Deep–sea Cartilaginous Fishes of the Indian Ocean, Vol. 1 – Sharks

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FAO Species Catalogue for Fishery Purposes No. 8, Vol. 1

ISSN 1020-8682

DEEPSEA CARTILAGINOUS FISHES

OF THE INDIAN OCEAN

Volume 1. Sharks


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DEEPSEACARTILAGINOUS FISHESOF THE INDIAN OCEAN

Volume 1

Sharks

by

David A. Ebert

Pacic Shark Research CenterMoss Landing Marine Laboratories

Moss Landing, California

United States of America

Support to the implementation of the International Guidelines on the

Management of Deep-Sea Fisheries in the High Seas (GCP/GLO/323/NOR)and

Fisheries management and marine conservation within a changing ecosystem

context (GCP/INT/253/JPN)

FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS

Rome, 2013


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The designations employed and the presentation of material in thisinformation product do not imply the expression of any opinion whatsoeveron the part of the Food and Agriculture Organization of the United Nations(FAO) concerning the legal or development status of any country, territory, cityor area or of its authorities, or concerning the delimitation of its frontiers orboundaries. The mention of specic companies or products of manufacturers,whether or not these have been patented, does not imply that these havebeen endorsed or recommended by FAO in preference to others of a similarnature that are not mentioned.

The views expressed in this information product are those of the author(s) anddo not necessarily reect the views or policies of FAO.

ISBN 978-92-5-107967-6 (print)E-ISBN 978-92-5-107968-3 (PDF)

FAO, 2013

FAO encourages the use, reproduction and dissemination of material in thisinformation product. Except where otherwise indicated, material may becopied, downloaded and printed for private study, research and teachingpurposes, or for use in non-commercial products or services, provided thatappropriate acknowledgement of FAO as the source and copyright holder isgiven and that FAOs endorsement of users views, products or services is notimplied in any way.

All requests for translation and adaptation rights, and for resale and othercommercial use rights should be made via www.fao.org/contact-us/licence-request or addressed to [email protected].

FAO information products are available on the FAO website (www.fao.org/publications) and can be purchased through [email protected].


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PREPARATION OF THIS DOCUMENT

This document was prepared under the FAO Deepsea Fisheries Programme, thanks to a generous funding from theGovernments of Norway and Japan (Support to the implementation of the International Guidelines on the Management ofDeep-Sea Fisheries in the High Seasand Fisheries management and marine conservation within a changing ecosystemcontext projects) for the purpose of assisting states, institutions, the shing industry and RFMO/As in the implementation ofFAO International Guidelines for the Management of Deep-sea Fisheries in the High Seas. This catalogue was developedin close collaboration with the FishFinder Programme of the Marine and Inland Fisheries Branch, Fisheries Department,Food and Agriculture Organization of the United Nations (FAO).

Its production follows the recommendations made during a workshop on Deep-sea Species Identication held in Rome in2009 organized in response to the need for a strategy for the development of appropriate deep-sea species identicationtools for shery purposes, in particular, to address the broadened requirements for reporting on not only target species,but also associated species following recent international developments with respect to sheries management guidanceand biodiversity conservation. The workshop recommended that a series of identication guides be developed forcertain vulnerable groups of species affected by bottom gear, with an initial focus on three of the most impacted groups:cartilaginous shes, corals and sponges. As a starting point, in consideration of the extensive information available oncartilaginous shes from other FAO guides and publications, it was decided to develop deep-sea identication guides forthis group at a regional level.

The present publication is the rst of a twovolume set dedicated to the identication of deepsea cartilaginous shesknown to occur in the Indian Ocean, encompassing FAO Fishing Areas 51 and 57, and that portion of Area 47 off SouthAfrica from about 1842E to 3000E. It focuses on sharklike species, providing accounts for all orders, families, andgenera and for one representative species of each genus. Moreover, fully illustrated keys to all taxa are included.

It is aimed at facilitating the species specic identication of deepsea shark shes occurring in the Indian Ocean by sheryobservers, crew members, scientists, shery ofcers and the interested public.

Supervision:Merete Tandstad, Jessica Sanders and Johanne Fischer (FAO, Rome)

Technical editor:Edoardo Mostarda (FAO, Rome)

Scientic illustrator (for material presented here for the rst time):Emanuela DAntoni (FAO, Rome)Distribution maps:Fabio Carocci (FAO, Rome)

Cover illustration: Emanuela DAntoni (FAO, Rome)

D.A. Ebert. 2013

Deepsea Cartilaginous Fishes of the Indian Ocean. Volume 1. Sharks.

FAO Species Catalogue for Fishery Purposes. No. 8, Vol. 1. Rome, FAO. 256 pp.

ABSTRACT

This volume is a comprehensive, fully illustrated Catalogue of the Deepsea Sharks of the Indian Ocean,encompassing FAO Fishing Areas 51 and 57, and that portion of Area 47 off South Africa from about 1842Eto 3000E. The present volume includes 8 orders, 23 families, 46 genera, and 117 species of sharklike shesoccurring in the Indian Ocean deepsea. It provides accounts for all orders, families, and genera and all keysto taxa are fully illustrated. A species representative account of each genus is also provided and includes:valid modern names and original citation of the species; synonyms; the English, French, and Spanish FAOnames for the species; a lateral view and often other useful illustrations; eld marks; diagnostic features;distribution, including a GIS map; habitat; biology; size; interest to sheries and human impact; local nameswhen available; a remarks sections; and literature. The volume is fully indexed and also includes sectionson terminology and measurements for sharks including an extensive glossary, and a dedicated bibliography.

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ACKNOWLEDGEMENTS

A volume of this magnitude and scope could not be accomplished without the generous help of others and the authorwishes to thank all of those who have been extremely helpful and generous with their time in responding to numerousquestions, providing data and information from their own research (some of it unpublished), and providing much neededliterature. The generic and species accounts were improved immensely from the contribution of colleagues and friends.

Apologies beforehand if somebody was forgotten, which will inevitably happen with a work of this magnitude, but all arethanked for their help and assistance.

Sincere thanks go to the FAO FishFinder team for providing assistance during the preparation of the manuscript and toMerete Tandstad and Jessica Sanders (FAO, Rome) who served as coordinators for the Deepsea Fisheries Programmeactivity related to the development of species identication guides for potentially vulnerable deep-sea species.

Thanks are extended to the following individuals for general discussions and information on various aspects of this project:Peter Kyne (Charles Darwin University), Colin Simpfendorfer (James Cook University), Rob Leslie (Department AgricultureForestry and Fisheries, Cape Town), Leonard Compagno, Wayne Florence, Dylan Clarke, and Michael Bougaardt (SouthAfrican Museum), Matthias Stehmann (ICHTHYS, Ichthyological Research Laboratory); Simon Wiegmann (UniversityHamburg), Hajime Ishihara (W and I Associates), Dean Grubbs (Florida State University), Michelle Hulls (Hobart,Tasmania), Cristina Oddone (Universidade Federal de Rio GrandeFRUG), Malcolm Francis (NIWA), Marcelo de Carvalho

(University of So Paulo), Hans Ho (National Museum of Marine Biology and Aquarium, Taiwan), Ivy Baremore and JohnCarlson (NOAA Southeast Fisheries Science Center, Panama City, Florida); David Catania, William Eschmeyer, and JonFong (California Academy of Sciences, San Francisco); Aaron Henderson (Sultan Qaboos University, Sultanate of Oman);Gavin Naylor and Nico Straube (College of Charleston, Charleston, South Carolina, USA); Bernard Sret (IRDMNHNParis, France); William White, Peter Last and Ross Daley (CSIRO Marine Laboratories, Hobart, Tasmania, Australia);Paul Cowley (South African Institute of Aquatic Biodiversity); Paul Clerkin (Pacic Shark Research Center, Moss LandingMarine Laboratories); Ross Shotton (Southern Ocean Deepsea Fishers Association); Graham Patchell, Daryl Smith,Phil Gaugler and crew of the F/V Will Watch (Sealord Group Ltd); Chris Heinecken and David Japp (CapFish & FOSScc); Daroomalingum Mauree (Director of Fisheries, Ministry of Fisheries, Mauritius); Rajiv Bacorisen, Asaid Morrarby,Nankishore Toofany, and the staff at the Albion Fisheries Research Center.

A special thanks to Joan Parker and the library staff (Moss Landing Marine Laboratories, Moss Landing, California, USA) fortracking down those hard to nd, usually obscure, references, and to Kelley Andrews, Jenny Bigman, Paul Clerkin, James

Knuckey, Mark BreenKlein and Melissa Nehmens (Pacic Shark Research Center, Moss Landing Marine Laboratories,Moss Landing, California, USA) who provided general assistance on various aspects of this project.

Finally, thanks to Kazuhiro Nakaya (Hoikkaido University), Sho Tanaka (Department of Marine Biology, School of MarineScience and Technology, Tokai University)andagain to Peter Kyne (Charles Darwin University) for critically reviewing thenal draft, and for their precious suggestions and useful information.

