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SMITHS' SEA fiSHES
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SMITHS' SEA FISHES
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-----.:..-- -~-.------. fDlHD BY MARGARET M. SM ITH
AND
PHilliP C. HEEMSTRA
Springer-verlag Berlin Heidelberg New York
London Paris Tokyo
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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 Africa 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 Insti tute 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 wi thout 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 of 76 scientists, representing 15 countries. Several skilled artists and photographers have also contributed to the numerous and beautiful illustrations. Research done in South Africa was supported by the Council for Scientific and Industrial Research. Work on the book at the J.L.B. Smith Institute was also funded by the Department of National Education and The Trustees of the Sea Fishes of Southern Africa Book Fund. Expenses of visits by ichthyologists from overseas were defrayed by a grant from the John S. Schlesinger Foundation.
Although this book is an impressive contribution to the zoology of southern Africa, few people will realise the enormous amount of work that has gone into it. Prof. Margaret 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 book; and third, to participate in the actual research, writing, editing, production of most of the illustrations, and ar-
ranging all of the colour plates for the book. For the past eight years, she has been assisted in the research, writing and editing of the book by Dr. P. C. Heemstra.
It is essential for the proper management of the marine fish resources of southern Africa that we know what species we are dealing with. This book will greatly facilitate the identification of our fishes. It will thus be of vital importance to the understanding and wise use of our tremendously diverse and valuable fish fauna.
We are proud of the affiliation of the J.L.B. Smith Institute with Rhodes University and happy to see the successful completion of this long-awaited work.
Dr. D. S. Henderson, Chairman of the Institute and Vice-Chancellor of Rhodes University
1. P. de Wit, Deputy President, CSIR
When J. L. B. Smith's monumental book The Sea Fishes of Southern Africa was first published in 1949, the first printing was sold out within three weeks. Due in large measure to the excellent and numerous illustrations (mostly by Margaret Smith), the book has been extremely popular in South Africa and also much in demand overseas. It has become an international ichthyological "classic" and is well known in museums and marine laboratories all over the world. In 1953 and 1961, J. L. B. Smith added a few additional species in appendices at the end of the book. Since 1961, many new species and new records of fishes have been discovered in the seas off southern Africa. Also, recent ichthyological research has considerably altered the classification 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
considered advisable to enlist the assistance of other ichthyologists who are experts on the various families of marine fishes represented in the seas of southern Africa. This team effort has produced a uniquely authoritative and comprehensive book on the marine fishes of southern Africa. To all of the many collaborators, we owe a debt of gratitude for their unstinting efforts to provide the latest and best fruits of their ichthyological research.
With this book, on which she laboured for over 15 years, Margaret Smith has produced not only a worthy tribute to her late husband's great achievements, but also a lasting testimonial to her own dedication to the fulfilment of his ideals.
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 Osteichthyes (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 A. R. Thorpe
W. M. Smith Dr I. Mackenzie Dr B. E. Hersov
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
Barclays National Bank Ltd Fisheries Development Corporation of
South Africa Ltd The Gencor Development Fund Hewlett Packard South Africa (Pty) Ltd
