the other fine artists who have African National Committee for Oceanographic Re• made drawings for us: Jean Michel Vincent, Virginia search. Additional funds were provided by the Smiths' McRostie, Anne Lefebvre, Moira Lambert, and from Sea Fishes Trust and Rhodes University. We are also the original Sea Fishes of Southern Africa: Patricia Par• grateful for the excellent facilities and resources of the kin, Denys M. Davis and Hester W. Locke. We are J.L.B. Smith Institute of Ichthyology and for the support especially grateful to John E. Randall for numerous of the Department of National Education. transparencies used in the colour plates. Gerald R. We thank the members of our Editorial Committee, Allen, Alan Emery, John Gunn, Sidney Kannemeyer, Brian R. Allanson, P. A. Hulley, John Wallace and Dennis King, Daniel Pelicier, A. R. Thorpe, Peter van A. R. Thorpe, for their valuable advice. We are also grate• Niekerk and Richard Winterbottom also supplied some ful for the excellent editorial assistance of Frances Perryer, transparencies used in this book. Matthew Seal and Basil van Rooyen of Macmillan, S.A. We are grateful to Robin Stobbs, whose abilities as a For donations of various kinds, we thank the Fisheries photographer, small boat captain, laboratory techni• Development Corporation, Sasolchem (Pty) Ltd and cian, computor operator and field assistant have been of the Hewlett-Packard Co. immeasurable help to us. We thank Billy Ranchod for his careful and excellent work as our collection manager. We are indebted to the Heemstra family (Valerie, Mrs Doris Cave, who was the Institute's librarian for Lydia and Julia) for much encouragement, help of vari• twenty-five years, assisted in the compilation of the bib• ous kinds, and their forbearance in tolerating the many liography and generally facilitated our work by main• weekends and evenings their father and husband de• taining a well-organised and superbly indexed library. voted to this book. Throughout the years the public, including the angling Finally, we wish to express our gratitude to the direc• fraternity, naturalists, divers and school children, have tor of our Institute, Prof. M. N. Bruton, for his constant sent in numerous specimens and contributed much useful support, advice and assistance in the completion of the information. Several persons have made a special effort manuscript.