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1. INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

1.1 Plan of the Catalogue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

1.2 Technical Terms and Measurements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

1.2.1 Picture Guide to External Terminology of Sharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61.2.2 Picture Guide to Skeletal Terminology of Sharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81.2.3 Measurements Used for Sharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101.2.4 Glossary of Technical Terms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

2. SYSTEMATIC CATALOGUE Subclass NEOSELACHII Cohort SELACHII. . . . . . . . . . . . . . . . . 31

2.1 Order HEXANCHIFORMES Frilled and cow sharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

2.1.1 Family CHLAMYDOSELACHIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

ChlamydoselachusGarman, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34Chlamydoselachus anguineus Garman, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

2.1.2 Family HEXANCHIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

HeptranchiasRanesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38Heptranchias perlo(Bonnaterre, 1788). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39HexanchusRanesque, 1810. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41Hexanchus griseus(Bonnaterre, 1788) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

2.2 Order SQUALIFORMES Dogsh sharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

2.2.1 Family ECHINORHINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

EchinorhinusBlainville, 1816 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49Echinorhinus brucus(Bonnaterre, 1788). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

2.2.2 Family SQUALIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

CirrhigaleusTanaka, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53Cirrhigaleusasper(Merrett, 1973) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54SqualusLinnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55Squalus chloroculusLast, White, and Motomura, 2007 . . . . . . . . . . . . . . . . . . . . . . . . . . 58Squalus edmundsiWhite, Last, and Stevens, 2007. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59Squalus hemipinnis White, Last, and Yearsley, 2007 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61Squalus montalbani Whitley, 1931 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63

2.2.3 Family CENTROPHORIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

CentrophorusMller and Henle, 1837 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67Centrophorus granulosus(Bloch and Schneider, 1801) . . . . . . . . . . . . . . . . . . . . . . . . . . 70Centrophorus moluccensisBleeker, 1860 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

DeaniaJordan and Snyder, 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74Deania calcea(Lowe, 1839) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75

2.2.4 Family ETMOPTERIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77

CentroscylliumMller and Henle, 1841 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

Centroscyllium ornatum(Alcock, 1889) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80EtmopterusRanesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81Etmopterus granulosus(Gnther, 1880) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

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2.2.5 Family SOMNIOSIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

CentroscymnusBocage and Capello, 1864. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89Centroscymnus coelolepisBocage and Capello, 1864. . . . . . . . . . . . . . . . . . . . . . . . . . . 90CentroselachusGarman, 1913. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91Centroselachus crepidater(Bocage and Capello, 1864) . . . . . . . . . . . . . . . . . . . . . . . . . 92ProscymnodonFowler, 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93Proscymnodon plunketi(Waite, 1910) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94ScymnodalatiasGarrick, 1956. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95Scymnodalatias albicaudaTaniuchi and Garrick, 1986. . . . . . . . . . . . . . . . . . . . . . . . . . 96Somniosus Lesueur, 1818. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97Somniosus antarcticusWhitley, 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98

ZameusJordan and Fowler, 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100Zameus squamulosus(Gnther, 1877) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101

2.2.6 Family OXYNOTIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102

OxynotusRanesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104Oxynotus bruniensis(Ogilby, 1893) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104

2.2.7 Family DALATIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105

DalatiasRanesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108Dalatias licha(Bonnaterre, 1788) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109EuprotomicrusGill, 1865 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110Euprotomicrusbispinatus(Quoy and Gaimard, 1824) . . . . . . . . . . . . . . . . . . . . . . . . . 111HeteroscymnoidesFowler, 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112HeteroscymnoidesmarleyiFowler, 1934. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113IsistiusGill, 1865. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114Isistiusbrasiliensis(Cuvier,InQuoy and Gaimard, 1824). . . . . . . . . . . . . . . . . . . . . . . 115SqualiolusSmith and Radcliffe, 1912. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117SqualiolusaliaeTeng, 1959 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118

2.3 Order PRISTIOPHORIFORMES Sawsharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120

2.3.1 Family PRISTIOPHORIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121

PliotremaRegan, 1906 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122PliotremawarreniRegan, 1906 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123PristiophorusMller and Henle, 1837 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125PristiophorusnancyaeEbert and Cailliet, 2011. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125

2.4 Order SQUATINIFORMES Angel sharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128

2.4.1 Family SQUATINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129

SquatinaDumril, 1806 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130Squatina africanaRegan, 1908 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131

2.5 Order HETERODONTIFORMES Bullhead sharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134

2.5.1 Family HETERODONTIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135

HeterodontusBlainville, 1816 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135Heterodontusramalheira(Smith, 1949) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136

2.6 Order ORECTOLOBIFORMES Carpet sharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138

2.6.1 Family PARASCYLLIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139

ParascylliumGill, 1862 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140ParascylliumsparsimaculatumGoto and Last, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . 141

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2.7 Order LAMNIFORMES Mackerel sharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142

2.7.1 Family ODONTASPIDIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144

OdontaspisAgassiz, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146Odontaspis ferox(Risso, 1810) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146

2.7.2 Family MITSUKURINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148

MitsukurinaJordan, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149Mitsukurina owstoniJordan, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

2.7.3 Family PSEUDOCARCHARIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151

PseudocarchariasCadenat, 1963 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152Pseudocarchariaskamoharai(Matsubara, 1936). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152

2.7.4 Family ALOPIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154

AlopiasRanesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155

Alopias superciliosusLowe, 1841 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155

2.7.5 Family CETORHINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157

CetorhinusBlainville, 1816 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158Cetorhinus maximus(Gunnerus, 1765) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158

2.8 Order CARCHARHINIFORMES Ground sharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161

2.8.1 Family SCYLIORHINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164

ApristurusGarman, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166Apristurus microps(Gilchrist, 1922) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169

BythaelurusCompagno, 1988. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171Bythaelurushispidus(Alcock, 1891) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172CephaloscylliumGill, 1862. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173CephaloscylliumalbipinnumLast, Motomura and White, 2008 . . . . . . . . . . . . . . . . . . . 175FigaroWhitley, 1928 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176Figaroboardmani(Whitley, 1928) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177GaleusRanesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178Galeus gracilisCompagno and Stevens, 1993. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179

HolohalaelurusFowler, 1934 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180HolohalaelurusfavusHuman, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182Holohalaeluruspunctatus(Gilchrist, 1914) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183ScyliorhinusBlainville, 1816 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184

Scyliorhinus capensis(Smith inMller and Henle, 1838) . . . . . . . . . . . . . . . . . . . . . . . 186

2.8.2 Family PROSCYLLIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187

CtenacisCompagno, 1973 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189Ctenacisfehlmanni(Springer, 1968) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189

EridacnisSmith, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190EridacnisradcliffeiSmith, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191

2.8.3 Family PSEUDOTRIAKIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192

PlanonasusWeigmann, Stehmann, and Thiel, 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . 194Planonasusparini Weigmann, Stehmann, and Thiel, 2013. . . . . . . . . . . . . . . . . . . . . . . 194Pseudotriakisde BritoCapello, 1868 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195Pseudotriakis microdonde Brito Capello, 1868. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195

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2.8.4 Family TRIAKIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197

Iago Compagno and Springer, 1971 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199Iago omanensis(Norman, 1939) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200MustelusLinck, 1790 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201Mustelus stevensiWhite and Last, 2008. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202

2.8.5 Family CARCHARHINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203

CarcharhinusBlainville, 1816 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205Carcharhinus altimus(Springer, 1950) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206

3. BIBLIOGRAPHY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208

4. INDEX OF SCIENTIFIC AND VERNACULAR NAMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239

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1. INTRODUCTION

The genesis of this Catalogue stems from a workshopon Deepsea Species Identication held in Rome, Italy,from 2 to 4 December 2009 (FAO, 2011). The workshopobjectives were to identify and review key issues forvulnerable deepsea species, e.g. corals, sponges, andchondrichthyans, that could be addressed when developingidentication tools to assist in implementing the FAOGuidelines for the management of deepsea sheries andenhance sheries management tools in general. A globalchecklist of deepsea chondrichthyans, that occur in thedeepsea from about 200 m to over 2000 m deep, wascompiled in relation to the 19 FAO major shing areas.Among the recommendations from the workshop wasthe production of comprehensive regional identicationguides to the deepsea chondrichthyan fauna. The presentcatalogue covers the deepsea Indian Ocean, primarilyFAO Fishing Areas 51 and 57, and that part of Area 47that extends from Cape Point, South Africa to the east, e.g.

the extreme southwestern Indian Ocean. The cataloguehas been divided into two volumes, the rst on the sharks(presented below) and a second volume on the batoidsand chimaeras. The catalogue includes species of major,moderate, minor, and minimal importance to sheries aswell as those of doubtful or potential use to sheries. It alsocovers those little known species that may be of research,educational, and ecological importance. The catalogue isintended to be a comprehensive review of the sharklikeshes of the deepsea Indian Ocean in a form accessibleto sheries workers as well as researchers on sharksystematics, biodiversity, distribution, and general biology.It also caters to other researchers that need comparativeinformation on sharks, and their relatives, and to peoplewho encounter sharks in the sea, and the general public.