Anglovaal Ltd Atlantic Fishing Enterprises Barlow Rand Limited CTP Book Printers (Pty) Ltd Durban City Council Hans Merensky Holdings (Pty) Ltd Irvin & Johnson Ltd Metal Box South Africa Ltd Monaco Institute of Oceanographic Research E. J. and A. L. Morris
Adcan Marine Suppliers Africana Book Collectors Dr B. R. Allanson Trevor Ashington L. G. Baumann The Brenthurst Library Professor J. W. Brommert Craig, Ryan and Tracey Bruton David Christie Pearl E. Coppin Bert Davids Claire Davids H. B. Dyer Johan Els Professor P. K. Faure
Corporate Donors Lennon Ltd Natal Parks Board Professor Margaret M. Smith William and Jenny Smith Total South Africa (Pty) Ltd
Corporate Subscribers National Parks Board Oceanographic Research Institute, Durban SAPPI Fine Papers (Pty) Ltd Smithsonian Institution The South African Association for
Marine Biological Research South African Permanent Building Society Standard Bank Investment Corporation A. R. Thorpe Wildlife Society of Southern Africa
Individual Donors Hedley Gerhardt Pat Goss Henry J. Gradwell Dr D. S. Henderson Basil Hersov Dr Brian Hulley Isowall SA (Pty) Ltd Alf Johnson Jessica and Len Jones Errol and Fredia Kreutzer Helen Leibel Robert E. Levitt Ian Mackenzie Natal Ski-Boat Light Tackle Game Fishing
Association
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Individual Donors
Richards Bay Minerals Mrs L. M. Russell S.A. Shipping News & Fishing Industry Review Derek and Linda Souter South African Ski-Boat Angling Association Nolan Sparg
George Brian Trow G. H. Tudor Umhlali Ski-Boat Club Pieter W. van Niekerk Zoological Society of Southern Africa
Individual Subscribers Christopher C. Appleton Peter J. Ashton Eugene K. Balon E. J. B. Bishop· Border Ski-Boat Angling Association Alan and Jiff Bowmaker The Braine family George and Margo Branch Alastair and Patricia Buchan Ian D. J. Cochrane-Murray Dr Allan Connell Douglas T. Cooke Mrs H. J. Creswell Brendan Davids John-Paul Davids Thomas Davids Durban Ski-Boat Club The Mayor of East London East London Museum East London Ski-Boat Club D. H. Eccles Karl Zachary Edwards Lloyd Edwards John Dudley Engelbrecht Director, Fishing Industry Research Institute Professor Vernon S. Forbes and W. H. Wood Mayor, Councillors and Citizens of Fort Beaufort Eugene and Lalie Fourie L. P. Dormehl Gertenbach M. C. V. Ginn Gonubie Marine Club Ken Gough Dr C. E. and Mrs D. Gow Grahamstown Municipality J. T. Hart Glen Harvey Alan S. Henderson Ian M. Henderson Ethnee Joy Hepburn (nee Amm) Viktor E. Hesse Dr P. Alexander Hulley Humansdorp Municipality Indian Ocean Export Company (Pty) Ltd P. B. N. Jackson J. W. Jagger Library, University of Cape Town
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City of Johannesburg Marthinus J. S. Jordaan Professor D. M. Joubert Hillel M. Kahn Sidney H. Kahn Dr P. G. Landsberg Dr B. E. la Trobe, Mayor Grahamstown 1985/86 R. H. Lefebvre In memory of the late Dr C. Mauritz Lombaard R. Loots W. McLevland E. K. Moorcroft Natal Ski-Boat Association National Zoological Gardens of South Africa Dan Niven Patrick Niven Oceana Fishing Group Ltd Mike and Mary-Lou Penrith Robin G. Piper Port Alfred Municipality Gavin, Howard, Charles, Mark and Jamie Pote R. McC Pott Ray Preen Requin Angling Club S.A. Dr G. J. B. Ross SAICCOR (Pty) Ltd R. Allan Scott Sea Fisheries Research Institute C. P. Skinner Rodney P. Smith W. H. Stoops Sandy Stretton Oscar B. Susman Swaziland Underwater Group Professor John Tew H. P. Thesen Umzimkulu Ski-Boat Club Rudy van der Elst Dr John H. Wallace E. G. Wanliss Mr and Mrs T. N. Webb City of Welkom Mr and Mrs K. C. Whyte Mr and Mrs J. O. M. Wilkinson Alastair and Gerry Williams
Acknowledgements
It is impossible to mention all of the many people who have assisted in the completion of this book. We are indebted to our numerous collaborators for their contributions and their patience in the long period that it has taken to complete this work. We are grateful to our assistant, Mrs Joan Wright, whose careful work, industry and skill with the word processor have been of tremendous help to us. We thanks Mrs Jean Pote for her unique ability to decipher the illegible handwriting and editorial ciphers presented to her; Mrs Pote has also done an enormous job on the word processor.
