FAO Species Catalogue for Fishery Purposes No. 7ISSN 1020-8682SHARKS, BATOIDS AND CHIMAERAS OF THE NORTH ATLANTICSHARKS, BATOIDS AND CHIMAERAS OF THE NORTH ATLANTIC byDavid A. EbertPacifc Shark Research CenterMoss Landing Marine LaboratoriesMoss Landing, CaliforniaUnited States of AmericaandMatthias F. W. StehmannCHTHYSchthyological Research Laboratory and ConsultantHildesheimer Weg 1322459 Hamburg, GermanyFOOD AND AGRCULTURE ORGANZATON OF THE UNTED NATONSRome, 2013The designations employed and the presentation of material in this information product do not imply the expression of any opinion whatsoever on the part of the Food and Agriculture Organization of the United Nations (FAO) concerning thelegalordevelopmentstatusofanycountry,territory,cityorareaorofits authorities,orconcerningthedelimitationofitsfrontiersorboundaries.The mentionofspecifccompaniesorproductsofmanufacturers,whetherornot these have been patented, does not imply that these have been endorsed or recommended by FAO in preference to others of a similar nature that are not mentioned.The views expressed in this information product are those of the author(s) and do not necessarily refect the views of FAO.SBN 978-92-5-107466-4All rights reserved. FAO encourages reproduction and dissemination of material inthisinformationproduct.Non-commercialuseswillbeauthorizedfreeof charge, upon request. Reproduction for resale or other commercial purposes, including educational purposes, may incur fees. Applications for permission to reproduce or disseminate FAO copyright materials, and all queries concerning rights and licences, should be addressed by e-mail to copyright@fao.org or to the Chief, Publishing Policy and Support Branch, Offce of Knowledge Exchange, Research and Extension, FAO, Viale delle Terme di Caracalla, 00153 Rome, taly. FAO 2013PREPARATION OF THIS DOCUMENTThisdocumentwaspreparedunderthegeneralsupervisionoftheFishFinderProgrammeoftheMarineResources Service, Fisheries Resources and Environment Division, Fisheries Department, Food and Agriculture Organization of the United Nations (FAO), Rome, taly. ts production has been supported by a generous Trust Fund (GCP/NT/105/EC) from the European Union. Thiscataloguerepresentsacomprehensiveandupdatedtreatmentoftheidentifcation,taxonomy,distribution, biologyandecologyofthesharks,rays,skatesandchimaerasoftheNorthAtlantic,FAOAreas21and27.nboth areas,cartilaginousfshesarecaughtbybothtargetandnon-targetfsheriesandaresubjecttoanextremelyhigh fishingpressure.Mostspeciesarevulnerabletooverfishingbecauseoftheirspecificbiologicalcharacteristicssuchasslow growth, late maturity, low reproductive potential and thus low capacity for population recovery. Such characteristics limittheircapacitytorecoverfromoverfishinganditisthusessentialtoimplementaneffectiveframeworkforthe managementoffisheriesonsharks,batoidsandchimaeras.ntheNorthAtlantictheEuropeanCommunity,United States of America and Canada have adopted Plans of Action for the Conservation and Management of Sharks that set downmeasuresforthesustainablemanagementofthefisheriesconcerned.Theplansincluderecommendationsto improve species-specific monitoring of catch and landings of cartilaginous fishes. Towardsthisdirection,thiscatalogueisaimedatfacilitatingthespeciesspecifcidentifcationofcartilaginousfshes occurring in the North Atlantic by scientists, fshery offcers and observers and the interested public.Programme manager: Johanne Fischer (FAO, Rome)Editorial assistance: Edoardo Mostarda (FAO, Rome) Scientic iIIustrator: Emanuela D'Antoni (FAO, Rome)Digitization of distribution maps: Fabio Carocci (FAO, Rome)Cover iIIustration: Emanuela D'Antoni (FAO, Rome)D.A. Ebert and M.F.W. Stehmann. 2013.Sharks, batoids, and chimaeras of the North Atlantic FAO Species Catalogue for Fishery Purposes. No. 7. Rome, FAO. 523 pp.ABSTRACTThis volume is a comprehensive, fully illustrated Catalogue of the Sharks, Batoid Fishes, and Chimaeras of the North Atlantic, encompassing FAO Fishing Areas 21 and 27. The present volume includes 11 orders, 32 families, 66 genera, and 148 species of cartilaginous fshes occurring in the North Atlantic. The Catalogue includes a section on standard measurements for a shark, batoid, and chimaera, with associated terms. t provides accounts for all orders, families, and genera and all keys to taxa are fully illustrated. nformation under each species account includes: valid modern names and original citation of the species; synonyms; the English, French, and Spanish FAO names for the species; a lateral view for sharks and chimaeras, dorsal and often also ventral view for batoids, and often other useful illustrations; feld marks; diagnostic features; distribution, including a GS map; habitat; biology; size; interest to fsheries and human impact; local names when available; a remarks sections; and literature. The volume is fully indexed and also includes sections onterminologyandmeasurementsincludinganextensiveglossary,alistofspeciesbyFAOStatistical Areas, and a dedicated bibliography.Sharks, Batoids and Chimaeras of the North Atlantic iiiACKNOWLEDGEMENTSAvolumeofthismagnitudeandscopecouldnotbeaccomplishedwithoutthegeneroushelpofothersandwewishto thank all of those who have been extremely helpful and generous with their time in responding to our numerous questions, providing data and information from their own research (some of it unpublished), and providing much needed literature. The species accounts were improved immensely from the contribution of colleagues and friends. We wish to apologize beforehand if we have forgotten anybody, which will inevitably happen with a work of this magnitude, but we do thank you all for your help and assistance. We would like to extend our thanks to the following individuals for general discussions and information on various aspects ofthisproject:Neil Aschliman(St. AmbroseUniversity); ArcadyBalushkin(ZN,St.Petersburg,Russia);vyBaremore, JohnCarlson,andEnricCortes(NationalMarineFisheriesServiceSoutheastFisheriesScienceCenter,Panama City,Florida);Odd AkselBergstad(nstituteofMarineResearch,His,Norway);RuiCoehlo(CentreofMarineScience, Universityof Algarve,Faro,Portugal);Charles"ChipCotton,KristeneParsons,and TraceySutton(Virginianstituteof Marine Science, College of William and Mary, USA); Jim Ellis (Cefas, United Kingdom); William D. Eschmeyer, (California Academy of Sciences, San Francisco); John D.M. Gordon (Scotland, UK); Karsten Hartel (Museum of Comparative Zoology HarvardUniversity,USA); AaronHenderson(SultanQaboosUniversity,SultanateofOman);Samuelglsias(MNHN-Station Biologie Marine de Concarneau, France); Graham Johnston (Marine nstitute, Galway, reland); Gunnar Jnsson (Reykjavik, celand); Dave Kulka and Steve Campana (Department of Fisheries and Oceans, Canada); Andrea Marshall (Foundation for the Protection of Marine Megafauna, Tofo Beach, nhambane, Mozambique); Lisa Natanson, Tobey Curtis, and John Galbraith (U.S. National Marine Fisheries Service, Northeast Fisheries Science Center); Gavin Naylor (College ofCharleston,Charleston,SouthCarolina,USA);AlexeiOrlov(VNROMoscow,Russia);SteveW.Ross(Centerfor Marine Science, Wilmington, NC, USA); Bernard Sret (RD-MNHN Paris, France); Greg Skomal (Massachusetts Marine Fisheries); and William White (CSRO Marine Laboratories, Hobart, Tasmania, Australia). AspecialthankstoDominiqueDidier(MillersvilleCollege)foruseofherunpublisheddataandinformationonthe Chimaeriformes,JoanParkerandthelibrarystaff(MossLandingMarineLaboratories,MossLanding,California,USA) fortrackingdownthosehardtofnd,usuallyobscure,references,andtoKelley Andrews,JennyBigman,PaulClerkin, and James Knuckey (Pacifc Shark Research Center, Moss Landing Marine Laboratories, Moss Landing, California, USA) who were of invaluable assistance and help in compiling the bibliography for this volume. Thanks to Peter Psomadakis for reviewing such a huge bibliography.David Ebert would like to especially thank Greg Cailliet (Pacifc Shark Research Center, Moss Landing Marine Laboratories, Moss Landing, California, USA) for his mentorship early in my career, and for his continued encouragement and support throughout my career, and to Kaaren Johansen for her patience and support while was writing this volume and fnally to my parents, Earl and Margaret (Peggy) Ebert, for their support and encouragement throughout my life.The David and Lucile Packard Foundation and Moss Landing Marine Laboratories provided funding and other support for this project.FAO Species Catalogue for Fishery Purposes No. 7 iv1. INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.1 Plan of the Catalogue. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31.2 Technical Terms and Measurements: Sharks, Batoids, and Chimaeras . . . . . . . . . . . . . . . . . . . . . . . . 61.2.1 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 Picture Guide to External Terminology and Measurements Used for Batoids. . . . . . . . . . . . . 131.2.5 Picture Guide to External Terminology and Measurements Used for Chimaeras. . . . . . . . . . . 141.2.6 Glossary of Technical Terms. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161.2.7 The egg capsules of skates (Arhynchobatidae and Rajidae) . . . . . . . . . . . . . . . . . . . . . . . . 332. SYSTEMATIC CATALOGUE - SubcIass NEOSELACHII - Cohort SELACHII . . . . . . . . . . . . . . 352.1 Order HEXANCHFORMES Frilled and Cow Sharks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 352.1.1 Family CHLAMYDOSELACHDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36Chlamydoselachus Garman, 1884. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37Chlamydoselachus anguineus Garman, 1884. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382.1.2 Family HEXANCHDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40Heptranchias Rafnesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41Heptranchias perlo (Bonnaterre, 1788) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .42Hexanchus Rafnesque, 1810. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .44Hexanchus griseus (Bonnaterre, 1788). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .45Hexanchus nakamurai Teng, 1962. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 482.2 Order SQUALFORMES Dogfsh sharks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 502.2.1 Family ECHNORHNDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .52Echinorhinus Blainville, 1816 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53Echinorhinus brucus (Bonnaterre, 1788) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .542.2.2 Family SQUALDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56Squalus Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57Squalus acanthias Linnaeus, 1758. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .59Squalus blainville (Risso, 1827) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62Squalus megalops (Macleay, 1881) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 642.2.3 Family CENTROPHORDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67Centrophorus Mller and Henle, 1837. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69Centrophorus granulosus (Bloch and Schneider, 1801) . . . . . . . . . . . . . . . . . . . . . . . . . . 70Centrophorus lusitanicus Bocage and Capello, 1864. . . . . . . . . . . . . . . . . . . . . . . . . . . 73Centrophorus niaukang Teng, 1959. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74Centrophorus squamosus (Bonnaterre, 1788) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76Deania Jordan and Snyder, 1902. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78Deania calcea (Lowe, 1839) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79Deania hystricosa (Garman, 1906) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81Deania profundorum (Smith and Radcliffe, 1912) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 832.2.4 Family ETMOPTERDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84Centroscyllium Mller and Henle, 1841 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86Table of ContentsSharks, Batoids and Chimaeras of the North Atlantic vCentroscyllium fabricii (Reinhardt, 1825). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87Etmopterus Rafnesque, 1810. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89Etmopterus gracilispinis Krefft, 1968. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91Etmopterus hillianus (Poey, 1861). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93Etmopterus princeps Collett, 1904 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .94Etmopterus pusillus (Lowe, 1839). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96Etmopterus spinax (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .992.2.5 Family SOMNOSDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101Centroscymnus Bocage and Capello, 1864 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103Centroscymnus coelolepis Bocage and Capello, 1864 . . . . . . . . . . . . . . . . . . . . . . . . . . 104Centroscymnus owstoni Garman, 1906. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106Centroselachus Garman, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107Centroselachus crepidater (Bocage and Capello, 1864) . . . . . . . . . . . . . . . . . . . . . . . . 108Scymnodalatias Garrick, 1956 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109Scymnodalatias garricki Kukuev and Konovalenko, 1988 . . . . . . . . . . . . . . . . . . . . . . . 110Scymnodon Bocage and Capello, 1864. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111Scymnodon ringens Bocage and Capello, 1864 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112Somniosus Lesueur, 1818 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113Somniosus microcephalus (Bloch and Schneider, 1801). . . . . . . . . . . . . . . . . . . . . . . . 114Somniosus rostratus (Risso, 1827) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117Zameus Jordan and Fowler, 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118Zameus squamulosus (Gnther, 1877) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1192.2.6 Family OXYNOTDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120Oxynotus Rafnesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122Oxynotus centrina (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122Oxynotus paradoxus Frade, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1242.2.7 Family DALATDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126Dalatias Rafnesque, 1810. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128Dalatias licha (Bonnaterre, 1788). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129Isistius Gill, 1865 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131Isistius plutodus Garrick and Springer, 1964 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131Squaliolus Smith and Radcliffe, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133Squaliolus laticaudus Smith and Radcliffe, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1342.3 Order SQUATNFORMES Angel sharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1372.3.1 Family SQUATNDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137Squatina Dumeril, 1806 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138Squatina dumeril Lesueur, 1818 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139Squatina squatina (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1412.4 Order ORECTOLOBFORMES. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1442.4.1 Family GNGLYMOSTOMATDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145Ginglymostoma Mller and Henle, 1837. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146Ginglymostoma cirratum (Bonnaterre, 1788). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1472.4.2 Family RHNCODONTDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149Rhincodon Smith, 1829 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150Rhincodon typus Smith, 1828. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150FAO Species Catalogue for Fishery Purposes No. 7 vi2.5 Order LAMNFORMES Mackerel sharks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1532.5.1 Family ODONTASPDDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155Carcharias Rafnesque, 1810. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156Carcharias taurus Rafnesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157Odontaspis Agassiz, 1838. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159Odontaspis ferox (Risso, 1810). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1602.5.2 Family MTSUKURNDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162Mitsukurina Jordan, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162Mitsukurina owstoni Jordan, 1898. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1632.5.3 Family ALOPDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164Alopias Rafnesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166Alopias superciliosus Lowe, 1841. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167Alopias vulpinus (Bonnaterre, 1788) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1682.5.4 Family CETORHNDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170Cetorhinus Blainville, 1816. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171Cetorhinus maximus (Gunnerus, 1765). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1712.5.5 Family LAMNDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174Carcharodon Smith, 1838. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175Carcharodon carcharias (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176Isurus Rafnesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178Isurus oxyrinchus Rafnesque, 1810. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179Isurus paucus Guitart, 1966. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182Lamna Cuvier, 1816. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183Lamna nasus (Bonnaterre, 1788) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1842.6 Order CARCHARHNFORMES Ground Sharks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1872.6.1 Family SCYLORHNDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188Apristurus Garman, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190Apristurus aphyodes Nakaya and Stehmann, 1998 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191Apristurus laurussonii (Saemundsson, 1922). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193Apristurus manis (Springer, 1979). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195Apristurus melanoasper Iglesias, Nakaya, and Stehmann, 2004. . . . . . . . . . . . . . . . . . . 196Apristurus microps (Gilchrist, 1922) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198Apristurus profundorum (Goode and Bean, 1896). . . . . . . . . . . . . . . . . . . . . . . . . . . . 199Galeus Rafnesque, 1810a. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201Galeus atlanticus (Vaillant, 1888). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202Galeus melastomus Rafnesque, 1810a. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203Galeus murinus (Collett, 1904). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205Scyliorhinus Blainville, 1816. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206Scyliorhinus canicula (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207Scyliorhinus retifer (Garman, 1881). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209Scyliorhinus stellaris (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2112.6.2 Family PSEUDOTRAKDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212Pseudotriakis Capello, 1868 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213Pseudotriakis microdon Capello, 1868. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2132.6.3 Family TRAKDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215Galeorhinus Blainville, 1816. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216Sharks, Batoids and Chimaeras of the North Atlantic viiGaleorhinus galeus (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217Mustelus Linck, 1790. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219Mustelus asterias Cloquet, 1821. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220Mustelus canis (Mitchill, 1815) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222Mustelus mustelus (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2252.6.4 Family CARCHARHNDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227Carcharhinus Blainville, 1816 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229Carcharhinus acronotus (Poey, 1860) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232Carcharhinus altimus (Springer, 1950). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233Carcharhinus brevipinna (Mller and Henle, 1839) . . . . . . . . . . . . . . . . . . . . . . . . . . . 235Carcharhinus falciformis (Mller and Henle, 1839) . . . . . . . . . . . . . . . . . . . . . . . . . . . 238Carcharhinus isodon (Mller and Henle, 1839) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240Carcharhinus leucas (Mller and Henle, 1839) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 242Carcharhinus limbatus (Mller and Henle, 1839). . . . . . . . . . . . . . . . . . . . . . . . . . . . 245Carcharhinus longimanus (Poey, 1861) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247Carcharhinus obscurus (Lesueur, 1818). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249Carcharhinus plumbeus (Nardo, 1827). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251Carcharhinus signatus (Poey, 1868) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254Galeocerdo Mller and Henle, 1837. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256Galeocerdo cuvier (Pron and Lesueur, 1822) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256Negaprion Whitley, 1940 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 259Negaprion brevirostris (Poey, 1868) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 259Prionace Cantor, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261Prionace glauca (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262Rhizoprionodon Whitley, 1929. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265Rhizoprionodon terraenovae (Richardson, 1836) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2662.6.5 Family SPHYRNDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 268Sphyrna Rafnesque, 1810a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269Sphyrna lewini (GriIfth and Smith, 1834). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 270Sphyrna mokarran (Rppell, 1837). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272Sphyrna tiburo (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275Sphyrna zygaena (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2773. SYSTEMATIC CATALOGUE - SubcIass NEOSELACHII - Cohort BATOIDEA. . . . . . . . . . . . 2793.1 Order TORPEDNFORMES Electric Rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2793.1.1 Family TORPEDNDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279Torpedo Houttuyn, 1764. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280Torpedo (Torpedo) marmorata Risso, 1810. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281Torpedo (Tetronarce) nobiliana Bonaparte, 1835 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283Torpedo (Torpedo) torpedo (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2843.2 Order PRSTFORMES Sawfshes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2863.2.1 Family PRSTDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 286Pristis Linck, 1790 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287Pristis pectinata Latham, 1794. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287Pristis pristis (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2893.3 Order RAJFORMES Skates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2913.3.1 Family RHNOBATDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 292Rhinobatos Linck, 1790 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293Rhinobatos (Glaucostegus) cemiculus Geoffroy Saint-Hilaire, 1817. . . . . . . . . . . . . . . . 294Rhinobatos (Rhinobatos) rhinobatos (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . 295FAO Species Catalogue for Fishery Purposes No. 7 viii3.3.2 Family ARHYNCHOBATDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297Bathyraja Ishiyama, 1958 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 298Bathyraja pallida (Forster, 1967) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299Bathyraja richardsoni (Garrick, 1961). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301Bathyraja spinicauda (Jensen, 1914). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3033.3.3 Family RAJDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305Amblyraja Malm, 1877. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308Amblyraja hyperborea (Collett, 1879) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 309Amblyraja jenseni (Bigelow and Schroeder, 1950) . . . . . . . . . . . . . . . . . . . . . . . . . . . 311Amblyraja radiata (Donovan, 1808) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313Dipturus Rafnesque, 1810. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 315Dipturus sp. cf. ossada (Risso, 1826). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 317Dipturus sp. cf. intermedia (Parnell, 1837) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 319Dipturus laevis (Mitchill, 1818) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 321Dipturus nidarosiensis (Storm, 1881). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323Dipturus oxyrinchus (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325Dipturus sp. Stehmann (in prep.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 328Leucoraja Malm, 1877. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329Leucoraja circularis (Couch, 1838). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 331Leucoraja erinacea (Mitchill, 1825) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 333Leucoraja fullonica (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 335Leucoraja garmani (Whitley, 1939). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337Leucoraja naevus (Mller and Henle, 1841) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 339Leucoraja ocellata (Mitchill, 1815). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 340Malacoraja Stehmann, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 343Malacoraja kreffti (Stehmann, 1977). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 344Malacoraja senta (Garman, 1885). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 346Malacoraja spinacidermis (Barnard, 1923). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347Neoraja McEachran and Compagno, 1982 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349Neoraja caerulea (Stehmann, 1976) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 351Neoraja iberica Stehmann, Seret, Costa and Baro, 2008. . . . . . . . . . . . . . . . . . . . . . . . 353Neoraja sp. sensu Stehmann et al. (2008). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 356Raja Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357Raja brachyura Lafont, 1873. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 360Raja clavata Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 362Raja eglanteria Bosc, 1800 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 364Raja maderensis Lowe, 1838. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 366Raja microocellata Montagu, 1818 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 368Raja miraletus Linnaeus, 1758. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 370Raja montagui Fowler, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372Raja undulata Lacepede, 1802. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 373Rajella Stehmann, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 375Rajella bathyphila (Holt and Byrne, 1908) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 378Rajella bigelowi (Stehmann, 1978) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 380Rajella dissimilis (Hulley, 1970). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382Rajella fyllae (Ltken, 1888) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384Rajella kukujevi (Dolganov, 1985) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386Rajella lintea (Fries, 1839) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 388Rostroraja Hulley, 1972 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 390Rostroraja alba (Lacepede, 1803). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3903.4 Order MYLOBATFORMES Stingrays. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3933.4.1 Family DASYATDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 394Dasyatis Rafnesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 395Dasyatis centroura (Mitchill, 1815). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 396Sharks, Batoids and Chimaeras of the North Atlantic ixDasyatis pastinaca (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 397Pteroplatytrygon Fowler, 1910. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 398Pteroplatytrygon violacea (Bonaparte, 1832). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3993.4.2 Family GYMNURDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 400Gymnura van Hasselt, 1823. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 401Gymnura altavela (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4013.4.3 Family MYLOBATDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403Myliobatis Cuvier, 1817 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 404Myliobatis aquila (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 405Pteromylaeus Garman, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 406Pteromylaeus bovinus (Geoffroy Saint-Hilaire, 1817) . . . . . . . . . . . . . . . . . . . . . . . . . . 407Rhinoptera Cuvier, 1829. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 408Rhinoptera bonasus (Mitchill, 1815) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409Rhinoptera marginata (Geoffroy Saint-Hilaire, 1817) . . . . . . . . . . . . . . . . . . . . . . . . . . 411Manta Bancroft, 1828. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 413Manta birostris (Walbaum, 1792) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 413Mobula Rafnesque, 1810 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 415Mobula mobular (Bonnaterre, 1788) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4154. SYSTEMATIC CATALOGUE - SubcIass HOLOCEPHALI . . . . . . . . . . . . . . . . . . . . . . . . . 4174.1 Order CHMAERFORMES Ghost sharks, Silver sharks, Ratfsh, Chimaeras . . . . . . . . . . . . . . . . . 