Biogeography of Region. The Indian Ocean regionincludes two major FAO Fishing Areas, the western IndianOcean (Area 51) and the eastern Indian Ocean (Area 57).The western Indian Ocean extends from the southeast coastof India at 7700E longitude where the states of Kerala andTamil Nadu meet at the sea, and extends southward to theequator, then east to 8000E longitude and south to 4500Slatitude where it runs parallel westward to 3000E longitudeand north to the coast of Africa. Also, included in this regionis the extreme southeastern portion of Area 47, extendingfrom Cape Point at about 1842E longitude eastwards to

the boundary of Area 51 at 3000E longitude. The easternIndian Ocean region extends essentially from the boundarywith Area 51 on its western edge, although it extends furthersouth to 5500S latitude, where it runs parallel to 15000Elongitude and then north to about 3731S latitude to thesoutheast coast of Australia. The eastern boundary of Area57 runs along the coast of southern Australia and aroundto the boundary of the states of Western Australia and theNorthern Territory at 12900E longitude. At that boundaryit extends northwards to 0800S latitude and then westalong 11328E longitude where it meets the south coastof Java at 0823S latitude and extends westwards alongthe coasts of Java and Sumatra running south in the Straitof Malacca, where it crosses the Strait at 0230N latitudeto meet the coast of the Malay Peninsula, where it goes ina northerly and westerly direction along the coasts facingthe Bay of Bengal to the point of departure. The IndianOcean encompasses 10 Large Marine Ecosystems, ve

in the western Indian Ocean and six in the eastern IndianOcean; one of the large marine ecosystems crosses theboundary of both areas off India. This includes the AgulhasCurrent, Somali Coastal Current, Arabian Sea, Red Sea,and a portion of the Bay of Bengal large marine ecosystemsin the western Indian Ocean (Area 51), and in the easternIndian Ocean (Area 57) this includes most of the Bay ofBengal, North Australia, Northwest Australia, WestCentralAustralia, Southwest Australia, and Southeast Australialarge marine ecosystems (Figure 1 Map of the IndianOcean FAO Areas 51 and 57).

The Indian Ocean is the warmest ocean with a minimumsurface temperature north of 20S of 22 C, but oftenexceeding 28 C. Two major circular ocean currents, aclockwise owing current in the northern hemisphere anda counterclockwise owing current south of the equator,dominate the ow patterns. However, during the winter

monsoon season, which heavily inuences these currents,the northern current reverses ow. The continental shelvessurrounding the Indian Ocean are generally narrow,averaging 200 km in width, except off the western coastof Australia where the shelf exceeds 1,000 km. Theaverage depth is 3,890 m with a maximum depth of 8,047m in the Diamantia Trench. The Indian Ocean topographyis quite dynamic as three crustal plates, African, Indianand Antarctic, converge with their junctures forming aninverted Y with the stem originating from the edge of thecontinental shelf off Mumbai, India and extending to thesouthern Indian Ocean where all three plates meet at ageologic feature called Rodrigues Triple Point, named afterthe island of Rodrigues that is located nearby. This geologicfeature subdivides the Indian Ocean into three basins,eastern, western and southern. Geologically the region,as the youngest of the major oceans, has active spreadingridges with several distinct subsurface features includingthe Ninety East Ridge that runs northsouth at the 90Emeridian, which bisects the Indian Ocean into its easternand western halves.

Classication and systematic arrangement used here.The higher classication of these shes includes the classChondrichthyes that is divided into two major groups,each with a long and separate preDevonian history,the chimaeroids, Holocephali (with a single living order

Chimaeriformes), and the sharks and batoids proper, withthe surviving group Neoselachii divided into two cohorts,the Selachii (sharks) and the Batoidea (rays and skates)and includes all of the modern living species. The Selachiiis further divided into two superorders, the Squalomorphiiand Galeomorphii. The superorder Squalomorphii includesthe orders Hexanchiformes, Squaliformes, Squatiniformes,and Pristiophoriformes, while the superorder Galeomorphiiincludes the Heterodontiformes, Orectolobiformes,Lamniformes, and Carcharhiniformes. The cohort Batoidearecognizes four orders, Torpediniformes, Pristiformes,Rajiformes, and Myliobatiformes. The ordinal classicationof the sharklike shes largely follows the arrangement ofCompagno (2001, 2005), Ebert and Compagno (In press), andEbert (In preparation) with some modications in recognizingeight orders and 34 families. The higher classication andthe taxonomic arrangement of the batoid shes and thechimaeras will be discussed in volume 2 of this catalogue.

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The following classication to order is based on the abovediscussion on higher ordinal classications (* starred ordersare covered in this volume):

Class Chondrichthyes (cartilaginous shes) Subclass Holocephali (chimaeras and fossil relatives)

Order Chimaeriformes (chimaeras or silver sharks)

Subclass Neoselachii (modern sharks and batoids) Cohort Selachii (modern sharks) Superorder Squalomorphii (squalomorph sharks) Order Hexanchiformes (cow and frilled sharks)* Order Squaliformes (dogsh sharks)*

Order Squatiniformes (angel sharks)*Order Pristiophoriformes (sawsharks)*

Superorder Galeomorphii (galeomorph sharks)Order Heterodontiformes (bullhead sharks)*

Order Lamniformes (mackerel sharks)*Order Orectolobiformes (carpet sharks)*

Order Carcharhiniformes (ground sharks)* Cohort Batoidea (batoids)

Order Torpediniformes (electric rays)

Order Pristiformes (sawshes) Order Rajiformes (skates and guitarshes) Order Myliobatiformes (stingrays)

Indian Ocean Deepsea Biodiversity. Sharks are amongone the most successful groups of shes having penetratedmost marine ecosystems, including continental and insularshelf waters from the intertidal out to, and into, the deepseaas well as occupying oceanic and pelagic habitats; somespecies have even penetrated into and occupy estuarineand freshwater river ecosystems. Recent reviews on thebiodiversity and distribution of sharks in various ecosystemshas shown that tropical marine ecosystems have the highest

diversity, followed by deepsea and temperate ecosystems,and with the epipelagic and freshwater ecosystems havinga lower diversity of species (Ebert and Winton, 2010; Kyneand Simpfendorfer, 2010; Rosa, Charvet-Almeida andQuijada, 2010; Stevens, 2010; White and Sommerville,2010). Worldwide there are more than 500 known living andvalid species of sharks comprising 8 orders, 34 families, and107 genera, with additional species requiring description(Nelson, 2006; Ebert and Winton, 2010; D.A. Ebert, pers.database, 1 March 2013). Globally, of these totals, all eightorders, 23 families, 58 genera, and 265 sharks species areconsidered to primarily inhabit the deepsea (FAO, 2011;D.A. Ebert, pers. database, 1 March 2013).

The deepsea Indian Ocean has a very diverse sharkfauna with 8 orders, 23 families, 46 genera, and at least 117species being represented (Table 1); while this cataloguewas in preparation a new genus and species of deepsea Pseudotriakidae from the northern Indian Ocean wasdescribed,Planonasus pariniWeigmann, Stehmann, andThiel, 2013, and is included here. The most specious group ofsharks in the deepsea Indian Ocean are the Squaliformesthat have at least 58 species represented, most of which arein the families Etmopteridae (n = 15) and the Centrophoridaeand Squalidae with 13 species in each family. TheCarcharhiniformes have 41 species representatives, withthe vast majority in the family Scyliorhinidae (n = 32). All

of the other shark orders only have six of fewer speciesrepresentatives. Of these totals, the diversity is remarkablysimilar in terms of numbers of families, genera, and speciesbetween the eastern and western Indian Ocean deepsea

(Table 1). The only families not represented in the easternIndian Ocean are the Pristiophoridae and Heterodontidae,while in the western Indian Ocean only the Parascylliidaeis not represented. The number of genera occurring in thewestern Indian Ocean (n = 42) is slightly higher than in theeastern Indian Ocean (n = 38), as well as the number ofspecies (n = 78 and 77, respectively). A total of 38 species

occurs in both regions, with the remaining numbers of speciesbeing unique to each region.

Indian Ocean

(Tot.)Sharks Batoids Chimaeras Tot.

Orders 8 3 1 12

Families 23 8 2 33

Genera 46 23 5 74

Species 117 61 17 195

Indian Ocean

(Area 51)

Orders 7 3 1 11

Families 22 5 2 29

Genera 42 14 5 61

Species 78 30 9 117

Indian Ocean

(Area 57)

Orders 6 3 1 10

Families 21 8 2 31

Genera 38 17 4 59

Species 77 34 11 122

Table 1 The families, genera, and species represented

within the Deepsea Indian Ocean (total for Areas 51and 57), western Indian Ocean (Area 51), and easternIndian Ocean (Area 57).