xi Contributors

Allen, G. R. Western Australian Museum, Francis Street, Gomon, J. R. Fish Division, National Museum of Natural Perth, WA 6000 (Australia). History, Washington, D.C. 20560 (USA). Anderson, M. E. Dept. of Ichthyology, California Academy Gon, O. J.L.B. Smith Institute of Ichthyology, Private Bag of Sciences, Golden Gate Park, San Francisco, CA 1015, Grahamstown 6140 (South Africa). 94118 (USA). Goodyear, R. H. Centro de Ciencias del Mar y Limnologia, Anderson, W. D. Jr. Grice Marine Biological Lab, 205 Fort Universidad de Panama (Republic of Panama). Johnson, Charleston, SC 29407 (USA). Haedrich, R. L. Memorial University of Newfoundland, St. Badcock, J. Institute of Oceanographic Sciences, Brook Johns, Newfoundland AlB 3X9 (Canada). Road, Godalming, Surrey GU8 5UB (England). Hensley, D. A. Department de Ciencias Marinas, Universidad Baird, R. C. Hayward-Schuster Woolen Mills Inc., East de Puerto Rico, Mayagiiez, Puerto Rico 00708 (USA). Douglas, MA 01516 (USA). Heemstra, P. C. J.L.B. Smith Institute of Ichthyology, Private Bass, A. J. Western Australian Museum, Francis Street, Bag 1015, Grahamstown 6140 (South Africa). Perth, WA 6000 (Australia). Hoese, D. F. The Australian Museum, P.O. Box A285, Syd• Ben-Tuvia, A. Dept. of Zoology, the Hebrew University of ney, NSW 2000 (Australia). Jerusalem (Israel). Holleman, W. Albany Museum, Grahamstown 6140 (South Bertelsen, E. Zoologisk Museum, Universitetsparken 15, Africa). DK 2100 Kl'lbenhavn (Denmark). Hulley, P. A. Dept. of Marine Biology, South African Bradbury, M. G. Dept. of Marine Biology, San Francisco Museum, P.O. Box 61, Cape Town 8000 (South Africa). State College, San Francisco, CA 94132 (USA). Hutchins, J. B. Western Australian Museum, Francis Street, Bray, D. J. The Australian Museum, P.O. Box A285, Sydney Perth, WA 6000 (Australia). South, NSW 2000 (Australia). IvantsotT, W. School of Biological Sciences, Macquarie Uni• Briggs, J. C. Dept. of Marine Sciences, University of South versity, North Ryde, NSW2113 (Australia). Florida, St. Petersburg, FL 33701 (USA). Iwamoto, T. Dept. of Ichthyology, California Academy of Caruso, J. H. Museum of Natural History, Tulane University, Sciences, Golden Gate Park, San Francisco, CA 94118 Belle Chasse, LA 70037. (USA). Castle, P. H. J. Dept. of Zoology, Victoria University, Private Johnson, R. K. Division of Fishes, Field Museum of Natural Bag, Wellington (New Zealand). History, Roosevelt Road at Lake Shore Drive, Chicago, Cohen, D. M. Division of Life Sciences, L.A. County IL 60605 (USA). Museum of Natural History, 900 Exposition Blvd, Los Kannemeyer, S. X. Dept. of Marine Biology, South African Angeles, CA 90007 (USA). Museum, P.O. Box 61, Cape Town 8000 (South Africa). Collette, B. B. Systematics Laboratory, National Marine Karrer, C. Ichthyologie Seefischerei, Zoologisches Institut Fisheries Service, National Museum of Natural History, und Museum, Martin-Luther-King-Platz 3, D-2000 Washington, D.C. 20560 (USA). Hamburg 13 (Federal Republic of Germany). Compagno, L. J. V. J. L. B. Smith Institute of Ichthyology, Keene, J. Route 1, Box 262, Shipman, VA 22971 (USA). Private Bag 1015, Grahamstown 6140 (South Africa). Knapp, L. W. Smithsonian Oceanographic Sorting Center, Cressey, R. F. Crustacea Division, National Museum of Washington, D.C. 20560 (USA). Natural History, Washington, D.C. 20560 (USA). KretTt, G. Ichthyologie Seefischerei, Zoologisches Institut Dawson, C. E. Gulf Coast Research Laboratory, P.O. Box und Museum, Martin-Luther-King-Platz 3, D-2000 218, Ocean Springs, MS 39564 (USA). Hamburg 13 (Federal Republic of Germany). de Sylva, D. E. School of Marine & Atmospheric Sciences, Leis, J. M. Fish Dept., The Australian Museum, P.O. Box University of Miami, 4600 Rickenbacker Causeway, A285, Sydney, NSW 2000 (Australia). Miami, FL 33149 (USA). McCosker, J. E. Steinhart Aquarium, California Academy Ebeling, A. W. Dept. of Biological Sciences, University of of Sciences, Golden Gate Park, San Francisco, CA California, Santa Barbara, CA 93106 (USA). 94118 (USA). Eschmeyer, W. N. Dept. ofIchthyology, California Academy McKay, R. J. Queensland Museum, Gregory Terrace, For• of Sciences, Golden Gate Park, San Francisco, CA 94118 titude Valley, QLD 4006 (Australia). (USA). Markle, D. F. Dept. of Fisheries & Wildlife, Oregon State Fernholm, B. Zoology Research Dept., Swedish University, Corvallis, OR 97331 (USA). Museum of Natural History, S-104 05 Stockholm (Swe• Martin, T. J. Dept. of Zoology, University of Natal, Durban den). (South Africa). Fricke, R. Saalestr. 3A, D-3300 Braunschweig (Federal Re• Matsui, T. Scripps Institution of Oceanography, La Jolla, public of Germany). CA 92093 (USA). Fritzsche, R. A. Dept. of Fisheries, Humboldt State Univer• Maul, G. E. Museu Municipal do Funchal, Madeira (Por• sity, Arcata CA 95521 (USA). tugal). Gibbs, R. H. Jr. Fish Division, National Museum of Natural Nakamura, I. Fisheries Research Station, Kyoto University, History, Washington, D.C. 20560 (USA). Maizuru Kyoto 625 (Japan). xu Nelson, J. S. Dept. of Zoology, University of Alberta, Ed• Smith, M. M. J .L.B. Smith Institute of Ichthyology, Private monton, Alberta T6G 2E9 (Canada). Bag 1015, Grahamstown 6140 (South Africa). Nielsen, J. Zoologisk Museum, Universitetsparken, 15 DK Smith-Vaniz, W. F. Dept. of Ichthyology, Academy of 2100 K!Ilbenhaven (Denmark). Natural Sciences, 19th & Parkway, Philadelphia, PA Olney, J. E. Virginia Institute of Marine Science, Gloucester 19103 (USA). Point, VA 23062 (USA). Springer, V. G. Fish Division, National Museum of Natural Parin, N. V. P. P. Shirshov Institute of Oceanography, History, Washington, D.C. 20560 (USA). Academy of Sciences, Krasikova 23, 117218 Moscow Stein, D. L. College of Oceanography, Oregon State Univer• (USSR). sity, Corvallis, Oregon 97331 (USA). Paxton, J. R. The Australian Museum, P.O. Box A285, Sulak, K. J. Virginia Institute of Marine Science, Gloucester Sydney, NSW 2000 (Australia). Point, VA 23062 (USA). Pietsch, T. W. College of Fisheries, University of Washington, Taylor, W. R. Fish Division, National Museum of Natural Seattle, WA 98195 (USA). History, Washington, D.C. 20560 (USA). Poss, S. G. Gulf Coast Research Laboratory, Ocean Springs, Tyler, J. C. National Museum of Natural History, Washing• MS 39564 (USA). ton, D.C. 20550 (USA). Post, A. Ichthyologie Seefischerei, Zoologisches Institut und Weitzman, S. H. Fish Division, National Museum of Natural Museum, Martin-Luther-King-Platz 3, D-2000 Ham• History, Washington, D.C. 20560 (USA). burg 13 (Federal Republic of Germany). Whitehead, P. J. P. Dept. of Zoology (Fishes), British Randall, J. E. Bernice P. Bishop Museum, P.O. Box 19000A Museum (Natural History), Cromwell Road, London Honolulu, HI 96818 (USA). SW7 5BD (England). Rosenblatt, R. H. Scripps Institution of Oceanography, La Williams, F. School of Marine & Atmospheric Science, Uni• Jolla, CA 92093 (USA). versity of Miami, 4600 Rickenbacker Causeway, Miami, Russell, B. C. Division of Natural Sciences, Northern Territ• FL 33149 (USA). ory Museum, P.O. Box 4646, Darwin, NT 5794 (Aus• Winterbottom, R. Dept. of Ichthyology, Royal Ontario tralia). Museum, 100 Queens Park, Toronto, Ontario M5S 2C6 Schaefer, S. A. Division of Fishes, Field Museum of Natural (Canada). History, Roosevelt Road at Lake Shore Drive, Chicago, Woodland, D. J. University of New England, Armidale, IL 60605 (USA). NSW 2351 (Australia). Shillington, F. A. Dept. of Oceanography, University of Wongratana, T. Dept. of Biology, Faculty of Science, Cape Town (South Africa). Chulalongkorn University, Bangkok (Thailand).