Special thanks and credit are due to the artists who have contributed the paintings and drawings that illustrate this work. We thank Elizabeth Tarr for her enormous contribution; Ms Tarr's accuracy and attention to detail are exemplary. We are much indebted to David Voorvelt, whose remarkable industry and skill as an artist have been essential to the completion of this work. We are also grateful to the other fine artists who have made drawings for us: Jean Michel Vincent, Virginia McRostie, Anne Lefebvre, Moira Lambert, and from the original Sea Fishes of Southern Africa: Patricia Parkin, Denys M. Davis and Hester W. Locke. We are especially grateful to John E. Randall for numerous transparencies used in the colour plates. Gerald R. Allen, Alan Emery, John Gunn, Sidney Kannemeyer, Dennis King, Daniel Pelicier, A. R. Thorpe, Peter van Niekerk and Richard Winterbottom also supplied some transparencies used in this book.
We are grateful to Robin Stobbs, whose abilities as a photographer, small boat captain, laboratory technician, computor operator and field assistant have been of immeasurable help to us. We thank Billy Ranchod for his careful and excellent work as our collection manager. Mrs Doris Cave, who was the Institute's librarian for twenty-five years, assisted in the compilation of the bibliography and generally facilitated our work by maintaining a well-organised and superbly indexed library.
Throughout the years the public, including the angling fraternity, naturalists, divers and school children, have sent in numerous specimens and contributed much useful information. Several persons have made a special effort
to collect fishes for us. In some cases these specimens have represented new species or new records for southern Africa. We are especially grateful to A. R. Thorpe, Dave Bickell, Rudy van der Elst, Malcolm Smale, Alan Connell, Colin Buxton, Dennis King, Len Jones, Alf Johnson, Elizabeth Tarr, Rob Tarr, Brian Trow, Hennie Kok, Wouter Holleman, Anton Bok, Tom Hecht, Robin Stobbs, Mike Bruton and Lynneth Beckley. We also thank the East London and Port Elizabeth Museums and the Oceanographic Research Institute in Durban for their efforts in sending specimens. We are indebted to P. A. Hulley and S. X. Kannemeyerfor their prompt loans of material from the South African Museum and for providing data on various fishes in their collection.
Most of the funds for the work done in Grahamstown were provided by the South African Council for Scientific and Industrial Research (CSIR) through the South African National Committee for Oceanographic Research. Additional funds were provided by the Smiths' Sea Fishes Trust and Rhodes University. We are also grateful for the excellent facilities and resources of the J.L.B. Smith Institute of Ichthyology and for the support of the Department of National Education.
We thank the members of our Editorial Committee, Brian R. Allanson, P. A. Hulley, John Wallace and A. R. Thorpe, for their valuable advice. We are also grateful for the excellent editorial assistance of Frances Perryer, Matthew Seal and Basil van Rooyen of Macmillan, S.A.
For donations of various kinds, we thank the Fisheries Development Corporation, Sasolchem (Pty) Ltd and the Hewlett-Packard Co.
We are indebted to the Heemstra family (Valerie, Lydia and Julia) for much encouragement, help of various kinds, and their forbearance in tolerating the many weekends and evenings their father and husband devoted to this book.
Finally, we wish to express our gratitude to the director of our Institute, Prof. M. N. Bruton, for his constant support, advice and assistance in the completion of the manuscript.
xi
Contributors
Allen, G. R. Western Australian Museum, Francis Street, Perth, WA 6000 (Australia).
Anderson, M. E. Dept. of Ichthyology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 (USA).
Anderson, W. D. Jr. Grice Marine Biological Lab, 205 Fort Johnson, Charleston, SC 29407 (USA).
Badcock, J. Institute of Oceanographic Sciences, Brook Road, Godalming, Surrey GU8 5UB (England).
Baird, R. C. Hayward-Schuster Woolen Mills Inc., East Douglas, MA 01516 (USA).
Bass, A. J. Western Australian Museum, Francis Street, Perth, WA 6000 (Australia).