1.1 Plan of the Catalogue

The format for this catalogue follows that of the FAOCatalogue of Sharks of the World (Compagno, 2001; Ebertand Compagno, In press; Ebert, In preparation), with ordersas the highest taxonomic group dealt with here, followed byfamily, genus, and species accounts. A key to the familiesand genera, where appropriate is also included. A differencein the present catalogue is that not all species accountsare dealt with in the same comprehensive detail. A list of alldeepsea species known to occur in the Indian Ocean ispresented for each family, with a representative species ofeach genus presented in detail and indicated by the presenceof a shark icon beside its scientic name. For those generawhere more than a single member of the genus has speciesthat are possibly subjected to intense shing pressures orare considered especially vulnerable (e.g. Centrophorus,Squalus), more than one species is presented in detail. Intotal, 51 of 117 shark species are presented in detail here.

The species specic information on the biology, conservationstatus, distribution, habitat, sheries, and systematics ofIndian Ocean sharks was compiled from primary literature

sources including, but not limited to, Compagno (2001),Compagno, Ebert and Smale (1989), Last, White andPogonosky (2007), Last, Motomura and White (2008), LastSret and White (2008), Last and Stevens (2009), Ebert and

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Winton (2010), Kyne and Simpfendorfer (2010), FAO (2011),and Ebert and Compagno (In press). Electronic sources werealso of invaluable help, these included, but were not limitedto, the California Academy of Sciences Catalogue of Fishes(http://www.calacademy.org/research/ichthyology/catalog/shcatsearch.html) and IUCN Shark Specialist Group (http://www.iucnredlist.org). A comprehensive bibliography of the

literature, including primary, grey, and electronic sources isprovided at the end of this volume.

Order accounts include the valid modern form of the ordername with author and year; the original citation of the ordername with its author, year, reference and pagination; thenumber of recognized families in the Indian Ocean deepsea; common order Synonyms mainly from the IndianOcean deepsea region with the name, author, year, andpagination; the FAO order Vernacular Names in English,French and Spanish; Field Marks and Diagnostic Featuresof members of the order; an account of the natural historyof the order under separate sections covering Distribution,Habitat and Biology; a section on Interest to Fisheries and

Human Impact, a synopsis of the human issues affectingshark families; Local Names when available; a Remarkssection mostly with systematic comments; and a Key toDeepsea Indian Ocean Families, when orders have morethan one family.

Family accounts include the valid modern form of thefamily name with author and year; the original citation of thefamily name with its author, year, reference and pagination;the valid type genus with author and date; the number ofrecognized deepsea Indian Ocean genera in the family;family Synonyms with names mostly associated with theIndian Ocean region and with the name, author, year, and

pagination; the FAO family Vernacular Names in English,French and Spanish; Field Marks and Diagnostic Featuresof members of the family; an account of the natural historyof the family under separate sections covering Distribution,Habitat and Biology; a section on Interest to Fisheries andHuman Impact, a synopsis of the human issues affectingshark families; Local Names when available; a Remarkssection mostly with systematic comments; a Literaturesection covering references to the entire family; and a Key toDeepsea Indian Ocean Genera, when families have morethan one genus.Generic accounts include the valid modern form of thegenus name with author and year; the original citation of the

genus (or subgenus), with its author, year, reference andpagination, and, if a subgenus, the original genus name withauthor and year that the subgenus was originally placed in;the type species and means of designating it (for example,by original designation, monotypy, absolute tautonymy, orsubsequent designation); the number of recognized IndianOcean species in the genus; the Synonyms of genera, withtheir rank (genus, subgenus, or other genusgroup ranking),author, year, pagination, and genus they were describedin, if originally ranked as subgenera or equivalents; FAONames if they exist; Field Marks if genera are large anddistinctive; Diagnostic Features of the genus; Local Nameswhere available; a Key to Deepsea Indian Ocean Species

if the genus has more than one species (is not monotypic);and a Remarks section.

Species accountsinclude the valid modern names of the

species, with author and date; the original citation of thespecies, with its author, year, reference pagination; theholotype, syntypes, lectotype or neotype of each species(paratypes are not listed in the present account), includingthe total length and sex of the specimen, its institutionaldeposition, and its catalogue number; the type localityincluding the location, coordinates and depth if available,

where the holotype, syntypes, lectotype or neotype werecaught; Synonyms of the species, including their names,authors and dates; a section listing other scientic namesrecently in use; the English, French, and Spanish FAONames for the species; a lateral view illustration, and oftenother useful illustrations (lateral view drawings are givenof each shark species, usually ventral views of heads,and often teeth and denticles of the shark in question);Field Marks; Diagnostic Features (except in monotypicgenera); Distribution, including a map; Habitat; Biology;Size; Interest to Fisheries and Human Impact; Local Nameswhen available; a Remarks section when necessary; andLiterature.

Synonyms commonly seen in the Indian Ocean literatureare listed, where appropriate, and include only truetaxonomic synonyms of the valid family, genus and speciesgiven. For species, another category, Other Combinations,is provided for common misidentications of a given specieswith another, valid species, as well as commonly usedcombinations that place a valid species in different genera.

FAO Family and Species Names. English, French andSpanish names for each family and species, primarily for usewithin FAO, were selected by the following criteria: (a) eachname applies to a single family or species worldwide; (b)the name conforms with FAO spelling nomenclature; (c) the

name conforms to prior usage when possible. FAO namesare not intended to replace local species names, but arenecessary to overcome the confusion caused by the use ofa single name for more than one species or several namesfor one species. The FAO names used here conform to priorFAO usage. The common French and Spanish names ofspecies from other FAO Catalogues, including the Sharks ofWorld (Compagno, 1984a, b, 2001; Ebert and Compagno, Inpress, Ebert, In preparation), and regional FAO Catalogueson the Sharks of the Western Indian Ocean (Compagno,1984c), Namibia (Bianchi et al., 1999), Western CentralPacic (Compagno, 1998), Red Sea and Gulf of Aden(Bonl and Abdallah, 2004), and North Atlantic (Ebert andStehmann, 2013), were used when appropriate.

Keys, Field Marks, and Diagnostic Features. Thesesections include identication data in different forms. Keysto families, genera and species are standard dichotomousbiological keys that are followed in steps of alternate choicesto single out the taxa covered. It should be noted that theKeys include only those families, genera, and species thatoccur in FAO Areas 51 and 57, or within the scope of thepresent Catalogue and does not include those groups notoccurring with the area. Field Marks generally include a fewobvious characters of use in eld identication, extractedfrom Diagnostic Features at various levels, but included in aseparate section. Field Marks are listed at the ordinal, familial

and species levels, and occasionally the generic level incases of large genera with many species. The arrangementof Field Mark characters is semihierarchical and pragmaticand may include characters from a higher level such as an



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order in lower level taxonomic accounts such as those ofspecies. Field Marks include characters that are obviousin live or freshcaught individuals but may be obscure infrozen or preserved material. Diagnostic Features are listsof characters at the ordinal, familial, generic, and specieslevel, with the character choice generally limited to externalcharacters, particularly at the species level, because of

their primary purpose of identication rather than indicationof relationships. The Diagnostic Features sections arehierarchical, with characters at the ordinal level generallynot duplicated at the family, genus and species levels.Monotypic orders with one family (such as Squatiniformes),monotypic families with one genus (Mitsukurinidae) ormonotypic genera with one species (Pseudotriakis) allhave the Diagnostic Features section present only in thehighest taxon covered.

Distribution.Geographic distributions for nearly all speciesof sharks are given by listing the countries or oceanographicfeatures, e.g. seamounts and troughs, off the coasts of whichthe sharks occur, and, in some instances with large countries

(Australia), more detailed data are given when available.In compiling distributional data and preparing maps it wasnoted that the distributions of many wideranging coastaland deepsea species are very patchily known as present.In many cases gaps in distribution may not indicate absenceof a given species but absence of knowledge. Continentalslope shark faunas are poorly known for much of the world,and a number of deepwater species probably have widerranges than are currently known. Much effort was made toscreen out distribution errors, based on misidentications ofspecies, at a cost of presenting distributional lists and mapsthat are patchy, but possibly more accurate.

Habitat.Habitat covers information on physical conditionswhere various sharks are found. The known depth range ofthe species (in metres), position in the water column, type ofsubstrate occupied, and preferences relative to coasts arenoted when available. In most cases data on salinity, oxygencontent, and specic temperature of the water in which theyoccur was not available or was not in an easily usable formand has not been regularly compiled here.

Biology. Includes data on reproduction, age and growth,diet, and behaviour and movement patterns. Compilationof these data suggests that very few deepsea species arebiologically well known, and several are known only a veryfew specimens that have ever been observed.