xiii Systematic contents

Taxon Common name Author Page

Superclass Agnatha (Cyclostomata) Class Pteraspidomorphi Myxiniformes Family No.1. Myxinidae hagfishes BoFernholm 35 Superclass Pisces () Class Chondrichthyes Subclass Elasmobranchii Superorder Squalomorphea Order Hexanchiformes Family No.2. Hexanchidae cow sharks A. J. Bass, P. C. Heemstra & L. J. V. Compagno 45 Family No.3. Chlamydoselachidae frill sharks A. J. Bass 47 Order Heterodontiformes Family No.4. Heterodontidae bullhead sharks A. J. Bass 48 Order Squaliformes Family No.5. Squalidae dogfishes A. J. Bass, L. J. V. Compagno & P. C. Heemstra 49 Family No.6. Echinorhinidae bramble sharks A. J. Bass & L. J. V. Compagno 63 Order Orectolobiformes Family No.7. Orectolobidae carpet sharks A. J. Bass 64 Family No.8. Rhincodontidae whale sharks A. J. Bass 66 Order Carcharhiniformes Family No.9. Carcharhinidae requiem sharks A. J. Bass, P. C. Heemstra & L. J. V. Compagno 67 Family No. 10. Proscylliidae proscyliids A. J. Bass& L. J. V. Compagno 87 Family No. 11. Scyliorhinidae catsharks A. J. Bass 88 Family No. 12. Pseudotriakidae false catsharks A. J. Bass 96 Family No. 13. Sphyrnidae hammerhead sharks A. J. Bass 96 Order Lamniformes Family No. 14. Lamnidae mackerel sharks A. J. Bass 98 Family No. 15. Cetorhinidae basking sharks A. J. Bass 101 Family No. 16. Alopiidae thresher sharks A. J. Bass 101 Family No. 17. Mitsukurinidae goblin sharks A. J. Bass & L. J. V. Compagno 103 Family No. 18. Pseudocarchariidae crocodile sharks A. J. Bass 103 Family No. 19. Odontaspididae ragged-tooth sharks A. J. Bass & L. J. V. Compagno 104 Order Pristiophoriformes Family No. 20. Pristiophoridae sawsharks A. J. Bass & P. C. Heemstra 106 Order Squatiniformes Family No. 21. Squatinidae angelsharks A. J. Bass 107 Superorder Batoidea Order Pristiformes Family No. 22. Pristidae sawfishes L. J. V. Compagno 110 Order Torpediniformes Family No. 23. Torpedinidae electric rays L. J. V. Compagno 112 Family No. 24. Narkidae numbfish L. J. V. Compagno 113