Ben-Tuvia, A. Dept. of Zoology, the Hebrew University of Jerusalem (Israel).
Bertelsen, E. Zoologisk Museum, Universitetsparken 15, DK 2100 Kl'lbenhavn (Denmark).
Bradbury, M. G. Dept. of Marine Biology, San Francisco State College, San Francisco, CA 94132 (USA).
Bray, D. J. The Australian Museum, P.O. Box A285, Sydney South, NSW 2000 (Australia).
Briggs, J. C. Dept. of Marine Sciences, University of South Florida, St. Petersburg, FL 33701 (USA).
Caruso, J. H. Museum of Natural History, Tulane University, Belle Chasse, LA 70037.
Castle, P. H. J. Dept. of Zoology, Victoria University, Private Bag, Wellington (New Zealand).
Cohen, D. M. Division of Life Sciences, L.A. County Museum of Natural History, 900 Exposition Blvd, Los Angeles, CA 90007 (USA).
Collette, B. B. Systematics Laboratory, National Marine Fisheries Service, National Museum of Natural History, Washington, D.C. 20560 (USA).
Compagno, L. J. V. J. L. B. Smith Institute of Ichthyology, Private Bag 1015, Grahamstown 6140 (South Africa).
Cressey, R. F. Crustacea Division, National Museum of Natural History, Washington, D.C. 20560 (USA).
Dawson, C. E. Gulf Coast Research Laboratory, P.O. Box 218, Ocean Springs, MS 39564 (USA).
de Sylva, D. E. School of Marine & Atmospheric Sciences, University of Miami, 4600 Rickenbacker Causeway, Miami, FL 33149 (USA).
Ebeling, A. W. Dept. of Biological Sciences, University of California, Santa Barbara, CA 93106 (USA).
Eschmeyer, W. N. Dept. ofIchthyology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 (USA).
Fernholm, B. Vertebrate Zoology Research Dept., Swedish Museum of Natural History, S-104 05 Stockholm (Sweden).
Fricke, R. Saalestr. 3A, D-3300 Braunschweig (Federal Republic of Germany).
Fritzsche, R. A. Dept. of Fisheries, Humboldt State University, Arcata CA 95521 (USA).
Gibbs, R. H. Jr. Fish Division, National Museum of Natural History, Washington, D.C. 20560 (USA).
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Gomon, J. R. Fish Division, National Museum of Natural History, Washington, D.C. 20560 (USA).
Gon, O. J.L.B. Smith Institute of Ichthyology, Private Bag 1015, Grahamstown 6140 (South Africa).
Goodyear, R. H. Centro de Ciencias del Mar y Limnologia, Universidad de Panama (Republic of Panama).
Haedrich, R. L. Memorial University of Newfoundland, St. Johns, Newfoundland AlB 3X9 (Canada).
Hensley, D. A. Department de Ciencias Marinas, Universidad de Puerto Rico, Mayagiiez, Puerto Rico 00708 (USA).
Heemstra, P. C. J.L.B. Smith Institute of Ichthyology, Private Bag 1015, Grahamstown 6140 (South Africa).
Hoese, D. F. The Australian Museum, P.O. Box A285, Sydney, NSW 2000 (Australia).
Holleman, W. Albany Museum, Grahamstown 6140 (South Africa).
Hulley, P. A. Dept. of Marine Biology, South African Museum, P.O. Box 61, Cape Town 8000 (South Africa).
Hutchins, J. B. Western Australian Museum, Francis Street, Perth, WA 6000 (Australia).
IvantsotT, W. School of Biological Sciences, Macquarie University, North Ryde, NSW2113 (Australia).
Iwamoto, T. Dept. of Ichthyology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 (USA).
Johnson, R. K. Division of Fishes, Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, IL 60605 (USA).
Kannemeyer, S. X. Dept. of Marine Biology, South African Museum, P.O. Box 61, Cape Town 8000 (South Africa).