Size. All size data are given as total lengths; this is themeasurement most often used as an independent variableand standard measurement in the shark literature, althoughparticularly in sheries papers precaudal lengths, forklengths, and other measurements have been used fromchoice or necessity. Unfortunately shark workers have notagreed on a standard method of measuring total length, sototal lengths from different sources in the literature may notbe strictly comparable. I prefer and advocate as a standardmethod a direct measurement, in which the shark is heldbelly down with its dorsal caudaln lobe depressed into linewith its body axis and total length measured as a point to

point distance (not over the curve of the body) from the snouttip to the tip of the dorsal caudaln lobe. This method lendsitself readily to quick use of a shboard with a perpendicularfront bar or plate to index the shs snout against, a one

metre or two metre ruler or folding ruler slipped under theshark, batoid, or chimaera or even a steel or cloth tape, andavoids the trouble of computation and possible errors andloss of data.

Total length data presented includes maximum size, sizeat maturity (in some cases, a size range at maturity, when

abundant data were available) and maximum size forboth sexes, and size at birth or hatching. Sometimes sizeat sexual maturity for either or both sexes is not known,in which cases reported minimum and maximum sizes ofadult individuals are given. In some cases maximum sizeexceeds that recorded for either sex, in which case the sexof the outsized individual or individuals representing themaximum size measurements was not indicated. In somepoorly known species only immature individuals are known,in which case the hypothetical maximum adult size is almostcertainly larger than the known immature maximum.

Interest to Fisheries and Human Impact. This sectionincludes Fisheries information, including whether the species

is taken in targeted or nontargeted (bycatch) sheries and iftaken as bycatch whether it is retained or discarded. Data onlocalities of sheries, gear used, and uses of the particularspecies are noted when available. National sheries datafor deepsea sharks is often sketchy and combined for anumber of species. Thus, catch statistics are generallyunavailable except for relatively few species of sharks butare noted when available, with particular emphasis on datafrom those species reported to FAO. Additional data areincreasingly available from national and regional sheriesbodies are presented when available. Other aspects ofhuman interaction is presented if available or known,although the average person rarely encounters most of

these deepsea sharks. The current conservation Red Liststatus of each species as evaluated by the IUCN SpeciesSurvival Commissions Shark Specialist Group (http://www.iucnredlist.org) is provided. At the end of the Bibliographysection an electronic reference section has been added witha link to the Red List Assessment for each species includedin the Catalogue.

Local Names. Many deepsea species have no vernacularnames whatsoever or are lumped under catchall names,while some species such as the basking sharks havedozens of names. Wherever possible common local namesare presented, especially for important widerangingsharks. The broadening interest in sharks and urgent need

to acquire speciesspecic data for their management andconservation should encourage sheries biologists andother researchers to compile local names for their owncountries or regions, and add to the sketchy knowledge oflocal names.

Remarks.Important information, especially on systematicsand nomenclature, are given in the remarks section. Also,the relative number of families per order, genera per family,and species per genus worldwide is given when appropriatefor comparison to the groups occurring within FAO Areas 51and 57.

Literature. References cited here include specic workswith important information for each species and family aswell as comprehensive accounts, but are not intended as acomprehensive bibliography.



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Fig. 1 Map of FAO Area 51 (Western Indian Ocean), Area 57 (Eastern Indian Ocean) and the

southeastern portion of Area 47 (Southeastern Atlantic Ocean)



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1.2 Technical Terms and Measurements

1.2.1 Picture Guide to External Terminology of Sharks

Fig. 2 Lateral view

head trunk

mouth

labial

furrows

gill

openings

pectoral n

pelvic n anal nclasper

(males)

caudal keel

precaudal pit

second dorsal n

rst dorsal ndorsaln

spine

caudal n

snout nostril eye spiracle

tail

gill slits

Fig. 3 Ventral view

trunk

mouth

pectoral n

anal ncaudal n

preanal ridges

pelvic n

(female, no claspers)

vent

snout nostrilprecaudal tail

Fig. 4 Head of an orectoloboid shark(ventral view)

Fig. 5 Nostril

incurrent aperture

excurrent aperture

anteriornasal ap

posteriornasal ap

mouth

nasoral groove

symphisial groove

anterior nasal ap

lower labial furrow

upper labial furrowbarbel

circumnarial

fold

circumnarial

groove

incurrentaperture

excurrentaperture

anterior nasal

ap (lifted)



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labial furrow

labial fold

n insertion

n origin

inner margin

free rear tip

posteriormarginanterior

margin

apex

base

spine

anteriormargin

origin insertioninner margin

free reartip

posterior

margin

apex

base

upper eye lid

secondaryeyelid

nictitatinglower eyelid

subocularpocket

notch

POSTERIOR

MEDIAL LATERAL

ANTERIOR

lower origin

preventralmargin

upper origin

dorsal margin

epaxial web

dorsal lobe

terminal lobe posterior tip

terminal margin

subterminal marginsubterminal notch

upper postventral margin

posterior notch

lower postventral margin

ventral tip

ventral lobe

hypaxial web

clasper tip

rhipidion

pseudosiphon

coverrhipidion

apopyle pelvic n

claspergroove

clasperspur

hypopyle

lateral fold

claspershaft

clasperglans

Fig. 6 Eyes

Fig. 7 Mouth corner

Fig. 8 Dorsal n

Fig. 9 Caudal n

Fig. 10 Pectoral nFig. 11 Dorsal view of clasper

(lamnid shark)



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1.2.2 Picture Guide to Skeletal Terminology of Sharks

internasal plate

rostrum

rostrum

rostralnode

nasal apertureorbital notch

optic nerve foramen

preorbital process

anterior fontanelle

cranial roof

parietal fossa

foramen magnum occipital centrum

hyomandibular facet

pterotic horn

sphenopterotic ridge

otic capsule

cranial roof

stapedialfenestrasuborbital

shelf

carotid foramen

stapedial fenestra

basal plate

orbitalnotch

nasal aperture

supraorbitalcrest

supraorbitalcrest

orbitnasalcapsule

suborbitalshelf

subethmoid fossanasal capsule

nasalfontanelle

hyomandibularfacet

otic capsule

postorbital

process

a) DORSAL VIEW b) VENTRAL VIEW

c) LATERAL VIEW

POSTERIOR

POSTERIORANTERIOR

ANTERIOR

Fig. 12 Chondrocranium

distal radials

intermediate radials

metapterygial axis

metapterygium

propterygium

mesopterygium

metapterygium

proximal radials

intermediate radialsdistal radials

radials

propterygium

radials

basalsbasals

a) APLESODIC b) PLESODIC

metapterygialaxis

Fig. 13 Aplesodic and plesodic pectoral ns



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betacartilage

basipterygium

intermediatesegments (2)

axialcartilage

axial cartilage

apopyle

ventralmarginal

ventralmarginal

ventralmarginal

terminal 3cartilage

dorsalmarginal

dorsalmarginal

dorsalmarginal

claspergroove

hypopyle

dorsalterminal

dorsalterminal

dorsalterminal

ventralterminal

ventralterminal

transversenotch

cuttingedge

mesial rootlobe

root

neck

blade

cusplet

cusplet

central foramen

transverse groovetransverseridges

serrations

root

root

crown

crown

pedicel

base

medial ridge

medial cusp

lateral cusplateral ridge

root

crowncusplet

cuspcusp

crown foot

crown foot

basal groove

basalledge

distal rootlobe

ventralterminal

endstyle ofaxial cartilage

endstyle ofaxial cartilage

appendixstemof axial cartilage

folded

erect

dorsalterminal 2

b) VENTRAL

b) DORSAL

a) LABIAL VIEW b) LINGUAL VIEW

DISTAL

BASAL

APICA L

ANTERIOR

POSTERIOR

MESIAL MESIAL

c) DORSAL (TERMINAL 3 CARTILAGE ANDDORSAL TERMINAL 2 REMOVED)

MEDIAL

MEDIAL

MEDIAL

LATERAL

LATERAL

LATERAL

Fig. 14 Clasper skeleton of lamnid shark (right side)

Fig. 15 Tooth terminology (left upper anterolateral tooth)

Fig. 16 Oblique anterolateral view of lateral trunk dermal denticle



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1.2.3 Measurements Used for Sharks

TL

FL

PCL

PD2

PD1

HDL

PG1

PSP

POB

PP1

PP2

SVL

PAL VCL

PPSPAS

IDS

DCS

ACS

PCA

PRNPOR

EYLEYHINGGS1GS2GS3GS4GS5GS6GS7P1AP1RP1BP1IP1PP1HP1LSOD

= PRENARIAL LENGTH= PREORAL LENGTH

= EYE LENGTH= EYE HEIGHT= INTERGILL LENGTH= FIRST GILL SLIT HEIGHT= SECOND GILL SLIT HEIGHT= THIRD GILL SLIT HEIGHT= FOURTH GILL SLIT HEIGHT= FIFTH GILL SLIT HEIGHT= SIXTH GILL SLIT HEIGHT= SEVENTH GILL SLIT HEIGHT= PECTORALFIN ANTERIOR MARGIN= PECTORALFIN RADIAL LENGTH= PECTORALFIN BASE= PECTORALFIN INNER MARGIN= PECTORALFIN POSTERIOR MARGIN= PECTORALFIN HEIGHT= PECTORALFIN LENGTH= SUBOCULAR POCKET DEPTH