4174.1.1 Family CHMAERDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 418Chimaera Linnaeus, 1758. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 419Chimaera monstrosa Linnaeus, 1758. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 420Hydrolagus Gill, 1862 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 421Hvdrolagus afhnis (de Brito Capello, 1868) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 422Hydrolagus lusitanicus Moura, Figueiredo, Bordalo-Machado, Almeida & Gordo, 2005 . . . . . 424Hydrolagus mirabilis (Collett, 1904). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 426Hydrolagus pallidus Hardy and Stehmann, 1990 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4274.1.2 Family RHNOCHMAERDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 429Harriotta Goode and Bean, 1895 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 430Harriotta haeckeli Karrer, 1972 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 431Harriotta raleighana Goode and Bean, 1895. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 432Rhinochimaera Garman, 1901 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 433Rhinochimaera atlantica Holt and Byrne, 1909 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4345. BIBLIOGRAPHY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4376. INDEX OF SCIENTIFIC AND VERNACULAR NAMES . . . . . . . . . . . . . . . . . . . . . . . . . . 493FAO Species Catalogue for Fishery Purposes No. 7 xSharks,andtheirrelativesthebatoidsandchimaeras,collectivelycomprisingtheClassChondrichthyesor cartilaginousfshesareamongonethemostsuccessful groupsoffshes(Compagno,1973;Nelson,2006)having penetratedmostmarineecosystems.Thisincludes continental and insular shelf waters from the intertidal out to a depth of 200 m, and into the deepsea as well as oceanic andpelagicforms.nsomeareas,somespecieshave even penetrated into and occupy estuarine and freshwater riverecosystems.Recentreviewsonthebiodiversityand distributionofchondrichthyansinvariousecosystemshas shownthattropicalmarineecosystemshavethehighest diversity, followed by deepsea and temperate ecosystems, and with the epipelagic and freshwater ecosystems having a lower diversity of species (Ebert and Winton, 2010; Kyne and Simpfendorfer, 2010; Rosa et al. 2010; Stevens, 2010; White and Sommerville, 2010). Worldwide there are at least 1200 known living and valid species of sharks, batoids, and chimaeroidscomprising13orders,58to60families,and 192genera,withadditionalspeciesrequiringdescription (Nelson,2006;EbertandWinton,2010;D.A.Ebert,pers. database,12October2011).Theseincludeover500 speciesofsharks,over650speciesofbatoids,orfat sharks,(ordersTorpediniformes,Pristiformes,Rajiformes, andMyliobatiformes),andatleast50speciesofghost sharks,silversharks,elephantfsh,chimaerasorratfsh (order Chimaeriformes). Thiscataloguecoversallthedescribedspeciesofliving sharks,batoids,andchimaerasfoundintheNorth Atlantic, FAO Areas 21 and 27 (Fig. 1). The catalogue is intended to be a comprehensive review of the shark-like fshes of the North Atlantic in a form accessible to fsheries workers as well as researchersonsharksystematics,biodiversity,distribution, and general biology. t also caters to other researchers that need comparative information on sharks, and their relatives, andtopeoplewhoencountersharksduringthecourseof workorplayinthesea,orinfreshwater,andthegeneral public.tincludesspeciesofmajor,moderate,minor,and minimal importance to fsheries as well as those of doubtful or potential use to fsheries. t also covers those species that havearesearch,recreational,educational,andaesthetic importance,aswellasthosespeciesthatoccasionallybite and threaten people in the water and the far more numerous speciesthatare'bitten'andthreatenedbypeoplethrough exploitation and habitat modifcation. Thespeciesspecifcinformationonthebiology, conservationstatus,distribution,habitat,fsheries,and systematicsofNorth AtlanticOceanchondrichthyanswas compiled from primary literature sources including, but not limitedto,BigelowandSchroeder(1948,1953),Hureau andMonod(eds.,1973),Whiteheadetal.(1984),Scott and Scott (1988), Branstetter, Burgess, and McEachran in ColletteandKlein-MacPhee(2002),Mooreetal.(2003), andHarteletal.(2008).RegionalFAOcataloguesand identifcationguidesfortheEasternCentral Atlantic(FAO Area34)andWesternCentral Atlantic(FAO Area31),as wellastheMediterraneanandBlackSea(FAOArea37) werealsohelpfulinroundingthespecieschecklistsfrom these adjacent areas. Several grey literature and electronic sourceswerealsoofinvaluablehelp,theseincluded,but werenotlimitedto,theCaliforniaAcademyofSciences CatalogueofFishes(http://www.calacademy.org/research/ichthyology/catalog/fshcatsearch.html);EuropeanUnion (EU),FAO(http://www.fao.org/f/default.asp),CES WorkingGroupofElasmobranchFishes(www.ices.dk), UCNSharkSpecialistGroup(http://www.iucnredlist.org), NOAAFisheries,andMar-Ecoreports.Acomprehensive bibliographyoftheliterature,includingprimary,grey,and electronic sources is provided at the end of this volume. BiogeographyofRegion.TheNorthAtlanticregion includestwomajorFAOFishing Areas,theEasternNorth Atlantic(Area27)andtheWesternNorthAtlantic(Area 21). TheEasternNorth Atlanticregionextendsessentially fromtheNorthPoletoabout36Ninthecentral Atlantic, andisboundedfromtheeastcoastofGreenlandand alongthe40Wmeridiansouthwardtowhereitintersects at 36N and to the east Area 27 is bounded in the eastern Barents Sea at about 68 30'E and by continental western Europe.TheWesternNorth Atlanticextendsfromeastern GreenlandwestwardstotheArcticwatersofnorthern Canada at 120W, and from northern Arctic waters at about 78 10'N southwards to about Cape Hatteras at 35N. The North Atlantic encompasses no less than 14 Large Marine Ecosystems,nineintheEasternNorth Atlanticandfvein the Western North Atlantic. This includes the East Greenland Shelf,NorwegianSea,BarentsSea,celandShelf,Faroe Plateau,NorthSea,BalticSea,CelticBiscayShelf,and berianCoastallargemarineecosystemsintheEastern North Atlantic (Area 27), and in the Western North Atlantic (Area 21) this includes the Northeast US Continental Shelf, Scotian Shelf, Newfoundland-Labrador Shelf, Hudson Bay, andWestGreenlandShelflargemarineecosystems(Fig. 1 Map of the North Atlantic FAO Areas 21 and 27). TheNorthAtlanticregionisessentiallyacoldtemperate regimethatextendsfromtheArcticCirclesouthwardsto warmertemperateboundariesatabout36N(Area27) and35N(Area21)latitude.ntheBorealmid-Atlantic theeastfowingGulfStreambecomestheNorthAtlantic Current, which forms the upper portion of the North Atlantic Gyre,aclockwiserotatingoceangyre.ThemainNorth Atlantic Current fows east towards the British sles where it branches into two major divisions, with the North Atlantic current continuing southward and a separate major branch fowingnorthwardstobecometheNorwegianCurrent (Briggs,1995).TheNorwegianCurrentbranchcarries relatively warm, saline water of Gulf Stream origin into the seas off the western coast of Europe (Briggs, 1995) and as a result, the border of the warm and cold-temperate regions occursfarthernorththaninthewesternNorth Atlantic.n the Western North Atlantic, the Florida Current becomes the Gulf Stream at about Cape Hatteras (35N), North Carolina, USA, and continues fowing northwards, off the continental shelf and over deeper water, but veers back into shallower waterovertheGrandBanks,southeastofNewfoundland. ThecoldLabradorCurrentfowssouthwardbetweenthe GulfStreamandthecoast,extendingcold-temperate 1. INTRODUCTIONSharks, Batoids and Chimaeras of the North Atlantic 1conditions farther south than they occur in the eastern North Atlantic (Briggs, 1995). As a result, the environment along thewesternNorthAmericancoastlinetransitionsrapidly from warm-temperate to polar regimes, completing the shift within 20 of latitude (Barnes and Hughes, 1982).CIassicationandsystematicarrangementusedhere. The higher classifcation of these fshes includes the class Chondrichthyesthatisdividedintotwomajorgroups, eachwithalongandseparate,pre-Devonianhistory, thechimaeroids,Holocephali(withasinglelivingorder Chimaeriformes),andthesharksandbatoidsproper,with thesurvivinggroupNeoselachiidividedintotwocohorts, theSelachii(sharks)andtheBatoidea(raysandskates) and includes all of the modern living species. The Selachii isfurtherdividedintotwosuperorders,theSqualomorphii and Galeomorphii. The superorder Squalomorphii includes the orders Hexanchiformes, Squaliformes, Squatiniformes, and Pristiophoriformes, while the superorder Galeomorphii includestheHeterodontiformes,Orectolobiformes, Lamniformes, and Carcharhiniformes. The cohort Batoidea recognizesfourorders,Torpediniformes,Pristiformes, Rajiformes, and Myliobatiformes. The ordinal classifcation oftheshark-likefsheslargelyfollowsthearrangement ofCompagno(2001,2005),EbertandCompagno(n press), and Ebert (n preparation) with some modifcations inrecognizingeightordersand34families.Thehigher classifcation of batoid fshes and the assignment of various familiestosubordersorordersarenotyetfullyclarifed. Nelson (2006) provided a table refecting different concepts andcomparinghisownclassifcationoftheprevious3rd edition (Nelson, 1994) with those of Compagno (1999) with sixordersand21families,byMcEachranand Aschliman (2004) with four orders and 14 families, and fnally his own concept of the 4th edition (Nelson, 2006) with four orders and 17 families. Three more families, as compared to McEachran and Aschliman (2004), in the latter concept concern a new familyPotamotrygonidaeofSouthAmericanfreshwater stingraysandthepreservationofthefamiliesRhinidae andRhynchobatidae.Theclassifcationusedherefollows Nelson(2006)withtheexceptionofaddingan18thfamily, Arhynchobatidae (Fowler 1934), resurrected by Compagno (1999,2005),butwithoutexplanation.Recently,Nayloret al. (2012) and Aschliman et al. (2012) revisited the validity ofthisfamilyusingclaspermorphologyandmolecular analysisandconcludedthatthisfamilyisvalid,andisa sister group to the family Rajidae. The higher classifcation ofchimaerasfollowsDidier(1995,2004)andDidier, Kemper,andEbert(2012).Therelationshipofthesharks to the batoids is still unresolved, with recent classifcations suggesting that the batoids are either sister to