xiv Taxon Common name Author Page

Order Rajiformes Suborder Rajoidei Family No. 25. Rajidae skates P. A. Hulley 115 Family No. 26. Anacanthobatidae smooth skates P. A. Hulley 127 Suborder Rhinobatoidei Family No. 27. Rhinobatidae guitarfishes L. 1. V. Compagno 128 Order Myliobatiformes Suborder Myliobatoidei Family No. 28. Myliobatidae eaglerays L. 1. V. Compagno 132 Family No. 29. Mobulidae mantas L. 1. V. Compagno 134 Family No. 30. Dasyatidae stingrays L. 1. V. Compagno 135 Suborder Hexatrygonoidei Family No. 31. Hexatrygonidae sixgill stingrays M. M. Smith & P. C. Heemstra 142 Subclass Holocephali Order Chimaeriformes Family No. 32. Chimaeridae chimaeras L. 1. V. Compagno 144 Family No. 33. Rhinochimaeridae longnose chimaeras L. 1. V. Compagno 146 Family No. 34. Callorhynchidae elephantfish L. 1. V. Compagno 147 Class Osteichthyes Subclass Sarcopterygii Superorder Crossopterygii Order Coelacanthiformes Family No. 35. Latimeriidae coelacanths M. M.Smith 152 Subclass Subdivision Teleostei Infradivision Order Family No. 36. Elopidae ladyfishes M. M.Smith 155 Family No. 37. Megalopidae tarpons M.M. Smith 156 Family No. 38. Albulidae M. M. Smith 157 Order Anguilliformes Family No. 39. Anguillidae freshwater P. H. 1. Castle 160 Family No. 40. Congridae conger eels P. H. 1. Castle 161 Family No. 41. Muraenidae moray eels P. H. 1. Castle & 1. E. McCosker 165 Family No. 42. Ophichthidae snake eels & worm eels 1. E. McCosker & P. H. 1. Castle 176 Family No. 43. Xenocongridae false morays P. H. 1. Castle 186 Family No. 44. Moringuidae spaghetti eels P. H. 1. Castle 187 Family No. 45. Muraenesocidae pike congers P. H. 1. Castle 188 Family No. 46. Synaphobranchidae cutth roat eels P. H. 1. Castle 188 Family No. 47. Nettastomatidae witch eels P. H. 1. Castle 190 Family No. 48. Serrivomeridae sawtooth eels P. H. 1. Castle 191 Family No. 49. Derichthyidae longneck eels P. H. 1. Castle 191 Family No. 50. Cyemidae arrow eels P. H. 1. Castle 192 Family No. 51. Nemichthyidae snipe eels P. H. 1. Castle 193 Order Family No. 52. Notacanthidae spiny eels K. Sulak 195 Family No. 53. Halosauridae halosaurs K. Sulak 196 Infradivision Clupeomorpha Order Family No. 54. Clupeidae herrings, sardines & pilchards P. 1. P. Whitehead & T. Wongratana 199 Family No. 55. Engraulidae anchovies P. 1. P. Whitehead & T. Wongratana 204 Family No. 56. Chirocentridae wolfherrings P. 1. P. Whitehead 207 Infradivision Euteleostei