TLFLPCLPD2PD1

HDLPG1PSPPOBPP1

= TOTAL LENGTH= FORK LENGTH= PRECAUDALFIN LENGTH= PRESECOND DORSALFIN LENGTH= PREFIRST DORSALFIN LENGTH

= HEAD LENGTH= PREBRANCHIAL LENGTH= PRESPIRACULAR LENGTH= PREORBITAL LENGTH= PREPECTORALFIN LENGTH

PP2SVLPALIDSDCS

PPSPASACSPCAVCL

= PREPELVICFIN LENGTH= SNOUTVENT LENGTH= PREANALFIN LENGTH= INTERDORSAL SPACE= DORSAL CAUDALFIN SPACE

= PECTORALFIN PELVICFIN SPACE= PELVICFIN ANALFIN SPACE= ANALFIN CAUDALFIN SPACE= PELVICFIN CAUDALFIN SPACE= VENT CAUDALFIN LENGTH

Fig. 17 Main longitudinal measures

Fig. 18 Measurements of pectoral n,gill slits, eye and snout

EYL

GS1

GS5

EYH

ING

POR

SOD

P1L

P1R

P1A P1P

P1H

P1IPRN

P1B



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= FIRST DORSALFIN MIDPOINT PECTORALFIN INSERTION= FIRST DORSALFIN MIDPOINT PELVICFIN ORIGIN= PELVICFIN MIDPOINT FIRST DORSALFIN

INSERTION= PELVICFIN MIDPOINT SECOND DORSALFIN

ORIGIN

PDI PDO

DPI

TRH

ABH

TAH

CPH

HDH

DPO

DAO DAI

D1L

D1H

D1P

D1A

D1I

P2B

P2H

ANH

ANP

P2A

ANA

P2P

ANB

P2I

ANIP2L

ANL

D2LD2B

D2A

D2H D

2P

D2I

D1B

CDM

CPU

CTL

CTR

CST

CPV

CPL

CFL

CFW

CSW

CDMCPVCPUCPLCFWCFLCSTCSW

CTRCTL

= DORSAL CAUDALFIN MARGIN= PREVENTRAL CAUDALFIN MARGIN= UPPER POSTVENTRAL CAUDALFIN MARGIN= LOWER POSTVENTRAL CAUDALFIN MARGIN= CAUDALFIN FORK WIDTH= CAUDALFIN FORK LENGTH= SUBTERMINAL CAUDALFIN MARGIN= SUBTERMINAL CAUDALFIN WIDTH

= TERMINAL CAUDALFIN MARGIN= TERMINAL CAUDALFIN LOBE

= FIRST DORSALFIN LENGTH= FIRST DORSALFIN ANTERIOR MARGIN= FIRST DORSALFIN BASE= FIRST DORSALFIN HEIGHT= FIRST DORSALFIN INNER MARGIN= FIRST DORSALFIN POSTERIOR MARGIN

= SECOND DORSALFIN LENGTH= SECOND DORSALFIN ANTERIOR MARGIN= SECOND DORSALFIN BASE= SECOND DORSALFIN HEIGHT

= SECOND DORSALFIN INNER MARGIN= SECOND DORSALFIN POSTERIOR MARGIN

= PELVICFIN LENGTH= PELVICFIN ANTERIOR MARGIN= PELVICFIN BASE= PELVICFIN HEIGHT= PELVICFIN INNER MARGIN [LENGTH]= PELVICFIN POSTERIOR MARGIN [LENGTH]

= ANALFIN LENGTH= ANALFIN ANTERIOR MARGIN= ANALFIN BASE= ANALFIN HEIGHT= ANALFIN INNER MARGIN

= ANALFIN POSTERIOR MARGIN

D1LD1AD1BD1HD1ID1P

D2LD2AD2BD2HD2ID2P

P2LP2AP2BP2HP2IP2P

ANLANAANBANHANI

ANP

HDHTRHABHTAH

CPHDAI

DAO

DPIDPO

PDI

PDO

= HEAD HEIGHT= TRUNK HEIGHT= ABDOMEN HEIGHT= TAIL HEIGHT

= CAUDALFIN PEDUNCLE HEIGHT= SECOND DORSALFIN INSERTION ANALFIN

INSERTION= SECOND DORSALFIN ORIGIN ANALFIN ORIGIN

Fig. 19 Measurements of caudal n

Fig. 20 Measurements of dorsal, pelvic and anal ns

Fig. 21 Other common measurements (lateral view)



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CLO

CLI

NOW

ANF

I

NW MOL

M

OW

LLA

ULA

GIR

HDW

TRW

ABW

TAW

CPWIN

O

SPL

ESL

CLB

MOLMOWULALLANOWINWANF

CLOCLICLB

= CLASPER OUTER LENGTH= CLASPER INNER LENGTH= CLASPER BASE WIDTH

= MOUTH LENGTH= MOUTH WIDTH= UPPER LABIALFURROW LENGTH= LOWER LABIALFURROW LENGTH= NOSTRIL WIDTH= INTERNARIAL SPACE= ANTERIOR NASALFLAP LENGTH

INOSPLESLHDWTRWABWTAWCPW

= INTERORBITAL SPACE= SPIRACLE LENGTH= EYE SPIRACLE SPACE= HEAD WIDTH= TRUNK WIDTH= ABDOMEN WIDTH= TAIL WIDTH= CAUDALFIN PEDUNCLE WIDTH

Fig. 22 Other common measurements (ventral and dorsal view)

GIR = GIRTH

b) CLASPER

d) ANGLE OF MOUTHc) VENTRAL VIEW

e) DORSOLATERAL VIEW

f) DORSAL VIEW

a) NOSTRIL



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PICTORIAL KEY OF DEEPSEASHARK ORDERS OCCURRING IN

THE INDIAN OCEAN(not a cladogram)

Fig. 23 Higher classication of Sharks (Orders)

SHARKS

HEXANCHIFORMES

SQUALIFORMES

PRISTIOPHORIFORMES

SQUATINIFORMES

CARCHARHINIFORMES

LAMNIFORMES

ORECTOLOBIFORMES

HETERODONTIFORMES

dorsal nspines

no dorsal nspines

5 gill slits2 dorsal ns

6 or 7 gill slits,1 dorsal n

mouth well infront of eyes

mouth behindfront of eyes

nictitating eyelids,spiral or scrollintestinal valve

no nictitatingeyelids, ring

intestinal valve

5 to 7 gillslits; anal n

present

body notraylike

body attened,raylike

snout short, notsawlike

snout elongated,sawlike

mouth terminal,pectoral ns notattached to head

5 or 6 gillslits; no anal

n


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1.2.4 Glossary of Technical Terms

The following glossary of terms used for the anatomyand biology of sharklike shes is modied from termsin Compagno (1984, 1988, 1999) and a short glossary inCompagno, Ebert and Smale (1989). The main glossaryduplicates that in volume 2 of the revised shark catalogue

(Compagno, 2001), except that additional terms for habitatand distribution used in the text are reinstated in the glossary.Terms for photophore patterns (including photomarks) thatpertain only to the family Etmopteridae and primarily to thegenusEtmopterusare discussed under that family.

Abdominal ridges or keels: In some sharks, pairedlongitudinal dermal ridges that extend from the bases of thepectoral ns to the pelvicn bases.

Abyss: The deep sea bottom, ocean basins or abyssalplain descending from 4000 m to about 6000 m.

Accessory dorsal marginal: In the clasper skeleton, a

at cartilage on the posterior end of the dorsal marginalcartilage that supports the cover rhipidion.

Adductor mandibulae muscles: Paired head musclesoriginating on the lateral faces of the quadrate process ofthe palatoquadrates and inserting on the lateral surface ofthe Meckels cartilages; the primary jawclosing muscles ofsharks.

Adelphophagy: Foetuseating, a mode of livebearingreproduction employing uterine cannibalism; early foetusesdeplete their yolk sacs early and subsist by rst eating theirsmaller siblings and then eating nutritive eggs produced by

the mother. At present only known for certain in the sandtiger shark (Carcharias taurus), but suspected in a fewother lamnoids.

Al ternate teeth: Small oral teeth with asymmetrical crownsthat form two interdigitated rows on the symphysis, with thecusps of each row hooked mesially towards the oppositerow. Additional paired rows of alternates may be presentdistal to the symphysial rows.

Amphitemperate: Referring to a species that occurs intemperate water in the northern and southern hemispheres,but is absent from the tropics.

Anal n: A single n on the ventral surface of the tailbetween the pelvic ns and caudal n of some sharks,absent in batoids, dogsh, sawsharks, angel sharks, andsome chimaeras.

Annular rings or annuli: In a vertebral centrum in crosssection, rings of calcied cartilage separated by uncalciedcartilage that occupy the intermedialia only, or concentricrings that cross both the intermedialia and basalia.

Anter ior: Forward, in the longitudinal direction of the snouttip. Also, cranial.