the modern sharkordersPristiophoriformesandSquatiniformes,and shareacommonancestrywiththeSqualiformes(e.g.the Hypnosqualean hypothesis), or follow traditional dichotomy of all modern sharks and batoids. The two hypothesis breaks down largely between traditional morphologist (Compagno, 1973,1977,1999,2001,2005;Shirai,1992a,1996;de Carvalho, 1996) and newer molecular evidence (Douady et al., 2003; Naylor et al., 2005; G. Naylor, pers. comm.). The following classifcation to order is based on the above discussion on higher ordinal classifcations (* starred orders are covered in this volume): Class Chondrichthyes (cartilaginous fshes)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 (dogfsh 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 (sawfshes)*Order Rajiformes (skates and guitarfshes)*Order Myliobatiformes (stingrays)*NorthAtIanticBiodiversity.TheNorthAtlantichas amoderatelydiversechondrichthyanfaunawith11 orders,32families,66genera,andatleast148species beingrepresented(Table1).Themostspeciousgroup ofchondrichthyansintheNorthAtlanticaretheskates (Rajiformes),whichhasatleast41species,represented. ThisisfollowedbytheCarcharhiniformeswith36species and the Squaliformes with 30 species. Of these totals, the EasternNorthAtlantichasahigherdiversityintermsof families (n = 32), genera (n = 63), and number of species (n = 127) in comparison to the Western North Atlantic that has 20 families, 50 genera, and 83 species. The number of batoidsrepresentedintheEasternNorth Atlanticismore than double that of the Western North Atlantic, 49 species versus 23, respectively. North Atlantic (Tot.)Sharks Batoids Chimaeras Tot.Orders 6 4 1 11Families 22 8 2 32Genera 42 20 4 66 Species 84 56 8 148North Atlantic (Area21) Orders 6 4 1 11Families 20 7 2 29Genera 32 14 4 50 Species 54 23 6 83North Atlantic (Area27) Orders 6 4 1 11Families 22 8 2 32Genera 39 20 4 63 Species 70 49 8 127Table 1 - The families, genera, and species represented withintheNorth Atlantic(totalfor Areas21and27), Eastern North Atlantic (Area 27), and Western North Atlantic (Area 21).FAO Species Catalogue for Fishery Purposes No. 7 21.1 PIan of the CataIogue TheformatforthiscataloguefollowsthatoftheFAO Catalogue of Sharks of the World (Compagno, 2001; Ebert and Compagno, n press; Ebert, n preparation), orders as thehighesttaxonomicgroupdealtwithhere,followedby family, genus, and species accounts. A key to the families and genera, where appropriate is also included. OrderAccountsincludethevalidmodernformofthe order name with author and year; the original citation of the order name with its author, year, reference and pagination; thenumberofrecognizedfamiliesintheNorthAtlantic; commonorderSynonymsmainlyfromtheNorthAtlantic region with the name, author, year, and pagination; the FAO order VernacuIar Names in English, French and Spanish; Field Marks and Diagnostic Features of members of the order; an account of the natural history of the order under separatesectionscoveringDistribution,Habitatand BioIogy;asectiononInteresttoFisheriesandHuman Impact,asynopsisofthehumanissuesaffectingshark families; Local Names when available; a Remarks section mostlywithsystematiccomments;andaKeytoNorth Atlantic Families, when orders have more than one family.FamiIyAccountsincludethevalidmodernformofthe familynamewithauthorandyear;theoriginalcitation ofthefamilynamewithitsauthor,year,referenceand pagination;thevalidtypegenuswithauthoranddate; thenumberofrecognizedNorthAtlanticgenerainthe family;familySynonymswithnamesmostlyassociated withtheNorthAtlanticregionandwiththename,author, year,andpagination;theFAOfamilyVernacuIarNames inEnglish,FrenchandSpanish;FieldMarksand Diagnostic Features of members of the family; an account of the natural history of the family under separate sections coveringDistribution,HabitatandBioIogy;asectionon InteresttoFisheriesandHumanImpact,asynopsisof thehumanissuesaffectingsharkfamilies;LocalNames when available; a Remarks section mostly with systematic comments; a Literature section covering references to the entirefamily;andaKeytoNorth AtIanticGenera,when families have more than one genus.GenericAccountsincludethevalidmodernformofthe genus name with author and year; the original citation of the genus(orsubgenus),withitsauthor,year,referenceand pagination, and, if a subgenus, the original genus name with author and year that the subgenus was originally placed in; the type species and means of designating it (for example, byoriginaldesignation,monotypy,absolutetautonymy,or subsequentdesignation);thenumberofrecognizedNorth Atlantic species in the genus; the Synonyms of genera, with their rank (genus, subgenus, or other genus-group ranking), author, year, pagination, and genus they were described in if originally ranked as subgenera or equivalents; FAO Names if they exist; Field Marks if genera are large and distinctive; DiagnosticFeaturesofthegenus;LocalNameswhere available; a Key to North AtIantic Species if the genus has more than one species (is not monotypic); and a Remarks section.Species Accounts include the valid modern names of the species,withauthoranddate;theoriginalcitationofthe species,withitsauthor,year,referencepagination;the holotype,syntypes,lectotypeorneotypeofeachspecies (paratypes are not listed in the present account), including thetotallengthandsexofthespecimen,itsinstitutional deposition,anditscataloguenumber;thetypelocality includingthelocation,coordinatesanddepthifavailable, wheretheholotype,syntypes,lectotypeorneotypewere caught;Synonymsofthespecies,includingtheirnames, authorsanddates;asectionlistingotherscientifcnames recentlyinuse;theEnglish,French,andSpanishFAO Names for the species; a lateral view illustration, and often otherusefulillustrations(lateralviewdrawingsaregiven of each shark species, usually ventral views of heads, and oftenteethanddenticlesofthesharkinquestion);Field Marks; Diagnostic Features (except in monotypic genera); Distribution,includingamap;Habitat;BioIogy;Size; InteresttoFisheriesandHumanImpact;LocalNames whenavailable;aRemarkssectionwhennecessary;and Literature.SynonymscommonlyseenintheNorth Atlanticliterature arelisted,whereappropriate,andincludeonlytrue taxonomic synonyms of the valid family, genus and species given. For species, another category, Other Combinations, is provided for common misidentifcations of a given species with another, valid species (for example, the requiem shark Carcharhinusbrachyurus(Carcharhinidae))wasoften termed C. remotus in the older literature, but the latter is a junior synonym of C. acronotus) as well as commonly used combinationsthatplaceavalidspeciesindifferentgenera (for example, Odontaspis taurus or Eugomphodus taurus for Carcharias taurus).FAOFamiIyandSpeciesNames.English,Frenchand Spanishnamesforeachfamilyandspecies,primarilyfor use within FAO, were selected by the following criteria: (a) each name applies to a single family or species worldwide; (b)thenameconformswithFAOspellingnomenclature; (c) the name conforms to prior usage when possible. FAO names are not intended to replace local species names, but arenecessarytoovercometheconfusioncausedbythe use of a single name for more than one species or several names for one species. The FAO names used here conform topriorFAOusage.ThecommonFrenchandSpanish names of species from the other FAO Catalogues, including theSharksofWorld(Compagno,1984,2001;Ebertand Compagno,npress;Ebert,npreparation),andregional FAO Catalogues on the Sharks, Rays, and Chimaeras of the Eastern Central Atlantic (Compagno, 1981a, b; Stehmann, 1981),WesternCentral Atlantic(Compagno,2002;Didier, 2002;McEachran,2002),andMediterraneanandBlack Sea (Serena, 2005), were used when appropriate. Keys,FieIdMarks,andDiagnosticFeatures.These sections include identifcation data in different forms. Keys to families, genera and species are standard dichotomous biological keys that are followed in steps of alternate choices to singleoutthetaxacovered.t shouldbenotedthat the Keys include only those families, genera, and species that occurinFAO Areas21and27,orwithinthescopeofthe presentCatalogueanddoesnotincludethosegroupsnot Sharks, Batoids and Chimaeras of the North Atlantic 3occurring with the area. Field Marks generally include a few obviouscharactersofuseinfeldidentifcation,extracted from Diagnostic Features at various levels, but included in a separate section. Field Marks are listed at the ordinal, familial andspecieslevels,andoccasionallythegenericlevelin cases of large genera with many species. The arrangement of Field Mark characters is semihierarchical and pragmatic and may include characters from a higher level such as an orderinlowerleveltaxonomicaccountssuchasthoseof species.FieldMarksincludecharactersthatareobvious inliveorfresh-caughtindividualsbutmaybeobscurein frozen or preserved material. Diagnostic Features are lists ofcharactersattheordinal,familial,generic,andspecies level, with the character choice generally limited to external characters,particularlyatthespecieslevel,becauseof their primary purpose of identifcation rather than indication ofrelationships.TheDiagnosticFeaturessectionsare hierarchical,withcharactersattheordinallevelgenerally notduplicatedatthefamily,genusandspecieslevels. Monotypic orders with one family (such as Squatiniformes), monotypicfamilieswithonegenus(Mitsukurinidae)or monotypicgenerawithonespecies(Pseudotriakis)all havetheDiagnosticFeaturessectionpresentonlyinthe highest taxon covered.Distribution. Geographic distributions for nearly all species of sharks are given by listing the countries off the coasts of or oceanographic features, e.g. seamounts and troughs, of wherethespeciesoccurs.ncompilingdistributionaldata andpreparingmapsitwasnotedthatthedistributionsof manywide-rangingcoastalanddeepseaspeciesarevery spottily known as present. n many cases gaps in distribution maynotindicateabsenceofagivenspeciesbutabsence ofknowledge.Continentalslopesharkfaunasarepoorly known for much of the world, and a number of deepwater speciesprobablyhavewiderrangesthanarecurrently known.Mucheffortwasmadetoscreenoutdistribution errors, based on misidentifcations of species, at a cost of presenting distributional lists and maps that are spotty, but possibly more accurate. Habitat. Habitat covers