xv Taxon Common name Author Page

Order Gonorhynchiformes Family No. 57. Gonorhynchidae beaked sandfish M. M. Smith 209 Family No. 58. Chanidae milkfish M. M. Smith 209 Order Siluriformes Family No. 59. Ariidae sea W. R. Taylor 211 Family No. 60. Plotosidae catfishes J. R. Gomon 213 Order Salmoniformes Suborder Argentinoidei Family No. 61. Argentinidae argentines D. M. Cohen 215 Family No. 62. Bathylagidae deepsea smelts D. M.Cohen 216 Family No. 63. Opisthoproctidae barreleyes P. C. Heemstra 216 Family No. 64. Alepocephalidae slickheads D. F. Markle 218 Family No. 65. Platytroctidae tubeshoulders T. Matsui & R. H. Rosenblatt 223 Family No. 66. Leptochilichthyidae leptochilichthyids D. F. Markle 225 Order Family No. 67. scaly dragonfishes R. H. Gibbs, Jr. 229 Family No. 68. Chauliodontidae R. H. Gibbs, Jr. 230 Family No. 69. Astronesthidae snaggletooths R. H. Gibbs, Jr. 231 Family No. 70. Idiacanthidae sawtail fishes P. A. Hulley 234 Family No. 71. Malacosteidae loosejaws R. H. Goodyear & R. H. Gibbs, Jr. 235 Family No. 72. Melanostomiidae scaleless dragonfishes R. H. Gibbs, Jr. 236 Family No. 73. Photichthyidae lightfishes S. Schaefer, R. K. Johnson & J. Badcock 243 Family No. 74. bristle mouths S. Schaefer, R. K. Johnson & J. Badcock 247 Family No. 75. hatchetfishes S. H. Weitzman 253 Order Family No. 76. Chlorophthalmidae greeneyes K. Sulak 261 Family No. 77. Scopelarchidae pear/eyes R. K. Johnson 265 Family No. 78. Notosudidae notosudids G. Krefft 268 Family No. 79. Synodontidae lizardfishes R. F. Cressey 270 Family No. 80. Giganturidae telescopefish R. K. Johnson 273 Family No. 81. Paralepididae barracudinas A. Post 274 Family No. 82. Anotopteridae daggertooths P. A. Hulley 278 Family No. 83. Evermannellidae sabretoothed fishes R. K. Johnson 278 Family No. 84. Omosudidae omosudids G.E.Maul 280 Family No. 85. Alepisauridae lancetfishes P. C. Heemstra & M. M. Smith 280 Order Family No. 86. Myctophidae lanternfishes P. A. Hulley 282 Family No. 87. N eoscopelidae blackchins P. A. Hulley 321 Order Family No. 88. Gadidae cods D. M. Cohen 324 Family No. 89. Merlucciidae hakes D. M. Cohen 324 Family No. 90. Moridae deepsea cods D. M. Cohen 326 Family No. 91. Melanonidae melanonids D. M. Cohen 328 Family No. 92. Bregmacerotidae codlets M. M. Smith 329 Family No. 93. Macrouridae grenadiers T. Iwamoto 330 Order Zoarciformes Family No. 94. Zoarcidae eelpouts M. E. Anderson 342 Family No. 95. Parabrotulidae false brotulas M. E. Anderson 343 Order Family No. 96. Ophidiidae cuskeels J. G. Nielsen & D. M. Cohen 345 Family No. 97. Carapidae pear/fishes J. E. Olney&D. F. Markle 350 Family No. 98. Bythitidae bythitids D. M. Cohen 354 Family No. 99. Aphyonidae aphyonids J. G. Nielsen 356 Order Batrachoidiformes Family No. 100. toadfishes J. B. Hutchins 358