Karrer, C. Ichthyologie Seefischerei, Zoologisches Institut und Museum, Martin-Luther-King-Platz 3, D-2000 Hamburg 13 (Federal Republic of Germany).
Keene, J. Route 1, Box 262, Shipman, VA 22971 (USA). Knapp, L. W. Smithsonian Oceanographic Sorting Center,
Washington, D.C. 20560 (USA). KretTt, G. Ichthyologie Seefischerei, Zoologisches Institut
und Museum, Martin-Luther-King-Platz 3, D-2000 Hamburg 13 (Federal Republic of Germany).
Leis, J. M. Fish Dept., The Australian Museum, P.O. Box A285, Sydney, NSW 2000 (Australia).
McCosker, J. E. Steinhart Aquarium, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 (USA).
McKay, R. J. Queensland Museum, Gregory Terrace, Fortitude Valley, QLD 4006 (Australia).
Markle, D. F. Dept. of Fisheries & Wildlife, Oregon State University, Corvallis, OR 97331 (USA).
Martin, T. J. Dept. of Zoology, University of Natal, Durban (South Africa).
Matsui, T. Scripps Institution of Oceanography, La Jolla, CA 92093 (USA).
Maul, G. E. Museu Municipal do Funchal, Madeira (Portugal).
Nakamura, I. Fisheries Research Station, Kyoto University, Maizuru Kyoto 625 (Japan).
Nelson, J. S. Dept. of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9 (Canada).
Nielsen, J. Zoologisk Museum, Universitetsparken, 15 DK 2100 K!Ilbenhaven (Denmark).
Olney, J. E. Virginia Institute of Marine Science, Gloucester Point, VA 23062 (USA).
Parin, N. V. P. P. Shirshov Institute of Oceanography, Academy of Sciences, Krasikova 23, 117218 Moscow (USSR).
Paxton, J. R. The Australian Museum, P.O. Box A285, Sydney, NSW 2000 (Australia).
Pietsch, T. W. College of Fisheries, University of Washington, Seattle, WA 98195 (USA).
Poss, S. G. Gulf Coast Research Laboratory, Ocean Springs, MS 39564 (USA).
Post, A. Ichthyologie Seefischerei, Zoologisches Institut und Museum, Martin-Luther-King-Platz 3, D-2000 Hamburg 13 (Federal Republic of Germany).
Randall, J. E. Bernice P. Bishop Museum, P.O. Box 19000A Honolulu, HI 96818 (USA).
Rosenblatt, R. H. Scripps Institution of Oceanography, La Jolla, CA 92093 (USA).
Russell, B. C. Division of Natural Sciences, Northern Territory Museum, P.O. Box 4646, Darwin, NT 5794 (Australia).
Schaefer, S. A. Division of Fishes, Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, IL 60605 (USA).
Shillington, F. A. Dept. of Oceanography, University of Cape Town (South Africa).
Smith, M. M. J .L.B. Smith Institute of Ichthyology, Private Bag 1015, Grahamstown 6140 (South Africa).
Smith-Vaniz, W. F. Dept. of Ichthyology, Academy of Natural Sciences, 19th & Parkway, Philadelphia, PA 19103 (USA).
Springer, V. G. Fish Division, National Museum of Natural History, Washington, D.C. 20560 (USA).
Stein, D. L. College of Oceanography, Oregon State University, Corvallis, Oregon 97331 (USA).
Sulak, K. J. Virginia Institute of Marine Science, Gloucester Point, VA 23062 (USA).
Taylor, W. R. Fish Division, National Museum of Natural History, Washington, D.C. 20560 (USA).
Tyler, J. C. National Museum of Natural History, Washington, D.C. 20550 (USA).
Weitzman, S. H. Fish Division, National Museum of Natural History, Washington, D.C. 20560 (USA).
Whitehead, P. J. P. Dept. of Zoology (Fishes), British Museum (Natural History), Cromwell Road, London SW7 5BD (England).
Williams, F. School of Marine & Atmospheric Science, University of Miami, 4600 Rickenbacker Causeway, Miami, FL 33149 (USA).