information on physical conditions where various sharks are found. The known depth range of the species (in metres), position in the water column, type ofsubstrateoccupied,andpreferencesrelativetocoasts arenotedwhenavailable.nmostcasesdataonsalinity, oxygencontent,andspecifctemperatureofthewaterin which they occur was not available or was not in an easily usable form and has not been regularly compiled here. BioIogy. ncludes data on reproduction, age and growth, diet, and behaviour and movement patterns. Compilation of these datasuggeststhatveryfewspeciesarebiologicallywell known, and even in the picked dogfsh (Squalus acanthias), perhapsthebest-knownoflivingcartilaginousfshes,there are areas of its biology that are very poorly known (such as its behaviour and sociobiology). Size.Allsizedataaregivenastotallengths;thisisthe measurement most often used as an independent variable and standard measurement in the shark literature, although particularlyinfsheriespapersprecaudallengths,fork lengths,andothermeasurementshavebeenusedfrom choice or necessity. Unfortunately shark workers have not agreedonastandardmethodofmeasuringtotallength, sototallengthsfromdifferentsourcesintheliterature maynotbestrictlycomparable.Wepreferandadvocate as a standard method a direct measurement, in which the shark, batoid, or chimaera is held belly down with its dorsal caudal-fnlobedepressedintolinewithitsbodyaxisand total length measured as a point to point distance (not over thecurveofthebody)fromthesnouttiptothetipofthe dorsalcaudal-fnlobe.Thismethodlendsitselfreadilyto quickuseofafshboardwithaperpendicularfrontbaror plate to index the fsh's snout against, a one metre or two metre ruler or folding ruler slipped under the shark, batoid, orchimaeraorevenasteelorclothtape,andavoidsthe trouble of computation and possible errors and loss of data. Totallengthdatapresentedincludesmaximumsize,size at maturity (in some cases, a size range at maturity, when abundantdatawereavailable)andmaximumsizefor bothsexes,andsizeatbirthorhatching.Sometimessize atsexualmaturityforeitherorbothsexesisnotknown, inwhichcasesreportedminimumandmaximumsizesof adultindividualsaregiven.nsomecasesmaximumsize exceeds that recorded for either sex, in which case the sex oftheoutsizedindividualorindividualsrepresentingthe maximumsizemeasurementswasnotindicated.nsome poorly known species only immature individuals are known, in which case the hypothetical maximum adult size is almost certainly larger than the known immature maximum.nsomespecieslength-weightequationsarepresented, usually of the form W = a + TLb, where W is weight, a and b are constants, and TL is total length.InteresttoFisheriesandHumanImpact.Thissection includesFisheriesinformation,includingwhetherthe speciesistakenintargetedornon-targeted(bycatch) fsheriesandiftakenasbycatchwhetheritisretainedor discarded.Dataonlocalitiesoffsheries,gearused,and usesoftheparticularspeciesarenotedwhenavailable. Nationalfsheriesdataforsharksisoftensketchyand combinedforanumberofspecies.Thus,catchstatistics wereavailableforrelativelyfewspeciesofsharksbutare notedwhenavailable,withparticularemphasisondata fromthosespeciesreportedtoFAO.Additionaldataare increasinglyavailablefromnationalandregionalfsheries bodiesarepresentedwhenavailable.Otheraspectsof humaninteraction,e.g.sharkattackorecotourism,are presented,andthecurrentconservationRedListstatus ofeachspeciesasevaluatedbytheUCNSpecies Survival Commission's Shark Specialist Group (http://www.iucnredlist.org). AttheendoftheBibliographysectionan electronic reference section has been added with a link to theRedList Assessmentforeachspeciesincludedinthe Catalogue. LocalNames.Manyspecieshavenovernacularnames whatsoeverorarelumpedundercatchallnames,while some species such as the white and basking sharks have dozens of names. Wherever possible common local names are presented, especially for important wide-ranging sharks. The broadening interest in sharks, batoids, and chimaeras, andurgentneedtoacquirespecies-specifcdatafortheir FAO Species Catalogue for Fishery Purposes No. 7 4management and conservation should encourage fsheries biologistsandotherresearcherstocompilelocalnames fortheirowncountriesorregions,andaddtothesketchy knowledge of local names.Remarks. mportant information, especially on systematics and 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 appropriate forcomparisontothegroupsoccurringwithinFAO Areas 21 and 27.Literature.Referencescitedhereincludespecifcworks withimportantinformationforeachspeciesandfamilyas well as comprehensive accounts, but are not intended as a comprehensive bibliography. Fig. 1Map of FAO Area 21 (Western North Atlantic) and Area 27 (Eastern North Atlantic)Sharks, Batoids and Chimaeras of the North Atlantic 51.2 TechnicaI Terms and Measurements: Sharks, Batoids, and Chimaeras1.2.1 Picture Guide to ExternaI TerminoIogy of SharksFig. 2 Lateral viewFig. 3 Ventral viewFig. 4 Head of an orectoloboid shark (ventral view)Fig. 5 Nostrilincurrent aperture excurrent aperture anterior nasal fapposterior nasal faphead trunkmouthlabial furrowsgill openingspectoral fnpelvic fnanal fnclasper (males)caudal keelprecaudal pitsecond dorsal fnfrst dorsal fndorsalfn spinecaudal fnsnout nostrileyespiracletailtrunkmouthpectoral fnanal fncaudal fnpreanal ridgespelvic fn (female, no claspers)ventsnoutnostrilgill slitsprecaudal tailmouthnasoral groovesymphisial grooveanterior nasal faplower labial furrowupper labial furrow barbelcircumnarial foldcircumnarial grooveincurrent apertureexcurrent apertureanterior nasal fap liItedFAO Species Catalogue for Fishery Purposes No. 7 6labial furrowlabial foldfn insertionfn origininner marginfree rear tipposterior marginanterior marginapexbasespineanterior marginorigin insertioninner marginfree rear tipposterior marginapexbaseupper eye lidsecondary eye lidnictitating lower eye lidsubocular pocketnotchPOSTERIORMEDIAL LATERALANTERIORlower originpreventral marginupper origindorsal marginepaxial webdorsal lobeterminal lobe posterior tipterminal marginsubterminal marginsubterminal notchupper postventral marginposterior notchlower postventral marginventral tipventral lobehypaxial webclasper tiprhipidionpseudosiphoncover rhipidionapopylepelvic fnclasper grooveclasper spurhypopylelateral foldclaspershaftclasperglansFig. 6 EyesFig. 7 Mouth cornerFig. 8 Dorsal nFig. 9 Caudal nFig. 10 Pectoral nFig. 11 Dorsal view of clasper (lamnid shark)Sharks, Batoids and Chimaeras of the North Atlantic 71.2.2 Picture Guide to SkeIetaI TerminoIogy of Sharksinternasal platerostrumrostrumrostral nodenasal apertureorbital notchoptic nerve foramenpreorbital processanterior fontanellecranial roofparietal fossaforamen magnum occipital centrumhyomandibular facetdistal radialsintermediate radialsmetapterygial axismetapterygiumpropterygiummesopterygiummetapterygiumproximal radialsintermediate radialsdistal radialsradialspropterygiumradialsbasalsbasalspterotic hornsphenopterotic ridgeotic capsulecranial roofstapedial fenestrasuborbital shelfcarotid foramenstapedial fenestrabasal plateorbital notchnasal aperturesupraorbital crestsupraorbital crestorbit nasal capsulesuborbital shelfsubethmoid fossa nasal capsulenasal fontanellehyomandibular facetotic capsulepostorbital processa) DORSAL VIEW b) VENTRAL VIEWc) LATERAL VIEWa) APLESODIC b) PLESODICPOSTERIORPOSTERIOR ANTERIORANTERIORmetapterygial axisFig. 12 ChondrocraniumFig. 13 Aplesodic and plesodic pectoral nsFAO Species Catalogue for Fishery Purposes No. 7 8beta cartilagebasipterygiumintermediate segments (2)axial cartilageaxial cartilageapopyleventral marginalventral marginalventral marginalterminal 3 cartilagedorsal marginaldorsal marginaldorsal marginalclasper groovehypopyledorsal terminaldorsal terminaldorsal terminalventral terminalventral terminaltransverse notchcutting edgemesial root loberootneckbladecuspletcuspletcentral foramentransverse groovetransverse ridgesserrationsrootrootcrowncrownpedicelbasemedial ridgemedial cusplateral cusp lateral ridgerootcrowncuspletcuspcuspcrown footcrown footbasal groovebasal ledgedistal root lobeventral terminalend-style of axial cartilageend-style of axial cartilageappendix-stem of axial cartilagefoldederectdorsal terminal 2b) VENTRAL b) DORSALa) LABIAL VIEW b) LINGUAL VIEWDISTALBASALAPICALANTERIORPOSTERIORMESIAL MESIALc) DORSAL (TERMINAL 3 CARTILAGE AND DORSAL TERMINAL 2 REMOVEDMEDIALMEDIALMEDIALLATERALLATERALLATERALFig. 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 denticleSharks, Batoids and Chimaeras of the North Atlantic 91.2.3 Measurements Used for SharksTLFLPCLPD2PD1HDLPG1PSPPOBPP1PP2SVLPAL VCLEYLGS1GS5EYHINGPORSODP1LP1RP1AP1PP1HP1L PRNPPSPASIDSDCSACSPCAPRN POR EYL EYH NG GS1 GS2 GS3 GS4 GS5 GS6 GS7 P1A P1R P1B P1 P1P P1H P1L SOD = PRENARAL LENGTH= PREORAL LENGTH= EYE LENGTH= EYE HEGHT= NTERGLL LENGTH= FRST GLL SLT HEGHT= SECOND GLL SLT HEGHT= THRD GLL SLT HEGHT= FOURTH GLL SLT HEGHT= FFTH GLL SLT HEGHT= SXTH GLL SLT HEGHT= SEVENTH GLL SLT HEGHT= PECTORAL-FN ANTEROR MARGN= PECTORAL-FN RADAL LENGTH= PECTORAL-FN BASE= PECTORAL-FN NNER MARGN= PECTORAL-FN POSTEROR MARGN= PECTORAL-FN HEGHT= PECTORAL-FN LENGTH= SUBOCULAR POCKET DEPTHTL FL PCL PD2 PD1 HDL PG1 PSP POB PP1 = TOTAL LENGTH = FORK LENGTH = PRECAUDAL-FN LENGTH = PRE-SECOND DORSAL-FN LENGTH = PRE-FRST DORSAL-FN LENGTH = HEAD LENGTH = PREBRANCHAL LENGTH = PRESPRACULAR LENGTH = PREORBTAL LENGTH = PREPECTORAL-FN LENGTH PP2 SVL PAL DS DCS PPS PAS ACS PCA VCL = PREPELVC-FN LENGTH= SNOUT-VENT LENGTH= PREANAL-FN LENGTH= NTERDORSAL SPACE= DORSAL CAUDAL-FN SPACE= PECTORAL-FN PELVC-FN SPACE= PELVC-FN ANAL-FN SPACE= ANAL-FN CAUDAL-FN SPACE= PELVC-FN CAUDAL-FN SPACE= VENT CAUDAL-FN LENGTHFig. 17 Main longitudinal measuresFig. 