xvi Taxon Common name Author Page

Order Lophiiformes Family No. 101. Lophiidae monks J. H. Caruso 363 Family No. 102. Antennariidae anglers T. W. Pietsch 366 Family No. 103. Chaunacidae coffinfishes M.M.Smith 369 Family No. 104. Ogcocephalidae seabats M. G. Bradbury 370 Family No. 105. Ceratiidae seadevils T. W. Pietsch 373 Family No. 106. Oneirodidae oneirodids T. W. Pietsch 375 Family No. 107. Melanocetidae devil-anglers T. W. Pietsch 375 Family No. 108. Himantolophidae footballfish T. W. Pietsch 376 Family No. 109. Diceratiidae horned anglers T. W. Pietsch 376 Order Gobiesociformes Family No. 110. clingfishes J. C. Briggs 378 Superorder Order Family No. 111. Atherinidae silvers ides W.Ivantsoff 381 Family No. 112. Notocheiridae surf sprites W.Ivantsoff 384 Order Family No. 113. Belonidae needlefishes B. B. Collette 385 Family No. 114. Scomberesocidae sauries M. M.Smith 387 Family No. 115. Hemiramphidae halfbeaks B.B.Collette 388 Family No. 116. Exocoetidae flyingfishes P. C. Heemstra & N. V. Parin 391 Order Family No. 117. Lampridae opahs P. C. Heemstra 398 Family No. 118. Veliferidae velifers P. C. Heemstra 398 Family No. 119. Trachipteridae ribbonfishes P. C. Heemstra & S. X. Kannemeyer 399 Family No. 120. Radiicephalidae tapertail P. C. Heemstra & S. X. Kannemeyer 402 Family No. 121. Lophotidae crestfishes P. C. Heemstra 402 Family No. 122. Regalecidae oarfishes P. C. Heemstra 403 Family No. 123. Stylephoridae tube-eye P. A. Hulley 404 Family No. 124. Ateleopodidae tadpole fishes M.M. Smith 404 Family No. 125. Mirapinnidae mirapinnids E. Bertelsen 406 Order Family No. 126. Berycidae berycids P. C. Heemstra 409 Family No. 127. Trachichthyidae slimeheads P. C. Heemstra 410 Family No. 128. Monocentridae pineapple fishes M.M. Smith 413 Family No. 129. flashlight fish J. E. McCosker 413 Family No. 130. Diretmidae diretmids A. Post 414 Family No. 131. Anoplogasteridae P. A. Hulley 415 Family No. 132. squirrelfishes, soldierfishes J. E. Randall & P. C. Heemstra 415 Family No. 133. Melamphaidae bigscale fishes A. W. Ebeling 427 Family No. 133a. Stephanoberycidae pricklefishes P. C. Heemstra 431 Family No. 134. Polymixiidae P. C. Heemstra 432 Order Cetomimiformes Family No. 135. whalefishes J. R. Paxton & D. J. Bray 433 Family No. 136. Rondeletiidae redmouth whalefishes J. R. Paxton & D. J. Bray 434 Family No. 137. Barbourisiidae red whalefish J. R. Paxton & D. J. Bray 434 Order Family No. 138. Zeidae dories P. C. Heemstra 435 Family No. 139. Oreosomatidae oreos Ch. Karrer 438 Family No. 140. Grammicolepididae tinselfishes Ch. Karrer & P. C. Heemstra 440 Family No. 141. Zeniontidae zeniontids P. C. Heemstra 441 Order Family No. 142. Pegasidae seamoths M.M.Smith 443 Family No. 143. Aulostomidae trumpetfishes P. C. Heemstra 444 Family No. 144. Fistulariidae flutemouths P. C. Heemstra 444 Family No. 145. Syngnathidae seahorses & pipefishes C. E. Dawson 445 Family No. 146. Solenostomidae ghost pipefishes R. A. Fritzsche 459