Anter ior fontanel le: On the elasmobranch neurocranium,an aperture on the anterodorsomedial surface, usually atthe rear of the ethmoid region and forming a passage intothe internal cranial cavity. It is closed by a tough membrane,

varies tremendously in shape, and may be pinched off bythe medially expanded orbits in a few sharks.

Anter ior margin:In precaudal ns, the margin from the norigin to its apex.

Anterior nasal ap:A ap on the front edges of the nostrils,

that serves to partially divide the nostril into incurrent andexcurrent apertures or openings.

Anter ior teeth: Enlarged, tall, narrowrooted oral teethnear the symphysis, often with lingually curved cusps.

Anterodorsal palpebral depressor muscle: In theorectoloboid family Parascylliidae, paired head musclesthat originate at the insertions of the preorbitalis muscles onthe anterolateroventral face of the Meckels cartilage, andinsert on the skin of the upper eyelid anterior to the eye.These are possibly for depressing the upper eyelids andclosing the eyes, and are not found in any other sharks.

Antorbital carti lages: On the neurocranium of sawsharksand batoids, separate cartilages attached to the sides of thenasal capsules that support the sides or front of the head.

Apex: In precaudal ns, the distal tip, which can be acutelypointed to broadly rounded.

Apical: In oral teeth, towards the tip of the crown or cusp.Can also be used as indicating direction towards the apexor tip of a n, nspine, etc.

Aplacental v iv ipari ty :Livebearing in which the young donot have a yolksac placenta. Found in all groups of live

bearing sharks.

Aplesodic n:A pectoral, pelvic, dorsal, or anal n in whichthe n radial cartilages do not extend into the distal n weband between the supporting ceratotrichia of the n web.Modern sharks always have aplesodic caudal ns, in whichthe haemal arches of the caudal vertebrae do not supportthe ventral caudal lobe.

Apopyle: The anterior opening of the clasper, on theanteromesial surface of the clasper and close to the vent.The apopyle receives sperm from the cloaca and uidfrom the siphons, which enter the clasper groove and aredischarged through the hypopyle. Apopyle is also used for

clasper skeletons for the anterior opening of the tubularshafts formed by enlarged marginal and axial cartilages.

Axial carti lage: In the clasper skeleton, the elongatedventral rod or plateshaped cartilage that forms the mainsupport of the clasper. Also termed appendixstem .

Barbels: Long conical paired dermal lobes on the snoutsof sharks, that may serve to locate prey. Sawsharks havebarbels on the underside of the snout in front of the nostrilsas in sturgeon, but most barbeled sharks have themassociated with the nostrils, either as an extension of theanterior nasal aps or as separate structures medial to the

nasal apertures.

Basal:In oral teeth, a proximal direction towards the crownfoot and roots.



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Basal cartilages or basals: In precaudal ns the largecartilages of the n bases, immediately distal to the pectoraland pelvic n girdles or the vertebral column (dorsal andanal ns), on which the radials articulate distally. The pairedpectoral ns of living sharks primitively have a tribasalpectoral n, with a propterygium , mesopterygium, andmetapterygiumas basals, although these may be fused;

in batoids, additional neopterygial basalsmay be addedbetween the mesopterygium and metapterygium andthe propterygium is variably expanded anterior witha propterygial basal and axis. The pelvic ns have abasipterygium that supports the pelvic radials and, inmales, the claspers. The caudal n has no basals, but theseare functionally replaced by expanded neural and haemalarches of the vertebral column.

Basal communicating canals:See subnasal fenestrae.

Basal groove:In oral teeth, a deep groove proximal to thebasal ledge on the labial surface of the crown neck andapical root margin.

Basal ledge: In oral teeth, a shelflike projection on thelabial surface of the crown foot.

Basal plate: The oor of the cranial cavity of theneurocranium, a ventral, medial plate extending from theethmoid region between the orbits and otic capsules andbelow the cranial cavity to the occipital condyles, occipitalcentrum and foramen magnum.

Basals or basalia: In a vertebral centrum, the diagonalspaces below the attachment surfaces of the basidorsalcartilages, above the basiventral cartilages, and between

the two halves of the double cone. Basalia may be lledwith uncalcied cartilage, may have diagonal calcicationspenetrating the uncalcied cartilage, or may have calciedannuli or solid calcied cartilage that are continuouswith calcication of the intermedialia. See diagonalcalcicationsand intermedialia.

Base:In precaudal ns, the proximal part of the n betweenthe origin and insertion, extending distally, and supportedby the cartilaginous n skeleton. In the caudal n, thatthickened longitudinal part of the n enclosing the vertebralcolumn and between the epaxial and hypaxial lobes or websof the n. In oral teeth, the proximal root and crown foot,in apposition to the distal cusp. In denticles, the proximal

anchoring structures, often with four or more lobes, holdingthe denticles in the skin.

Basidorsal cartilages: A pair of wedgeshaped arched,thin cartilages articulating with the dorsolateral surfaces ofa vertebral centrum and forming a continuous neural archwith the interdorsal cartilages to protect the spinal cord.

Basipterygium:The large elongate longitudinal cartilage atthe n base of the pelvic n, attached to the posterolateralends of the pelvic girdle or puboischiadic bar. Thebasipterygium has pelvic radials attached along its distaledge and has the clasper skeleton attached posteriorly in

males.

Basiventral cartilages:A pair of rounded or wedgeshapedcartilages on the ventrolateral surfaces of a vertebral

centrum that form the bases for attachment of ribs inmonospondylous precaudal vertebrae. In diplospondylousprecaudal and caudal vertebrae the basiventrals formhaemal arches along with the interventral cartilages forprotecting the caudal artery and vein.

Bathypelagic zone:That part of the oceans beyond the

continental and insular shelves, from about 1000 m to 4000m and above the middle and lower continental rises and theabyssal plain, the sunless zone. Some oceanic sharks maytransit the epipelagic, mesopelagic and bathypelagic zonesto the bottom while migrating vertically.

Batoid:A ray or at or winged shark, a neoselachian of thecohort Batoidea, a sawsh, sharkray, wedgesh, guitarsh,thornray, panray, electric ray, skate, stingray, buttery ray,eagle ray, cownose ray, devil ray or Manta. Rays areclosely allied to the sawsharks (Pristiophoriformes) andangel sharks (Squatiniformes), but differ from them inhaving the pectoral ns fused to the sides of the head overthe gill openings, which are ventral rather than laterally or

ventrolaterally placed.

Benthic or Demersal: referring to organisms that arebottomdwelling.

Beta cartilage:In the clasper skeleton, a single, dorsolateralattened, wedgeshaped or cylindrical cartilage connectingthe pelvic basipterygium and axial cartilage and reinforcingthe intermediate segments, possibly derived from a pelvicradial.

Blade:In oral teeth, an arcuate, convexedged section ofthe cutting edge of the crown foot, without cusplets.

Body ridges:Elongated longitudinal dermal ridges on thesides of the trunk and precaudal tail in certain carpet sharks(Orectolobiformes), in the whale, zebra and some bamboosharks.

Body:Can refer to an entire shark, sometimes restricted tothe trunk and precaudal tail.

Branchial arches: The paired visceral arches behind thehyoid arch and just in front of the scapulocoracoid thatsupport the gills. In elasmobranchs the ve to seven branchialarches primitively consist of a pair of dorsomedial and wedgeshaped cartilages, the pharyngobranchials , closely situated

against the roof of the pharynx, a pair of dorsolateral and morecylindrical epibranchials that are connected dorsomediallyto the pharyngobranchials , a pair of ventrolateral cylindricalceratobranchials that are connected ventrolaterally to theepibranchials, a pair of ventromedial hypobranchials thatare connected ventrolaterally to the ceratobranchials, andunpaired ventromedial basibranchials that are connectedventrolaterally to the hypobranchials. The hypobranchialsand basibranchials along with the expanded ventral endsof the ceratobranchials form the basibranchial skeletonof the oor of the branchial pharynx. The branchial skeletonis variably modied in elasmobranchs, with basibranchialsand sometimes hypobranchials often lost, the last two

pharyngobranchials and the last epibranchial often fusedtogether, and the last basibranchial often expanded into along, broad copula with which the anterior hypobranchialsand posterior ceratobranchials articulate.



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Calcied cartilage:Shark skeletons are formed of hyalinecartilage or gristle, but this is often reinforced with layersof calcied cartilage, cartilage impregnated with a mineral,hydroxyapatite, similar to that of bone but organizeddifferently, in a hard, tilelike pavement of tiny tesserae,or more compactly as in the calcied structures of vertebralcentra.

Calcied double cones: In vertebrae, the primarycalcications of the notochordal sheath, in lateral viewresembling two hollow, horizontal cones with their apicesmerged, or an hourglass.

Cannibal viviparity:See uterine cannibalism.

Carcharhinoid: A ground shark, a member of the orderCarcharhiniformes, and including the catsharks, falsecatsharks, nbacked catsharks, barbeled houndsharks,houndsharks, weasel sharks, requiem sharks andhammerheads.

Carina:On the crowns of oral teeth, a low blunt mesodistalridge replacing the cusp and cutting edge, in sharks that eathardshelled invertebrate prey.