18FAO Species Catalogue for Fishery Purposes No. 7 10= FRST DORSAL-FN MDPONT PECTORAL-FN NSERTON= FRST DORSAL-FN MDPONT PELVC-FN ORGN= PELVC-FN MDPONT FRST DORSAL-FN NSERTON= PELVC-FN MDPONT SECOND DORSAL-FN ORGNPDI PDODPITRHABHTAHCPHHDHDPODAO DAID1LD1HD1PD1AD1IP2BP2HANHANPP2AANAP2PANBP2IANIP2LANLD2LD2BD2AD2H D2PD2ID1BCDMCPUCTLCTRCSTCPVCPLCFLCFWCSW= DORSAL CAUDAL-FN MARGN= PREVENTRAL CAUDAL-FN MARGN= UPPER POSTVENTRAL CAUDAL-FN MARGN= LOWER POSTVENTRAL CAUDAL-FN MARGN= CAUDAL-FN FORK WDTH= CAUDAL-FN FORK LENGTH= SUBTERMNAL CAUDAL-FN MARGN= SUBTERMNAL CAUDAL-FN WDTH= TERMNAL CAUDAL-FN MARGNCDM CPV CPU CPL CFW CFL CST CSW CTR = FRST DORSAL-FN LENGTH= FRST DORSAL-FN ANTEROR MARGN= FRST DORSAL-FN BASE= FRST DORSAL-FN HEGHT= FRST DORSAL-FN NNER MARGN= FRST DORSAL-FN POSTEROR MARGN= SECOND DORSAL-FN LENGTH= SECOND DORSAL-FN ANTEROR MARGN= SECOND DORSAL-FN BASE= SECOND DORSAL-FN HEGHT= SECOND DORSAL-FN NNER MARGN= SECOND DORSAL-FN POSTEROR MARGN= PELVC-FN LENGTH= PELVC-FN ANTEROR MARGN= PELVC-FN BASE= PELVC-FN HEGHT= PELVC-FN NNER MARGN [LENGTH]= PELVC-FN POSTEROR MARGN [LENGTH]= ANAL-FN LENGTH= ANAL-FN ANTEROR MARGN= ANAL-FN BASE= ANAL-FN HEGHT= ANAL-FN NNER MARGND1L D1A D1B D1H D1 D1P D2L D2AD2B D2H D2 D2P P2L P2A P2B P2H P2 P2PANLANAANBANHANHDH TRH ABH TAH CPH DA DAO DPDPOPDPDO= HEAD HEGHT= TRUNK HEGHT= ABDOMEN HEGHT= TAL HEGHT= CAUDAL-FN PEDUNCLE HEGHT= SECOND DORSAL-FN NSERTON ANAL-FN NSERTON= SECOND DORSAL-FN ORGN ANAL-FN ORGNFig. 19 Measurements of caudal nFig. 20 Measurements of dorsal, pelvic and anal nsFig. 21 Other common measurementsSharks, Batoids and Chimaeras of the North Atlantic 11CLOCLINOWANFINWMOLMOWLLAULAGIRHDWTRWABWTAWCPWINOSPLESLCLBCLO CL CLB MOL MOW ULA LLA NOW NW ANF NOSPLESLHDWTRWABWTAWCPW= CLASPER OUTER LENGTH= CLASPER NNER LENGTH= CLASPER BASE WDTH= MOUTH LENGTH= MOUTH WDTH= UPPER LABAL-FURROW LENGTH= LOWER LABAL-FURROW LENGTH= NOSTRL WDTH= NTERNARAL SPACE= ANTEROR NASAL-FLAP LENGTH= NTERORBTAL SPACE= SPRACLE LENGTH= EYE SPRACLE SPACE= HEAD WDTH= TRUNK WDTH= ABDOMEN WDTH= TAL WDTH= CAUDAL-FN PEDUNCLE WDTHFig. 22FAO Species Catalogue for Fishery Purposes No. 7 12

pectoral Iin malar, or cheek thorns of maturemales)spiracle1.2.4 Picture Guide to ExternaI TerminoIogy and Measurements Used for Batoidsalar thorns (or wing thorns of mature malespelvic fn, anterior lobepelvic fn, posterior lobeclasper of males1st dorsal fn2nd dorsal fncaudal fnanuslateral tail foldinner margin oI pelvic fntail spine (sting)dorsal cutaneous foldventral cutaneous foldthorns of median rowlength of snout, preorbitallength of snout, preorallength of tailwidth of discmouthgill slitsnasal aperturestotal 41 rowssnout angle in front of level of spiracleslength of discaxil of pectoral fneyeFig. 23 Upper side of a typical skate (family Rajidae)Fig. 24 Base of tail in stingrays (family Dasyatidae)Fig. 25 Lower side of a typical skateFig. 26 Anterior part of disc of a skateFig. 27Teeth of a stingray (arrows indicate method of counting pavement pattern in batoids)Sharks, Batoids and Chimaeras of the North Atlantic 131.2.5 Picture Guide to ExternaI TerminoIogy and Measurements Used for Chimaeras1st dorsal fnlateral-line canalsfrontal tenaculum(mature males)snoutnostrilmouthsoft gill covergillopeningpectoralfnprepelvic tenaculum (mature males)pelvic fnanal fnclasper (male sex organ)fn spinetotal lengthhead length body length2nd dorsal fncaudal fncaudal flamentFig. 28 Lateral view of a typical ChimaeraFAO Species Catalogue for Fishery Purposes No. 7 14snout elongated, sawlikedisc subrhombic to lozenge-shaped; pelvic fns uni-lobed; tail thin, mostly long and whip-like, often with serrated sting on rootpectoral fns forming a large oval disc; uni-lobed pelvic fns; tail massive, with two large dorsals and a large caudal fn; large electric organ on each side of headexcept Guitarfshes,pectoral fns fused with head and trunk to form mostly subrhombic disc; pelvic fns bilobed; tail rather slender, with two small dorsals and rudimentary caudal fnCARTILAGINOUS FISHESPCTURE KEY OF SHARKS, BATODS AND CHMAERAS ORDERS (not a cladogram)HEXANCHFORMESSQUALFORMESSQUATNFORMESRAJFORMESTORPEDNFORMESMYLOBATFORMESPRSTFORMESCHMAERFORMESCARCHARHNFORMESLAMNFORMESORECTOLOBFORMESone gill slit5 to 7 gill slitsanal fn present5 gill slits, 2 dorsal fns, no fn spines6 or 7 gill slits, 1 dorsal fnmouth behind front of eyesmouth well in front of eyesnictitating eyelids, spiral or scroll intestinal valveno nictitating eyelids, ring intestinal valveno anal fnbody not raylikebody fattened, raylikesnout short, not sawlikemouth terminal, pectoral fns not attached to headmouth ventral, pectoral fns attached to sides of headFig. 29 Higher classication of Sharks, Batoids and Chimaeras (Orders)Sharks, Batoids and Chimaeras of the North Atlantic 151.2.6 GIossary of TechnicaI TermsThefollowingglossaryoftermsusedfortheanatomy andbiologyofshark-likefshesismodifedfromtermsin Compagno(1984,1988,1999)andashortglossaryin Compagno,EbertandSmale(1989).Themainglossary (1.2.4.1)duplicatesthatinvolume2oftherevisedshark catalogue (Compagno, 2001), except that additional terms forhabitatanddistributionusedinthetextarereinstated intheglossary.Termsforphotophorepatterns(including photomarks)thatpertainonlytotheFamilyEtmopteridae andprimarilytothegenusEtmopterusarediscussed under that family.Abdominalridgesorkeels:nsomesharks,paired longitudinal dermal ridges that extend from the bases of the pectoral fns to the pelvic-fn bases.Abyss:Thedeepseabottom,oceanbasinsorabyssal plain descending from 4500 m to about 6000 m.AccessorydorsaImarginaI:ntheclasperskeleton,a fatcartilageontheposteriorendofthedorsalmarginal cartilage that supports the cover rhipidion.AdductormandibuIaemuscIes:Pairedheadmuscles originatingonthelateralfacesofthequadrateprocessof the palatoquadrates and inserting on the lateral surface of the Meckel's cartilages; the primary jaw-closing muscles of sharks.AdeIphophagy:Foetus-eating,amodeoflive-bearing reproduction employing uterine cannibalism; early foetuses deplete their yolk-sacks early and subsist by frst eating their smaller siblings and then eating nutritive eggs produced by themother. Atpresentonlyknownforcertaininthesand tigershark(Carchariastaurus),butsuspectedinafew other lamnoids.Alternate teeth: Small oral teeth with asymmetrical crowns that form two interdigitated rows on the symphysis, with the cuspsofeachrowhookedmesiallytowardstheopposite row.Additionalpairedrowsofalternatesmaybepresent distal to the symphysial rows.Amphitemperate:Referringtoaspeciesthatoccursin temperate water in the northern and southern hemispheres, but is absent from the tropics.AnaIn:Asinglefnontheventralsurfaceofthetail betweenthepelvicfnsandcaudalfnofsomesharks, absentinbatoids,dogfsh,sawsharks,angelsharks,and some chimaeras.AnnuIarringsorannuIi:navertebralcentrumincross section, rings of calcifed cartilage separated by uncalcifed cartilagethatoccupytheintermedialiaonly,orconcentric rings that cross both the intermedialia and basalia.Anterior: Forward, in the longitudinal direction of the snout tip. Also, cranial.Anterior fontanelle: On the elasmobranch neurocranium, anapertureontheanterodorsomedialsurface,usuallyat the rear of the ethmoid region and forming a passage into the internal cranial cavity. t is closed by a tough membrane, variestremendouslyinshape,andmaybepinchedoffby the medially expanded orbits in a few sharks.Anterior margin: n precaudal fns, the margin from the fn origin to its apex.Anterior nasaI ap: A fap on the front edges of the nostrils, thatservestopartiallydividethenostrilintoincurrentand excurrent apertures or openings.Anterior teeth: Enlarged, tall, narrow-rooted oral teeth near the symphysis, often with lingually curved cusps.AnterodorsaIpaIpebraIdepressormuscIe:nthe orectoloboidfamilyParascylliidae,pairedheadmuscles that originate at the insertions of the preorbitalis muscles on theanterolateroventralfaceoftheMeckel'scartilage,and insertontheskinoftheuppereyelidanteriortotheeye. Thesearepossiblyfordepressingtheuppereyelidsand closing the eyes, and are not found in any other sharks.Antorbital cartilages: On the neurocranium of sawsharks and batoids, separate cartilages attached to the sides of the nasal capsules that support the sides or front of the head.Apex: n precaudal fns, the distal tip, which can be acutely pointed to broadly rounded.ApicaI: n oral teeth, towards the tip of the crown or cusp. Can also be used as indicating direction towards the apex or tip of a fn, fn-spine, etc.ApIacentaI viviparity: Live-bearing in which the young do nothaveayolk-sacplacenta.Foundinallgroupsoflive-bearing sharks.ApIesodic n: A pectoral, pelvic, dorsal, or anal fn in which the fn radial cartilages do not extend into the distal fn web andbetweenthesupportingceratotrichiaofthefnweb. Modern sharks always have aplesodic caudal fns, in which the haemal arches of the caudal vertebrae do not support the ventral caudal lobe.ApopyIe:Theanterioropeningoftheclasper,onthe anteromesialsurfaceoftheclasperandclosetothevent. Theapopylereceivesspermfromthecloacaandfuid fromthesiphons,whichentertheclaspergrooveandare discharged through the hypopyle. Apopyle is also used for clasperskeletonsfortheanterioropeningofthetubular shafts formed by enlarged marginal and axial cartilages.AxiaIcartiIage:ntheclasperskeleton,theelongated ventralrodorplate-shapedcartilagethatformsthemain support of the clasper. Also termed appendix-stem.Barbels:Longconicalpaireddermallobesonthesnouts ofsharks,thatmayservetolocateprey.Sawsharkshave barbels on the underside of the snout in front of the nostrils asinsturgeon,butmostbarbeledsharkshavethem associatedwiththenostrils,eitherasanextensionofthe anterior nasal faps or as separate structures medial to the nasal apertures.FAO Species Catalogue for Fishery Purposes No. 7 16Basal: n oral teeth, a proximal direction towards the crown foot and roots.Basalcartilagesorbasals:nprecaudalfnsthelarge cartilages of the fn bases, immediately distal to the pectoral andpelvicfngirdlesorthevertebralcolumn(dorsaland anal fns), on which the radials articulate distally. The paired pectoralfnsoflivingsharksprimitivelyhaveatribasal pectoralfn,withapropterygium,mesopterygium,and metapterygiumasbasals,althoughthesemaybefused; inbatoids,additionalneopterygiaIbasaIsmaybeadded betweenthemesopterygiumandmetapterygiumand thepropterygiumisvariablyexpandedanteriorwith apropterygiaIbasaIandaxis.Thepelvicfnshavea basipterygiumthatsupportsthepelvicradialsand,in males, the claspers. The caudal fn has no basals, but these arefunctionally replaced by expanded neural and haemal arches of the vertebral column.BasaI communicating canaIs: See subnasaI fenestrae.Basal groove: n oral teeth, a deep groove proximal to the basalledgeonthelabialsurfaceofthecrownneckand apical root margin.Basalledge:noralteeth,ashelf-likeprojectiononthe