xvii Taxon Common name Author Page

Family No. 147. Macroramphosidae snipefishes P. C. Heemstra 459 Family No. 148. Centriscidae shrimpfishes P. C. Heemstra 461 Order Family No. 149. Scorpaenidae scorpionfishes W. M. Eschmeyer 463 Family No. 150. Tetrarogidae waspfishes S. Poss 479 Family No. 151. Aploactinidae velvetfishes S. Poss 479 Family No. 152. Congiopodidae horsefishes S. Poss 480 Family No. 153. Caracanthidae crouchers W. N. Eschmeyer 481 Family No. 154. Bembridae deepwater flatheads L. W.Knapp 481 Family No. 155. Platycephalidae flatheads L. W. Knapp 482 Family No. 156. Hoplichthyidae spiny flatheads M.M.Smith 486 Family No. 157. Triglidae gurnards P. C. Heemstra 486 Family No. 158. Peristediidae armoured gurnards P. C. Heemstra 489 Family No. 159. Dactylopteridae helmet gurnards W. N. Eschmeyer 490 Family No. 160. Psychrolutidae fatheads J. S. Nelson 491 Family No. 161. Liparididae snailfishes D. L. Stein 492 Order Family No. 162. boarfishes P. C. Heemstra 506 Family No. 163. Ambassidae glassies P. C. Heemstra & T. J. Martin 507 Family No. 164. Kuhliidae flagtails M. M.Smith 508 Family No. 165. Polyprionidae wreckfishes P. C. Heemstra 509 Family No. 166. Serranidae rockcods & seabasses P. C. Heemstra&J. E. Randall 509 Family No. 167. Grammistidae soapfishes J. E. Randall 537 Family No. 168. Callanthiidae goldies P. C. Heemstra & W. D. Anderson, Jr. 538 Family No. 169. Pseudochromidae dottybacks M.M. Smith 539 Family No. 170. Acanthoclinidae spiny basslets M. M. Smith 541 Family No. 171. Pseudogrammidae podges M. M. Smith 541 Family No. 172. Plesiopidae longfins P. C. Heemstra 542 Family No. 173. Teraponidae thornfishes P. C. Heemstra 543 Family No. 174. Priacanthidae bigeyes P. C. Heemstra 544 Family No. 175. Apogonidae cardinal fishes O.Gon 546 Family No. 176. Acropomatidae acropomatids P. C. Heemstra 561 Family No. 177. Scombropidae gnomefishes P. C. Heemstra 563 Family No. 178. Pomatomidae elf M. M. Smith&J. L. B. Smith 564 Family No. 179. Haemulidae rubberlips & grunters M. M. Smith & R. J. McKay 564 Family No. 180. Dinopercidae cavebass P. C. Heemstra 571 Family No. 181. Lutjanidae snappers W. D. Anderson, Jr. 572 Family No. 182. Caesionidae fusiliers M. M.Smith 579 Family No. 183. Sparidae sea breams J. L. B. Smith & M. M. Smith 580 Family No. 184. Centracanthidae picarels P. C. Heemstra 594 Family No. 185. Lethrinidae emperors M. M. Smith 595 Family No. 186. Nemipteridae butterfly breams & spinecheeks B. C. Russell 600 Family No. 187. Coracinidae galjoens M. M.Smith 601 Family No. 188. Parascorpididae jutjaw fish P. C. Heemstra 602 Family No. 189. Kyphosidae sea chubs M.M. Smith 603 Family No. 190. Scorpididae stonebreams M. M.Smith 604 Family No. 191. Scatophagidae scatties P. C. Heemstra 604 Family No. 192. Ephippidae batfishes M.M. Smith 605 Family No. 193. Monodactylidae moonies P. C. Heemstra 607 Family No. 194. Gerreidae pursemouths D. J. Woodland 608 Family No. 195. Drepanidae sicklefishes M. M.Smith 609 Family No. 196. Mullidae goatfishes A. Ben-Tuvia 610 Family No. 197. Malacanthidae tilefishes P. C. Heemstra 613 Family No. 198. Sillaginidae siUagos R.J. McKay 615 Family No. 199. Sciaenidae kobs P. C. Heemstra 616 Family No. 200. Menidae moonfish M. M.Smith 619 Family No. 201. Leiognathidae soapies M. M. Smith 620 Family No. 202. Lobotidae tripletail P. C. Heemstra 621 Family No. 203. armourheads P. C. Heemstra 622 Family No. 204. Pomacanthidae angelfishes M. M. Smith & P. C. Heemstra 623