Carotid foramen: A single foramen or one of a pair offoramina that penetrate the basal plate usually near itsmidlength and allow passage of the internal carotid arteriesinto the cranial cavity. In some advanced elasmobranchsthe carotid foramina shift through the stapedial foraminaand onto the medial wall of the orbit.

Cartilaginous shes:Members of the class Chondrichthyes.

Caudal crest: A prominent sawlike row of enlargedpointed denticles along the dorsal caudal margin andsometimes along the ventral caudal margin of the caudaln. Found in certain sharks including hexanchoids andsome carcharhinoids.

Caudal n:The n on the end of the tail in sharklike shes,lost in some batoids.

Caudal keels:A dermal keel on each side of the caudalpeduncle that may extend onto the base of the caudal n,and may, in a few sharks, extend forward as a body keel tothe side of the trunk.

Caudal peduncle:That part of the precaudal tail extendingfrom the insertions of the dorsal and anal ns to the front ofthe caudal n.

Central foramen:In oral teeth, a nutrient foramen on themidline of the lingual surface of the root, in the transversegroove.

Centrum (plural, Centra): A spoolshaped, partially orusually fully calcied structure that forms as a segmentalconstriction in the notochordal sheath of neoselachians, andwhich as an articulated string forms the principal structuralunits of the vertebral column. Centra are primarily formed bythe calcied double cones in the notochordal sheath, whichmay be their only calcication, but additional secondarycalcication may occur in the centrum between the outersurfaces of the calcied double cones, including calciedintermedialia, radii, annuli, and diagonal calcications.

Ceratotrichia: Slender soft or stiff laments of an elasticprotein, supercially resembling keratin or horn, from theGreek keratos, horn, and trichos, hair. Ceratotrichia run inparallel and radial to the n base and support the n webs.The prime ingredient of sharkn soup.

Chimaera: A member of the order Chimaeriformes,

subclass Holocephali, see also Chimaeroid, Holocephali.

Chimaeroid: A chimaera, ratsh, silver shark, ghostshark, spooksh or elephant sh, a member of the orderChimaeriformes.

Chondrichthyan:Referring to the class Chondrichthyes.

Chondrichthyes: The class Chondrichthyes, from Greekchondros, cartilage, and ichthos, sh, a major taxonomicgroup of aquatic, gillbreathing, jawed, nned vertebrateswith primarily cartilaginous skeletons, 1 to 7 externalgill openings, oral teeth in transverse rows on their jaws,and mostly small, toothlike scales or dermal denticles.

Chondrichthyes include the living elasmobranchs andholocephalans and their numerous fossil relatives, and alsocan be termed sharklike shes or simply sharks.

Chondrocranium:See neurocranium.

Circumglobal:Occurring around the world.

Circumnarial fold :A raised semicircular, lateral ap of skinaround the incurrent aperture of a nostril, in heterodontoids,orectoloboids, and a few batoids, dened by a circumnarialgroove.

Circumnarial groove:A shallow groove dening the lateralbases of the circumnarial folds.

Circumtropical:Occurring around the tropical regions ofthe world.

Clasper claws:In parascylliid orectoloboids, a longitudinalrow of large anterolaterally directed clawlike denticles onthe dorsolateral surface of the clasper glans, supported bythe terminal ventral.

Clasper dactyl:In parascylliid orectoloboids, a large ngerlike process on the medial face of the clasper, supported bythe dorsal terminal and having a mesospur, an analogue to

the lateral spur or spine of the terminal 3 cartilage of otherorectoloboids and other sharks.

Clasper gaff or hook: In the external clasper glans, aposterior hooklike structure, like a clasper spur but formedfrom the dorsal terminal cartilage, found in squaloids of thefamily Squalidae.

Clasper glans:The distal and dorsal part of the externalclasper from the hypopyle to its tip, and including variousmovable terminal structures; also, the same area of theclasper skeleton.

Clasper groove: The longitudinal groove through theclasper, surrounded by the axial and marginal cartilages,and connecting the apopyle and hypopyle.



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Clasper hooks:In the clasper glans of some carcharhinoidsharks, small clawlike dermal denticles arranged in a rowalong the ventral surface of the free edge of the exorhipidion.

Clasper sacs: Dermal sacs with longitudinally ribbedwalls on the ventral and medial surfaces of the claspers ofhexanchoids.

Clasper shaft:That part of the clasper skeleton from itsorigin on the pelvic n basipterygium to the hypopyle;also, that part of the external clasper from its base to thehypopyle.

Clasper spine:In the external clasper, a projection of theterminal 3 cartilage on the lateral surface of the clasperglans, which forms a short to long, acutely pointed, spinethat is covered with shiny hard tissue, possibly enameloid,dentine or both. In some squaloids other terminal cartilagesmay have spines.

Clasper spur: In the external clasper, a projection of the

terminal 3 cartilage on the lateral surface of the clasperglans, which may be pointed but is not covered with shinyhard tissue.

Clasper tip:The posterior end of a clasper.

Claspers: The paired copulatory organs present onthe pelvic ns of male cartilaginous shes, for internalfertilization of eggs, also termed mixopterygia.

Classication:The ordering of organisms into groups onthe basis of their relationships, which may be by similarityor common ancestry.

Cloaca: The common chamber at the rear of the bodycavity of elasmobranchs through which body wastes andreproductive products including sperm, eggs, and youngpass, to be expelled to the outside through a commonopening or vent.

Cover rhipidion: On the external clasper glans, anelongated, longitudinal blade or ap on its dorsomedialexternal edge, often supported by an accessory dorsalmarginal cartilage.

Cranial cavity: The central cavity of the neurocranium,containing the brain, pituitary gland, and roots of the cranial

nerves. It extends posteriorly between the orbits and oticcapsules to the foramen magnum.

Cranial roof :The anterior roof of the cranial cavity of theneurocranium, a dorsomedial, arched or attened plateextending from the anterior fontanelle and between theorbits to the parietal fossa of the otic capsule. Sometimesperforated by a frontal or parietal foramen or fenestra,which may be continuous with the anterior fontanelle andcan occupy most of the cranial roof.

Craniomandibular muscles: Paired head muscles inheterodontoid sharks that originate from long tendons

on the medial walls of the orbits that extend below andtransverse to the levator palatoquadrati and spiracularconstrictor muscles and behind the spiracles to insert onthe posterodorsolateral face of the Meckels cartilages.

They are found in no other sharks and may serve to retractor elevate the jaws.

Crown: The distal part of the oral tooth, almost entirelycovered with shiny enameloid except for the neck. Indenticles, a at dorsal platelike or thornlike structure,elevated above the denticle base on a stalk or pedicle or

conuent with the base.

Crown foot: The expanded, proximal, basal part of thecrown, often bearing cusplets or blades.

Cusp:A usually pointed large distal projection of the crown.A primary cuspis situated on the midline of the crown foot.Multicuspidrefers to oral teeth or denticles with more thanone cusp. In lateral trunk denticles, the posterior ends of thecrown may have medialand lateral cusps, sharp or bluntprojections associated with the medial and lateral ridges.

Cusplet: As with a cusp, but a small projection in associationwith a cusp, and usually mesial and distal but not medial on

the crown foot.

Cutting edge: In oral teeth, the compressed sharplongitudinal ridge on the mesodistal edges of the crown.

Dentine:The primary material of shark oral teeth, a hardtissue with numerous vascular and nonvascular canals.

Dermal denticle orplacoid scale:A small toothlike scalefound in cartilaginous shes, covered with enameloid, witha core and base of dentine and usually small and oftencloseset to one another and covering the body. A few nonbatoid sharks, many batoids, and chimaeroids generally

have them enlarged and sparse or reduced in numbers.

Dermal lobes: In wobbegongs, family Orectolobidae,narrow or broadbased, simple or branched projections ofskin along the horizontal head rim and on the chin.

Diagonal calcications: In a vertebral centrum in crosssection, platelike (diagonal calcied lamellae) or knoblike (diagonal calcied lobes) structures of calciedcartilage that partially lls the uncalcied basalia. Thesehave a radial orientation from the centre of the centrum.

Diphycercal: A caudal n with the vertebral axis runninghorizontally into the n base, which is not elevated.

Diplospondylous vertebrae: Vertebrae of the tail withtwo centra and two basidorsal and basiventral elementsper segment, and mostly with a haemal arch formed bythe basiventral and interventral elements. These includediplospondylous precaudal vertebrae between themonospondylous vertebrae and the base of the caudal n,and diplospondylous caudal vertebrae in the caudal n.

Distal:In any direction, at the far end of a structure. In oralteeth, used in a special sense for structures on the teethtowards the posterolateral mouth corners or rictuses. Seeapicaland basal.

Dorsal:Upwards, in the vertical direction of the back. Seeventral.



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Dorsal n:A n located on the trunk or precaudal tail orboth, and between the head and caudal n. Most sharks

Documents

Deep–sea Cartilaginous Fishes of the Indian Ocean, Vol. 1 – Sharks