labial surface of the crown foot.BasaIpIate:Thefoorofthecranialcavityofthe neurocranium,aventral,medialplateextendingfromthe ethmoidregionbetweentheorbitsandoticcapsulesand belowthecranialcavitytotheoccipitalcondyles,occipital centrum and foramen magnum.Basalsorbasalia:navertebralcentrum,thediagonal spacesbelowtheattachmentsurfacesofthebasidorsal cartilages,abovethebasiventralcartilages,andbetween thetwohalvesofthedoublecone.Basaliamaybeflled with uncalcifed cartilage, may have diagonal calcifcations penetrating the uncalcifed cartilage, or may have calcifed annuliorsolidcalcifedcartilagethatarecontinuous withcalcifcationoftheintermedialia.Seediagonal caIcications and intermedialia.Base: n precaudal fns, the proximal part of the fn between theoriginandinsertion,extendingdistally,andsupported bythecartilaginousfnskeleton.nthecaudalfn,that thickened longitudinal part of the fn enclosing the vertebral column and between the epaxial and hypaxial lobes or webs ofthefn.noralteeth,theproximalrootandcrownfoot, inappositiontothedistalcusp.ndenticles,theproximal anchoring structures, often with four or more lobes, holding the denticles in the skin.Basidorsalcartilages:Apairofwedge-shapedarched, thin cartilages articulating with the dorsolateral surfaces of avertebralcentrumandformingacontinuousneuralarch with the interdorsal cartilages to protect the spinal cord.Basipterygium: The large elongate longitudinal cartilage at the fn base of the pelvic fn, attached to the posterolateral endsofthepelvicgirdleorpuboischiadicbar.The basipterygiumhaspelvicradialsattachedalongitsdistal edgeandhastheclasperskeletonattachedposteriorlyin males.Basiventral cartilages: A pair of rounded or wedge-shaped cartilagesontheventrolateralsurfacesofavertebral centrumthatformthebasesforattachmentofribsin monospondylousprecaudalvertebrae.ndiplospondylous precaudalandcaudalvertebraethebasiventralsform haemalarchesalongwiththeinterventralcartilagesfor protecting the caudal artery and vein.BathypeIagiczone:Thatpartoftheoceansbeyondthe continental and insular shelves, from about 1000 m to 3000 to 6000 m and above the middle and lower continental rises andtheabyssalplain,thesunlesszone.Someoceanic sharksmaytransittheepipelagic,mesopelagicand bathypelagic zones to the bottom while migrating vertically. Batoid: A ray or fat or winged shark, a neoselachian of the cohort Batoidea, a sawfsh, sharkray, wedgefsh, guitarfsh, thornray,panray,electricray,skate,stingray,butterfyray, eagleray,cownoseray,devilrayorManta.Raysare closelyalliedtothesawsharks(Pristiophoriformes)and angelsharks(Squatiniformes),butdifferfromthemin having the pectoral fns fused to the sides of the head over thegillopenings,whichareventralratherthanlaterallyor ventrolaterally placed. BenthicorDemersal:referringtoorganismsthatare bottom-dwelling.Beta cartilage: n the clasper skeleton, a single, dorsolateral fattened, wedge-shaped or cylindrical cartilage connecting the pelvic basipterygium and axial cartilage and reinforcing theintermediatesegments,possiblyderivedfromapelvic radial.Blade:noralteeth,anarcuate,convex-edgedsectionof the cutting edge of the crown foot, without cusplets.Body ridges: Elongated longitudinal dermal ridges on the sides of the trunk and precaudal tail in certain carpet sharks (Orectolobiformes), in the whale, zebra and some bamboo sharks.Body: Can refer to an entire shark, sometimes restricted to the trunk and precaudal tail.Branchialarches:Thepairedvisceralarchesbehindthe hyoidarchandjustinfrontofthescapuIocoracoidthat support the gills. n elasmobranchs the fve to seven branchial arches primitively consist of a pair of dorsomedial and wedge-shaped cartilages, the pharyngobranchiaIs, closely situated against the roof of the pharynx, a pair of dorsolateral and more cylindricalepibranchiaIsthatareconnecteddorsomedially to the pharyngobranchiaIs, a pair of ventrolateral cylindrical ceratobranchialsthatareconnectedventrolaterallytothe epibranchials,apairofventromedialhypobranchialsthat areconnectedventrolaterallytotheceratobranchials,and unpairedventromedialbasibranchialsthatareconnected ventrolaterallytothehypobranchials.Thehypobranchials andbasibranchialsalongwiththeexpandedventralends oftheceratobranchialsformthebasibranchialskeleton of the foor of the branchial pharynx. The branchial skeleton isvariablymodifedinelasmobranchs,withbasibranchials andsometimeshypobranchialsoftenlost,thelasttwo pharyngobranchialsandthelastepibranchialoftenfused together,andthelastbasibranchialoftenexpandedintoa Sharks, Batoids and Chimaeras of the North Atlantic 17long,broadcopuIawithwhichtheanteriorhypobranchials and posterior ceratobranchials articulate.CaIcied cartiIage: Shark skeletons are formed of hyaline cartilageorgristle,butthisisoftenreinforcedwithlayers of calcifed cartilage, cartilage impregnated with a mineral, hydroxyapatite,similartothatofbonebutorganized differently, in a hard, tile-like pavement of tiny tesserae, or morecompactlyasinthecalcifedstructuresofvertebral centra.CaIcieddoubIecones:nvertebrae,theprimary calcifcationsofthenotochordalsheath,inlateralview resemblingtwohollow,horizontalconeswiththeirapices merged, or an hourglass.CannibaI viviparity: See uterine cannibaIism.Carcharhinoid:Agroundshark,amemberoftheorder Carcharhiniformes,andincludingthecatsharks,false catsharks,fnbackedcatsharks,barbeledhoundsharks, houndsharks,weaselsharks,requiemsharksand hammerheads.Carina: On the crowns of oral teeth, a low blunt mesodistal ridge replacing the cusp and cutting edge, in sharks that eat hard-shelled invertebrate prey.Carotidforamen:Asingleforamenoroneofapairof foraminathatpenetratethebasalplateusuallynearits midlength and allow passage of the internal carotid arteries intothecranialcavity.nsomeadvancedelasmobranchs thecarotidforaminashiftthroughthestapedialforamina and onto the medial wall of the orbit.CartiIaginous shes: Members of the class Chondrichthyes.CaudaI crest: A prominent saw-like row of enlarged pointed denticlesalongthedorsalcaudalmarginandsometimes alongtheventralcaudalmarginofthecaudalfn.Found incertainsharksincludinghexanchoidsandsome carcharhinoids.CaudaI n: The fn on the end of the tail in shark-like fshes, lost in some batoids.CaudaIkeeIs:Adermalkeeloneachsideofthecaudal peduncle that may extend onto the base of the caudal fn, and may, in a few sharks, extend forward as a body keel to the side of the trunk.CaudaI peduncIe: That part of the precaudal tail extending from the insertions of the dorsal and anal fns to the front of the caudal fn.Centralforamen:noralteeth,anutrientforamenonthe midline of the lingual surface of the root, in the transverse groove.Centrum(pIuraI,Centra):Aspool-shaped,partiallyor usuallyfullycalcifedstructurethatformsasasegmental constriction in the notochordal sheath of neoselachians, and which as an articulated string forms the principal structural units of the vertebral column. Centra are primarily formed by the calcifed double cones in the notochordal sheath, which maybetheironlycalcifcation,butadditionalsecondary calcifcationmayoccurinthecentrumbetweentheouter surfacesofthecalcifeddoublecones,includingcalcifed intermedialia, radii, annuli, and diagonal calcifcations.Ceratotrichia:Slendersoftorstiffflamentsofanelastic protein,superfciallyresemblingkeratinorhorn,fromthe Greek keratos, horn, and trichos, hair. Ceratotrichia run in parallel and radial to the fn base and support the fn webs. The prime ingredient of shark-fn soup.Chimaera:AmemberoftheorderChimaeriformes, subclass Holocephali, see also Chimaeroid, HoIocephaIi.Chimaeroid:Achimaera,ratfsh,silvershark,ghost shark,spookfshorelephantfsh,amemberoftheorder Chimaeriformes.Chondrichthyan: Referring to the class Chondrichthyes.Chondrichthyes:TheclassChondrichthyes,fromGreek chondros,cartilage,andichthos,fsh,amajortaxonomic groupofaquatic,gill-breathing,jawed,fnnedvertebrates withprimarilycartilaginousskeletons,1to7external gillopenings,oralteethintransverserowsontheirjaws, andmostlysmall,tooth-likescalesordermaldenticles. Chondrichthyesincludethelivingelasmobranchsand holocephalans and their numerous fossil relatives, and also can be termed shark-like fshes or simply sharks.Chondrocranium: See neurocranium.CircumgIobaI: Occurring around the world.CircumnariaI foId: A raised semicircular, lateral fap of skin around the incurrent aperture of a nostril, in heterodontoids, orectoloboids, and a few batoids, defned by a circumnarial groove.CircumnariaI groove: A shallow groove defning the lateral bases of the circumnarial folds.CircumtropicaI:Occurringaroundthetropicalregionsof the world.CIasper cIaws: n parascylliid orectoloboids, a longitudinal rowoflargeanterolaterallydirectedclaw-likedenticleson the dorsolateral surface of the clasper glans, supported by the terminal ventral.CIasper dactyI: n parascylliid orectoloboids, a large fnger-like process on the medial face of the clasper, supported by the dorsal terminal and having a mesospur, an analogue to the lateral spur or spine of the terminal 3 cartilage of other orectoloboids and other sharks.CIaspergafforhook:ntheexternalclasperglans,a posterior hook-like structure, like a clasper spur but formed from the dorsal terminal cartilage, found in squaloids of the family Squalidae.CIaspergIans:Thedistalanddorsalpartoftheexternal clasperfromthehypopyletoitstip,andincludingvarious movableterminalstructures;also,thesameareaofthe clasper skeleton.FAO Species Catalogue for Fishery Purposes No. 7 18Claspergroove:Thelongitudinalgroovethroughthe clasper,surroundedbytheaxialandmarginalcartilages, and connecting the apopyle and hypopyle.CIasper hooks: n the clasper glans of some carcharhinoid sharks, small claw-like dermal denticles arranged in a row along the ventral surface of the free edge of the exorhipidion.CIaspersacs:Dermalsacswithlongitudinallyribbed walls on the ventral and medial surfaces of the claspers of hexanchoids.CIaspershaft:Thatpartoftheclasperskeletonfromits originonthepelvicfnbasipterygiumtothehypopyle; also,thatpartoftheexternalclasperfromitsbasetothe hypopyle.CIasper spine: n the external clasper, a projection of the terminal3cartilageonthelateralsurfaceoftheclasper glans,whichformsashorttolong,acutelypointed,spine that is covered with shiny hard tissue, poss