XVlll Taxon Common name Author Page

Family No. 205. Chaetodontidae P. C. Hef!mstra 627 Family No. 206. Oplegnathidae knifejaws P. C. Heemstra 632 Family No. 207. Bramidae pomfrets M.M.Smith 633 Family No. 208. Caristiidae manefishes P. C. Heemstra 636 Family No. 209. Emmelichthyidae rovers P. C. Heemstra 637 Family No. 210. Carangidae kingfishes W. F. Smith-Vaniz 638 Family No. 211. Coryphaenidae dolphinfish M.M.Smith 661 Family No. 212. Rachycentridae cobia M.M.Smith 661 Family No. 213. Echeneidae remoras P. C. Heemstra 662 Family No. 214. Cirrhitidae hawkfishes J. E. Randall 664 Family No. 215. Cheilodactylidae fingerfins M. M.Smith 667 Family No. 216. Pempheridae sweepers P. C. Heemstra 668 Family No. 217. Bathyclupeidae bath yclupeids P. C. Heemstra 669 Family No. 218. Cichlidae cichlids P. C. Heemstra 670 Family No. 219. Pomacentridae damselfishes G. R. Allen 670 Family No. 220. Labridae wrasses J. E. Randall 683 Family No. 221. Scaridae parrotfishes J. E. Randall 706 Family No. 222. Mugilidae mullets M. M. Smith &J. L. B. Smith 714 Family No. 223. Polynemidae threadfins M.M.Smith 720 Family No. 224. Sphyraenidae barracudas D. P. de Sylva & F. Williams 721 Family No. 225. Opistognathidae jawfishes W. F. Smith-Vaniz 726 Family No. 226. Cepolidae bandfishes W. F. Smith-Vaniz 727 Family No. 227. Congrogadidae snakelets R. Winterbottom 729 Family No. 228. Chiasmodontidae swallowers R. K. Johnson & M. J. Keene 731 Family No. 229. Champsodontidae gapers P. C. Heemstra 734 Family No. 230. Uranoscopidae stargazers P. C. Heemstra 735 Family No. 231. Trichonotidae sand divers P. C. Heemstra 736 Family No. 232. Creediidae sand burrowers J. S. Nelson 736 Family No. 233. Percophidae duckbills P. C. Heemstra&J. S. Nelson 737 Family No. 234. Mugiloididae sandsmelts P. C. Heemstra 739 Family No. 235. Blenniidae blennies V. G. Springer 742 Family No. 236. Tripterygiidae threefin blennies W. Holleman 755 Family No. 237. Clinidae klipfishes M.M.Smith 758 Family No. 238. Ammodytidae sandlances P. C. Heemstra 769 Family No. 239. Callionymidae dragonets R. Fricke 770 Family No. 240. Gobiidae gobies D. F. Hoese 774 Family No. 241. Eleotridae sleepers D. F. Hoese 807 Family No. 242. Kraemeriidae sand darts D. F. Hoese 811 Family No. 243. Acanthuridae surgeonfishes & unicornfishes J. E. Randall 811 Family No. 244. Zanclidae Moorish idol P. C. Heemstra & M. M. Smith 823 Family No. 245. Siganidae rabbitfishes D. J. Woodland 824 Family No. 246. Scombrolabracidae black mackerel I. Nakamura 825 Family No. 247. Gempylidae snake mackerels I. Nakamura 825 Family No. 248. Trichiuridae frostfishes I. Nakamura 829 Family No. 249. Scombridae tunas, mackerels & bonitos B.B.Collette 831 Family No. 250. Luvaridae louvars P. C. Heemstra 838 Family No. 251. Xiphiidae swordfishes P. C. Heemstra 839 Family No. 252. Istiophoridae sailfish, spearfishes & marlins P. C. Heemstra 839 Suborder Stromateoidei Family No. 253. Amarsipidae amarsipids R. L. Haedrich 842 Family No. 254. Stromateidae ruffs R. L. Haedrich 842 Family No. 255. Nomeidae driftfishes R. L. Haedrich 846 Family No. 256. Tetragonuridae squaretails R. L. Haedrich 851 Order Pleuronectiformes Suborder Psettodoidei Family No. 257. Psettodidae adalahs P. C. Heemstra 853 Suborder Pleuronectoidei Family No. 258. Citharidae largescale flounders P. C. Heemstra 853 Family No. 259. Bothidae lefteye flounders D. A. Hensley 854

XiX Taxon Common name Author Page

Family No. 260. Pleuronectidae righteye flounders P. C. Heemstra 863 Suborder Soleoidei Family No. 261. Cynoglossidae tonguefishes P. C. Heemstra 865 Family No. 262. Soleidae soles P. C. Heemstra & O. Gon 868 Order Suborder Balistoidei Family No. 263. Balistidae M. M. Smith & P. C. Heemstra 876 Family No. 264. Monacanthidae filefishes 1. B. Hutchins 882 Family No. 265. Triacanthodidae spikefishes 1. C. Tyler 887 Family No. 266. Ostraciidae boxfishes M.M.Smith 890 Suborder Tetraodontoidei Family No. 267. Triodontidae threetooth puffer M. M. Smith 894 Family No. 268. Tetraodontidae blaasops M. M. Smith & P. C. Heemstra 894 Family No. 269. Diodontidae burrfishes & porcupinefishes 1. M. Leis 903 Family No. 270. Molidae ocean sunfishes P. C. Heemstra 907

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