Winterbottom, R. Dept. of Ichthyology, Royal Ontario Museum, 100 Queens Park, Toronto, Ontario M5S 2C6 (Canada).
Woodland, D. J. University of New England, Armidale, NSW 2351 (Australia).
Wongratana, T. Dept. of Biology, Faculty of Science, Chulalongkorn University, Bangkok (Thailand).
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Systematic contents
Taxon Common name Author Page
Superclass Agnatha (Cyclostomata) Class Pteraspidomorphi
Order Myxiniformes Family No.1. Myxinidae hagfishes BoFernholm 35 Superclass Pisces (Gnathostomata) 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 Actinopterygii Subdivision Teleostei Infradivision Elopomorpha
Order Elopiformes Family No. 36. Elopidae ladyfishes M. M.Smith 155 Family No. 37. Megalopidae tarpons M.M. Smith 156 Family No. 38. Albulidae bonefishes M. M. Smith 157
Order Anguilliformes Family No. 39. Anguillidae freshwater eels 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 Notacanthiformes Family No. 52. Notacanthidae spiny eels K. Sulak 195 Family No. 53. Halosauridae halosaurs K. Sulak 196 Infradivision Clupeomorpha
Order Clupeiformes 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 catfishes W. R. Taylor 211 Family No. 60. Plotosidae eel 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 Stomiiformes Family No. 67. Stomiidae scaly dragonfishes R. H. Gibbs, Jr. 229 Family No. 68. Chauliodontidae viper fishes 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. Gonostomatidae bristle mouths S. Schaefer, R. K. Johnson & J. Badcock 247 Family No. 75. Sternoptychidae hatchetfishes S. H. Weitzman 253
Order Aulopiformes 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 lancet fishes P. C. Heemstra & M. M. Smith 280
Order Myctophiformes Family No. 86. Myctophidae lanternfishes P. A. Hulley 282 Family No. 87. N eoscopelidae blackchins P. A. Hulley 321
Order Gadiformes 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 Ophidiiformes 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. Batrachoididae toadfishes J. B. Hutchins 358
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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. Gobiesocidae clingfishes J. C. Briggs 378 Superorder Acanthopterygii
Order Atheriniformes Family No. 111. Atherinidae silvers ides W.Ivantsoff 381 Family No. 112. Notocheiridae surf sprites W.Ivantsoff 384
Order Beloniformes 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 Lampriformes 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 crest fishes 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 Beryciformes 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. Anomalopidae flashlight fish J. E. McCosker 413 Family No. 130. Diretmidae diretmids A. Post 414 Family No. 131. Anoplogasteridae fangtooth P. A. Hulley 415 Family No. 132. Holocentridae squirrel fishes, 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 beardfishes P. C. Heemstra 432
Order Cetomimiformes Family No. 135. Cetomimidae 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 Zeiformes 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 Syngnathiformes 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
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Taxon Common name Author Page
Family No. 147. Macroramphosidae snipefishes P. C. Heemstra 459 Family No. 148. Centriscidae shrimpfishes P. C. Heemstra 461
Order Scorpaeniformes 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 coral 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 Perciformes Family No. 162. Caproidae 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 wreck fishes 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 long fins 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 jut jaw 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. Pentacerotidae armourheads P. C. Heemstra 622 Family No. 204. Pomacanthidae angel fishes M. M. Smith & P. C. Heemstra 623
XVlll
Taxon Common name Author Page
Family No. 205. Chaetodontidae butterflyfishes 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 finger fins 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 damsel fishes G. R. Allen 670 Family No. 220. Labridae wrasses J. E. Randall 683 Family No. 221. Scaridae parrot fishes J. E. Randall 706 Family No. 222. Mugilidae mullets M. M. Smith &J. L. B. Smith 714 Family No. 223. Polynemidae thread fins 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 rabbit fishes 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 frost fishes 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 &
Suborder Stromateoidei marlins P. C. Heemstra 839
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 Tetraodontiformes Suborder Balistoidei Family No. 263. Balistidae triggerfishes 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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